The bees of Michigan - Isaacs Lab - Michigan State University

Zootaxa 4352 (1): 001–160 http://www.mapress.com/j/zt/ Copyright © 2017 Magnolia Press

Monograph

ISSN 1175-5326 (print edition)

ZOOTAXA

ISSN 1175-5334 (online edition)

https://doi.org/10.11646/zootaxa.4352.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:7C684128-FFA7-48AA-B395-B9C6BC39353A

ZOOTAXA 4352

The bees of Michigan (Hymenoptera: Apoidea: Anthophila), with notes on distribution, taxonomy, pollination, and natural history JASON GIBBS1,2, JOHN S. ASCHER3, MOLLY G. RIGHTMYER4 & RUFUS ISAACS1 1

Department of Entomology, Michigan State University, East Lansing, MI, USA 48824. Current address: Department of Entomology, University of Manitoba, Winnipeg, MB, Canada R3T 2N2. E-mail: [email protected] 3 Department of Biological Science, National University of Singapore, Singapore 117543. 4 San Diego, CA, USA, 92116.

2

Magnolia Press Auckland, New Zealand Accepted by A. Smith-Pardo: 28 Aug 2017; published: 21 Nov. 2017

JASON GIBBS, JOHN S. ASCHER, MOLLY G. RIGHTMYER & RUFUS ISAACS The bees of Michigan (Hymenoptera: Apoidea: Anthophila), with notes on distribution, taxonomy, pollination, and natural history (Zootaxa 4352) 160 pp.; 30 cm. 21 Nov. 2017 ISBN 978-1-77670-268-8 (paperback) ISBN 978-1-77670-269-5 (Online edition)

FIRST PUBLISHED IN 2017 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/j/zt

© 2017 Magnolia Press All rights reserved. No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing. This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use. ISSN 1175-5326

(Print edition)

ISSN 1175-5334

(Online edition)

2 · Zootaxa 4352 (1) © 2017 Magnolia Press

GIBBS ET AL.

Table of contents Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 Results and discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Bees of Michigan annotated checklist . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 ANDRENIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Andreninae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Andrenini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Genus Andrena Fabricius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Subgenus Andrena Fabricius s. s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 Subgenus Callandrena Cockerell s. l. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 Subgenus Cnemidandrena Hedicke . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Subgenus Conandrena Viereck . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Subgenus Euandrena Hedicke . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 Subgenus Gonandrena Viereck . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 Subgenus Holandrena Pérez . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 Subgenus Iomelissa Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Subgenus Larandrena LaBerge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Subgenus Leucandrena Hedicke . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 Subgenus Melandrena Pérez . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23 Subgenus Micrandrena Ashmead . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24 Subgenus Parandrena Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Subgenus Plastandrena Hedicke . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Subgenus Ptilandrena Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Subgenus Rhacandrena LaBerge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 Subgenus Scaphandrena Lanham . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 Subgenus Scrapteropsis Viereck . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27 Subgenus Simandrena Pérez . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 Subgenus Taeniandrena Hedicke . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 Subgenus Thysandrena Lanham . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 Subgenus Trachandrena Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30 Subgenus Tylandrena LaBerge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 Panurginae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 Calliopsini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 Genus Calliopsis Smith . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 Subgenus Calliopsis Smith s. s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 Subgenus Verbenapis Cockerell & Atkins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 Perditini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 Genus Perdita Smith . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 Subgenus Cockerellia Ashmead . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 Subgenus Perdita Smith s. s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34 Protandrenini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 Genus Protandrena Cockerell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 Genus Pseudopanurgus Cockerell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 APIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 Apinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 Anthophorini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 Genus Anthophora Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 Subgenus Clisodon Patton . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38 Subgenus Melea Sandhouse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 Subgenus Mystacanthophora Brooks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 Genus Habropoda Smith . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 Apini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 Genus Apis Linnaeus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 Bombini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 Genus Bombus Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 Subgenus Bombias Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 Subgenus Cullumanobombus Vogt . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 Subgenus Psithyrus Lepeletier . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 Subgenus Pyrobombus Dalla Torre . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 Subgenus Subterraneobombus Vogt . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 Subgenus Thoracobombus Dalla Torre . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 Eucerini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

THE BEES OF MICHIGAN

Zootaxa 4352 (1) © 2017 Magnolia Press ·

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Genus Eucera Scopoli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 Subgenus Synhalonia Patton . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 Genus Florilegus Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 Subgenus Florilegus Robertson s. l. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 Genus Melissodes Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 Subgenus Apomelissodes LaBerge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 Subgenus Eumelissodes LaBerge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 Subgenus Heliomelissodes LaBerge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47 Subgenus Melissodes Latreille s. s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 Genus Peponapis Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 Subgenus Peponapis Robertson s. l. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 Genus Svastra Holmberg . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 Subgenus Epimelissodes Ashmead . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 Nomadinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 Ammobatoidini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 Genus Holcopasites Ashmead . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 Epeolini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 Genus Epeolus Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 Genus Triepeolus Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 Nomadini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 Genus Nomada Scopoli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 Osirini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65 Genus Epeoloides Giraud . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65 Xylocopinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 Ceratinini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 Genus Ceratina Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 Subgenus Zadontomerus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66 Xylocopini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 Genus Xylocopa Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 Subgenus Xylocopoides Michener . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 COLLETIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 Colletinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 Genus Colletes Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 Hylaeinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 Genus Hylaeus Fabricius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 Subgenus Cephalylaeus Michener . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70 Subgenus Hylaeus Fabricius s. s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71 Subgenus Metziella Michener . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72 Subgenus Paraprosopis Popov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72 Subgenus Prosopis Fabricius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73 Subgenus Spatulariella Popov . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74 HALICTIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74 Halictinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74 Augochlorini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 Genus Augochlora Smith . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 Subgenus Augochlora Smith s. s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 Genus Augochlorella Sandhouse . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 Genus Augochloropsis Cockerell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 Subgenus Paraugochloropsis Schrottky . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 Halictini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76 Genus Agapostemon Guerin-Meneville . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76 Subgenus Agapostemon Guerin-Meneville s. s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76 Genus Halictus Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77 Subgenus Nealictus Pesenko . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77 Subgenus Odontalictus Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77 Subgenus Protohalictus Pesenko . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77 Subgenus Seladonia Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78 Genus Lasioglossum Curtis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78 Subgenus Dialictus Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78 Subgenus Evylaeus Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88 Subgenus Hemihalictus Cockerell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88 Subgenus Lasioglossum Curtis s. s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90 Subgenus Leuchalictus Warncke . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91 Subgenus Sphecodogastra Ashmead . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92 Genus Sphecodes Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93 Nomiinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97


GIBBS ET AL.

Dieunomiini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 Genus Dieunomia Cockerell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 Subgenus Dieunomia Cockerell s. s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 Rophitinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 Genus Dufourea Lepeletier . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 MEGACHILIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98 Megachilinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98 Anthidiini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98 Genus Anthidiellum Cockerell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98 Subgenus Loyolanthidium Urban . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98 Genus Anthidium Fabricius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99 Subgenus Anthidium Fabricius s. s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99 Subgenus Proanthidium Friese . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 Genus Dianthidium Cockerell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 Subgenus Dianthidium Cockerell s. s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100 Genus Stelis Panzer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101 Subgenus Dolichostelis Parker & Bohart . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101 Subgenus Stelis Panzer s. s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101 Genus Trachusa Panzer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102 Subgenus Heteranthidium Cockerell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102 Megachilini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103 Genus Coelioxys Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103 Subgenus Boreocoelioxys Mitchell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103 Subgenus Coelioxys Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104 Subgenus Cyrtocoelioxys Mitchell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104 Subgenus Paracoelioxys Gribodo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105 Subgenus Synocoelioxys Mitchell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105 Subgenus Xerocoelioxys Latreille s. s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105 Genus Megachile Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106 Subgenus Acentron Mitchell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106 Subgenus Callomegachile Michener . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106 Subgenus Chelostomoides Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106 Subgenus Eutricharaea Thomson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107 Subgenus Leptorachis Mitchell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108 Subgenus Litomegachile Mitchell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108 Subgenus Megachile Latreille s. s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109 Subgenus Megachiloides Mitchell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110 Subgenus Sayapis Titus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110 Subgenus Xanthosarus Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111 Osmiini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112 Genus Ashmeadiella Cockerell . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112 Subgenus Ashmeadiella Cockerell s. s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112 Genus Chelostoma Latreille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113 Subgenus Prochelostoma Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113 Genus Heriades Spinola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113 Subgenus Neotrypetes Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113 Genus Hoplitis Klug . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114 Subgenus Alcidamea Cresspm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114 Genus Osmia Panzer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115 Subgenus Cephalosmia Sladen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115 Subgenus Diceratosmia Robertson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115 Subgenus Helicosmia Thomson . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115 Subgenus Melanosmia Schmiedeknecht . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116 Subgenus Osmia Panzer s. s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120 MELITTIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121 Mellitinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121 Genus Macropis Panzer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121 Subgenus Macropis Panzer s. s. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121 Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122 Appendix 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139 Appendix 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141 Supplementary table 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143 Supplementary table 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144



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Abstract The state of Michigan occupies an area between the Great Plains and the northeastern United States, bordering four Great Lakes, with diverse biogeographical regions. Michigan also has the second most diverse agriculture in the country, with many crops that depend on bees for pollination. This unique combination provides a wide range of opportunities for bees to persist, yet there is no current published checklist of these important insects. This study was conducted to provide the first annotated checklist of the bee (Apoidea: Anthophila) fauna of Michigan, summarizing aspects of their taxonomy and behavior and to provide provisional conservation assessment. The list was compiled from a critical review of published literature, museum specimens, and database records, supplemented by new collections. In total, 465 species are included in the checklist, including 38 new records, however evidence for 13 species is poor, several more species require taxonomic revision, and the presence of additional species is expected. The exotic megachilid species Megachile apicalis Spinola, M. pusilla Pérez (=concinna Smith, auct.) and Osmia taurus Smith are reported from Michigan for the first time. New state records of native species include Anthidium tenuiflorae Cockerell and Nomada alpha alpha Cockerell, both previously undocumented from eastern North America, and Nomada sphaerogaster Cockerell, which has rarely been recognized. The taxonomy of some bee species is clarified by the formal publication of 11 new synonymies (some previously reported online or in manuscripts). The following list cites junior synonyms first followed by the valid name: Andrena chippewaensis Mitchell 1960 = A. (Simandrena) wheeleri Graenicher 1904; Osmia hendersoni Cockerell 1907 = O. (Melanosmia) tarsata Provancher 1888; Osmia michiganensis Mitchell 1962 = O. (M.) subarctica Cockerell 1912 (new status, removed from synonymy with O. (M.) tersula Cockerell 1912); Sphecodes persimilis Lovell and Cockerell 1907 = S. davisii Robertson 1897; Sphecodes knetschi Cockerell 1898 = S. dichrous Smith 1853; Sphecodes carolinus Mitchell 1956 = S. coronus Mitchell 1956; Sphecodes stygius Robertson 1893 = S. mandibularis Cresson 1872; Sphecodes prostygius Mitchell 1960 = S. fattigi Mitchell 1956; Stelis vernalis Mitchell 1962 = S. coarctatus Crawford 1916; and Stelis michiganensis Mitchell 1962 = S. foederalis Smith 1854. Poorly known Andrena (Cnemidandrena) are discussed, including A. parnassiae Cockerell, a new state record, A. robervalensis Mitchell, and the extralimital A. runcinatae Cockerell. Of these, only A. robervalensis was considered in the subgeneric revision, but we recognize all three as valid species pending further study. Nomada binotata (Robertson 1903) and N. quadrimaculata (Robertson 1903) are removed from synonymy with N. ovata (Robertson 1903), based on examination of the lectotypes. A new species, Triepeolus eliseae Rightmyer, the eastern representative of the verbesinae species group, is described. A putative undescribed species, Osmia aff. trevoris, is documented, but requires additional study for its status to be fully resolved. A rich bee fauna is documented that includes geographically-restricted species, rare and regionally-declining species, and economically-important species, providing information for ongoing conservation planning and future analysis of trends in bee populations. Key words: check list, faunal list, new species, nomenclature, pollinators, synonymies

Introduction The bee fauna of North America remains incompletely documented, despite widespread interest in native bee conservation (Cariveau & Winfree 2015; Matheson et al. 1996). Assessing changing bee populations remains a challenging task (Williams et al. 2001), particularly given the limited published information on species distributions, incomplete digitization of historical collections, and lack of modern revisionary studies for many bee genera. However, recent analyses of these trends suggest a range of dynamics: declining populations in some taxa, increasing in others, and stable in the majority (Bartomeus et al. 2013a; Colla et al. 2012). Unfortunately, a considerable proportion of regional species cannot be analyzed due to a lack of adequate samples, a problem compounded by identification difficulties resulting from a lack of comprehensive taxonomic revisions. Future examination of bee population trends will depend on improved documentation of regional bee diversity (Berenbaum et al. 2007), including publication of annotated checklists of species for each state. Such efforts provide baseline data on species distribution and can spur further research on bee conservation and diversity. Printed catalogs (Hurd 1979; Moure & Hurd 1987) and monographs on bees of the Eastern United States (Mitchell 1960, 1962) provide state-level distributional data for North American bees, but taxonomic and distributional data provided therein are now outdated. Subsequent publications have clarified the status of many genera including the species-rich and taxonomically challenging Lasioglossum (Gibbs 2010b, 2011; Gibbs et al. 2013; McGinley 1986), Andrena (Bouseman & LaBerge 1979; Donovan 1977; LaBerge 1977, 1980, 1987, 1989; Ribble 1968), and Nomada (Schwarz & Gusenleitner 2004; Droege et al. 2010). A compilation of taxonomic and distributional data for world bees assembled by JSA and published online at http://discoverlife.org (Ascher & Pickering 2017) is updated regularly to reflect a continuous influx of recently published records and reports, with


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many hundreds of new state records mapped online together with specimen records from diverse sources with varying data quality. Contributions of verified state records come from multiple sources but particularly S. Droege (USGS Patuxent Wildlife Research Center (PWRC)) and T. Griswold (USDA Bee Biology and Systematics Laboratory (BBSL)), who each maintain specimen databases, and records verified by JSA and captured in the Arthropod Easy Capture (AEC) database (http://sourceforge.net/p/arthropodeasy Version: 1.34, 2013; see Schuh et al. 2010). Specimen records captured by collaborative databasing efforts have recently become more accessible through data portals such as the USGS-run BISON (Biodiversity Information Serving Our Nation), but the quality of these data is inconsistent. Furthermore, a large proportion of records need to be digitized before they will be accessible for assembling state, county, and site lists of bees. State checklists allow for an inclusive and critical review of all available records that can support future studies of the bee fauna of that state and they can inform regional- and national-scale studies. Several state lists for bees have been published recently in the United States, including Wisconsin (Wolf & Ascher 2009), Pennsylvania (Donovall & VanEngelsdorp 2010), Indiana (Jean 2010), and Colorado (Scott et al. 2011). Other state lists are in development or have been disseminated online (Droege 2016; Pascarella & Hall 2012). Cory Sheffield (Royal Saskatchewan Museum) is leading an effort to document Canadian bees. Additional publications such as Zarrillo et al. (2016) for Connecticut supplement the historical baseline (Mitchell, 1960, 1962, Hurd, 1979) by detailing new and interesting state records. Some reports of new state records, however, are less reliable, e.g., most records of native bees reported for the first time from New Hampshire by Tucker & Rehan (2016) require additional scrutiny. Michigan is nested within the Great Lakes Region and has biogeographic features of both the central eastern regions and northern tier states. For example, the Upper Peninsula (UP) and Isle Royale have affinities to more northern flora and fauna than does the Lower Peninsula (LP) (Husband et al. 1980). A floral tension zone crosses the LP, separating it into northern (NLP) and southern (SLP) zones that are distinct in climate, soils, and plant communities. Some bumble bees have northern or southern range limits which correspond well with this floral tension zone (Husband et al. 1980). Coastal habitats on the western edge of the LP include sand dunes adjacent to Lake Michigan, which influences the climate that is suitable for regionally unique agriculture as well as flora and fauna not seen in other parts of the state (Reznicek 1994). This region also represents the northern range limits of some bee species that are typically found further south (Tuell et al. 2009). Historical collections of Michigan bees come from a variety of sources, one of the most important being that of Robert R. Dreisbach (Dow Chemical, Midland, Michigan), who collected extensively across the state and beyond, often with his wife Kathryn. Reported bee species richness is high for Midland County where he lived and worked (see map, Fig. 1). Dreisbach’s personal collection of over 250,000 insect specimens was willed to Michigan State University (Fischer 1965). Unfortunately, Dreisbach did not always maintain careful collection data (Cantrall 1968), and many of his labels only list the county, date, and collector without more specific locality data (Fig. 2A, B). It is rumored among local entomologists that Dreisbach intentionally sought to obtain multiple county records in a single collection bout by focusing his efforts around the boundaries between counties. The Dreisbachs’ Michigan collections have been used in the description of many insect species including numerous bees (Mitchell 1960, 1962), resulting in 23 Michigan holotypes of bees, nine of which are currently valid, and many additional paratypes of bees from Michigan. Details are provided in the species accounts below for the holotypes, which reside at the National Museum of Natural History (NMNH) in Washington, D.C. With the exception of type series, Mitchell (1960, 1962) did not provide more location-specific information nor did he indicate the source of state records. Mitchell (1960, 1962) recorded 398 species present in Michigan, based on his concepts of species, which is nearly twice the species totals he reported for adjacent states such as Indiana and Ohio. A long-term study by Francis C. Evans (University of Michigan, Ann Arbor) summarized extensive bee collections initiated by Urless N. Lanham in the 1950’s, including floral association records from the Edwin S. George Reserve in Livingston County, Michigan (Evans 1986). Much of this important material, which includes several unique bee species records for the state, is housed at the University of Michigan, Museum of Zoology (UMMZ). The W.K. Kellogg Biological Station (KBS) of Michigan State University (MSU) located next to Gull Lake in Kalamazoo County has also been visited repeatedly by MSU collectors (Matthews 1965). Specimens from KBS, including many uncommon species for the state, frequently bear labels recording the collection locality as “Gull Lake Biol. Sta.” (Fig. 2C) and were collected by Roland L. Fischer who taught Introductory Entomology during the summer at KBS during his tenure at MSU from 1953 to 1992 (Stehr & Nielsen 1994). The names of his



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students including the melittologist George C. Eickwort and the lepidopterist Ronald W. Hodges are frequently seen on labels from KBS and other sites. Ronald Hodges also collected important bee records from Isle Royale in Keweenaw County, the most northerly region of the state. Specimens from a dissertation on strawberry pollination are housed at MSU and contribute records to a number of counties, particularly the otherwise under sampled Manistee County (Connor 1973). During the 1980’s and early 1990’s, environmental impact studies related to testing of an extremely low frequency communications system by the U.S. Navy were conducted in the UP (Scott 1994, 1996; Strickler et al. 1996; Strickler & Scriber 1994). This resulted in numerous specimens from Dickinson and Iron Counties made by Roland Fischer and graduate students Karen Strickler and Virginia Scott. Records from Ingham and Washtenaw Counties are numerous due to the presence of MSU and the University of Michigan in these two counties, respectively. The high relative abundance of records from Cheboygan County, many of which are more than a century old, are due in part to the University of Michigan Biological Station located at Pellston. The station sits on the south end of Douglas Lake and many labels from the station list this as the locality. Hundreds of these records are housed in institutions outside of Michigan, including the American Museum of Natural History and Ohio State University. Other frequently collected localities include Rose Lake Wildlife Area in Clinton and Shiawassee Counties in central Michigan near MSU and the Yankee Springs Recreation Area in Barry County in the southwest. Most of the remaining counties in the state have been less frequently sampled for bees historically.

FIGURE 1. Map of Michigan showing counties colored by their known bee species richness.

Collections that are more recent come from a variety of sources. Mark O’Brien (UMMZ) has recently published studies on the distribution and nesting biology of megachilid bees in Michigan (O’Brien 2007; O’Brien et al. 2013; O’Brien & Craves 2008). Unpublished bee surveys by Ann Fraser and students from Kalamazoo


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College in Kalamazoo and Barry Counties provide some valuable records (Arnosky 2009), including species not previously recorded for the state. Recent studies focused on documenting bees, primarily crop pollinators, in highbush blueberry fields in western Michigan have provided numerous records from Van Buren, Ottawa, and Allegan Counties (Gibbs et al. 2016; Tuell et al. 2008, 2009; Tuell & Isaacs 2009). Tuell et al. (2009) reported 166 species present in blueberry fields, including some new state records such as Andrena confederata Viereck and A. neonana Viereck. More recent sampling as part of the Integrated Crop Pollination Project (http://ProjectICP.org) in apple, blueberry, and cherry farms from the southwest region and north to Oceana and Leelanau Counties has produced several thousand new specimen records of crop pollinators (Gibbs et al. 2016, 2017). A survey of potato fields provided additional records from Montcalm County (Buchanan et al. 2017). The Great Lakes Bioenergy Research Center has funded pollinator research in Michigan biofuel crops (corn, switchgrass and prairie) including recent sampling of a number of additional counties across the Lower Peninsula (Bennett et al. 2014; Bennett & Isaacs 2014; Gardiner et al. 2010). Several recent and ongoing projects at MSU that extend earlier research on blueberries, old fields (Carson et al. 2016), prairie restoration, biofuels, and opportunistic collecting by the lead author, primarily near MSU, have contributed additional material for this study. These collection efforts have produced several-thousand bee specimens that have been summarized in this checklist.

FIGURE 2. Historical collection labels for type material of bees from Michigan at the National Museum of Natural History, Smithsonian Instituion A) Robert Dreisbach label. B) Robert and Kathryn Dreisbach label. C) Roland Fischer label. Photographs provided with the permission of the National Museum of Natural History, Smithsonian Institution, 10th and Constitution Ave. N.W., Washington, DC 20560-0193. (http://www.nmnh.si.edu/).

Methods We compiled a list of bee and county records from a variety of sources including the scientific literature, museum specimens, online image databases (http://bugguide.net—only images readily identifiable by JSA), and our own collections. A preliminary list of species and county records was compiled from published studies, primarily taxonomic revisions (Baker 1975; Bouseman & LaBerge 1979; Broemeling & Moalif 1988; Brooks 1983; Droege et al. 2010; Gibbs 2010b, 2011; Gonzalez & Griswold 2013; LaBerge 1961, 1969, 1973, 1977, 1980, 1987, 1989, 1956a; b; LaBerge & Bouseman 1970; LaBerge & Ribble 1972, 1975, McGinley 1986, 2003, Milliron 1971, 1973a; b, Mitchell 1960, 1962; Onuferko 2017; Ordway 1966a; Ribble 1967; Rightmyer 2008; Rightmyer et al. 2010; Stephen 1954; Timberlake 1954, 1958, 1960, 1968, 1969, 1973, 1976). Other studies based on identifications by trained taxonomists were also consulted (Arduser 1986; Bartomeus et al. 2013a; Buchanan et al. 2017; Dreisbach 1945; Evans 1986; Gardiner et al. 2010; Gibbs et al. 2016, 2017; Miller et al. 2002; Milliron 1939;



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O’Brien 2007; O’Brien & Craves 2008; Scott 1996; Tuell et al. 2008, 2009; Tuell & Isaacs 2009). Records from older studies were updated based on current taxonomic understanding. We also examined specimens from regional collections, particularly the A. J. Cook Arthropod Research Collection at Michigan State University, East Lansing, Michigan (MSUC), the University of Michigan Museum of Zoology, Ann Arbor, Michigan (UMMZ), and the insect collection at Kalamazoo College, Kalamazoo, Michigan (KCIC). Some type material at the Illinois Natural History Survey (INHS), NMNH, and California Academy of Sciences (CAS) were examined to clarify species concepts. MSUC collections currently housed in research labs are indicated using the abbreviation of the principal investigator (RI: Rufus Isaacs, DL: Doug Landis). These include the majority of specimens from recent collections in blueberry fields, biofuel plantings, and at pollinator habitat restoration sites. A number of interesting records from unpublished studies were available from research conducted at the Southwest Michigan Research and Extension Center (SWMREC) in Berrien County and Clarksville Research Center (CRC) in Ionia County. Many bumble bee records from Michigan were digitized as part of another study (Cameron et al. 2011) and 2,131 MSUC records were shared for six species by S. Cameron. Specimen data in the Digital Bee Collections Network captured using Arthropod Easy Capture (AEC) Software (2013; Schuh et al. 2010) resulted in several hundred additional records that are deposited in the American Museum of Natural History (AMNH, 562 records), Cornell University Insect Collection (CUIC, 96), Bohart Museum of Entomology, University of California, Davis (56), University of Massachusetts, Amherst (21), Connecticut Agricultural Experimental Station (15); and other collections combined for a total of 21 records: Field Museum of Natural History, Newark Museum, New York State Museum, Rutgers University, University of Connecticut at Storrs, University of New Hamsphire and the NMNH. Records from the AMNH and other digitized collections also enhance our knowledge of broader distribution patterns of the regional bee fauna. Additional database records were gathered from the Illinois Natural History Survey (INHS, 497, of which 389 are Bombus), Snow Entomological Museum Collection, University of Kansas (SEMC, 518), C. A. Triplehorn Insect Collection, Ohio State University (OSUC, 528), USGS Patuxent Wildlife Research Center, Bee Monitory and Inventory Laboratory, located in Beltsville, Maryland (PWRC, 1,385; most ultimately to be deposited at the NMNH), University of Colorado (UCMC, 4) and USDA-ARS Bee Biology and Systematics Laboratory (BBSL, 5,466, of which 5,365 were Bombus see Cameron et al. 2011). These records were filtered to remove species that have undergone recent taxonomic changes making their identity uncertain. The type database maintained at the NMNH was also used for information related to type specimens originating from Michigan. These have been photographed and images are available online at http://collections.nmnh.si.edu/search/ento/. Hadel Go (AMNH) shared images of holotype specimens to help clarify taxonomic limits of some taxa. The Barcode of Life Data Systems (Ratnasingham & Hebert 2007) were checked for additional Michigan records of bees included in molecular diagnostic studies, most vouchers of which are housed at the Packer Collection at York University (PCYU). Recent collections by JG are deposited at MSUC: RI or the J. B. Wallis/R. E. Roughley Museum of Entomology (JBWM). A call for additional bee records was made to the Michigan Entomological Society (Gibbs 2015), resulting in a small number of additional county records, including Houghton County specimens donated to MSUC by Dana Richter (Michigan Technological University) and several county records from the private collection of Mark VanderWerp (MVWC). Additional records for Houghton County came as part of pilot study using citizen scientists, specimens are deposited at JBWM. Mike Arduser provided additional records and notes on Michigan bees from his personal collection. Thomas Wood (MSU) provided several valuable records from 2017 collections, which are currently deposited in his personal collection (TJWC). Species names from earlier studies were updated to reflect current taxonomic and nomenclatural understanding when necessary. As examples, Heriades is treated as feminine and both Melissodes and Coelioxys are considered masculine (ICZN 1999: art. 30.1.2 and 30.1.4.4), although in the literature Heriades has often been treated as masculine and Melissodes and Coelioxys nearly always as feminine (LaBerge 1961; Rocha Filho & Packer 2016). However, prevailing usage is not considered relevant with respect to gender agreement in this situation by the International Code of Zoological Nomenclature, so we are compelled to adopt the correct gender as verified by Commissioner D. Yanega (personal communication). The spelling of changeable specific epithets for species in those genera therefore differs from most previous works. Subgeneric categories for Bombus and Lasioglossum follow recent studies of these genera (Gibbs 2016; Gibbs et al. 2013; Williams et al. 2008). Family-group taxonomy follows Michener (2007) and the World Bee Checklist (http://itis.gov/) from 2007, with some subsequent updates following Scott et al. (2011) and Ascher & Pickering (2017). Our list is organized


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alphabetically by family, subfamily, tribe, genus, subgenus, and species. The species description follows the format of Rightmyer (2008). Specimens were loaned from NMNH, INHS, UCMC, Oregon State Arthropod Collection (OSAC), Rob Jean personal collection (RJPC), and CAS. Since many specimens from Michigan have no locality information below the level of county, including those obtained by its most prolific historical collector, we have organized our list by county, compiling a list of all bee species occurrences for each of Michigan’s eighty-three counties. We included records based on our critical review of the literature as well as specimens examined from historical and recent collections. In all cases, we attempted to verify each record by examining specimens ourselves. Particular attention was paid to species of uncertain taxonomic status, known to have been misidentified by historical or recent specialists, or with a known range that made occurrence in Michigan implausible. We document separately from our main list those species likely to occur in, but not yet recorded from, Michigan based on known ranges which make occurrence in Michigan probable, e.g., occurrence in both southern Ontario and any combination of Midwestern states to the south or west of Michigan (Appendix 1). We also exclude from the main list those species that have been recorded in the literature but for which the current evidence does not support their occurrence (Appendix 2). We provide additional notes and details for a subset of bee species that are new state records, species of conservation concern, valid species and their junior synonyms where the name-bearing type is from Michigan (Supplementary table 1), and where recent taxonomic or nomenclatural confusion necessitates some additional discussion or clarification. Label information is provided in a standard format for consistency. Host plant records are updated to current taxonomic understanding based on the USDA plants list (plants.usda.gov). We provide some citations related to biological information for the species in our checklist when available. This is not intended as an exhaustive review of the biological literature, but is intended to provide readers with a starting point for more detailed study and stronger basis for analyses using biological data.

Results and discussion Analysis of a wide range of sources and specimens resulted in 464 valid bee species and 1 putative undescribed species occurring in Michigan (Supplementary Table 2). This total is substantially more than the 398 species recorded in Mitchell’s (1960, 1962) distributional tables, but the totals are not directly comparable due to the differing concepts of species in his era. The substantial increase in species richness in our final list highlights both the contributions of subsequent authors and the importance of reviewing unpublished historical collections and the ongoing discovery of state-level bee records even in a relatively well-sampled region. Despite this, most counties in Michigan remain undersampled (Fig. 1), and the continuing discovery of new state records and of species and morphospecies new to science suggests that future additions can be expected, including native bees and additional exotic introductions (Gibbs et al. 2017; Martins et al. 2017). However, evidence for 13 species in our species list is poor, and some taxa such as various Nomada in the ruficornis species group require taxonomic revision that may change the total species documented here. Museum collections housed substantial undocumented diversity. New state records, including Anthidium tenuiflorae Cockerell, Ashmeadiella bucconis (Say), Coelioxys immaculatus (Cockerell), Megachile pusilla Pérez (as concinna), M. dakotensis Mitchell, and M. petulans Cresson, were found already identified at MSUC. Reexamination of specimens confirmed these unpublished records. New records for other species such as Lasioglossum fedorense (Crawford) were found among undetermined material (UMMZ in this case). Several new species records for Michigan, including Anthidiellum notatum (Latreille), Lasioglossum oenotherae (Stevens), Megachile apicalis Spinola, M. mucida Cresson, Nomada sphaerogaster Cockerell, and Pseudopanurgus compositarum (Robertson), were first discovered based on collections made by JG. These were mostly found in Ingham County, near MSU, which is the best-documented county in the state highlighting that new species may be detected with greater sampling even in areas with previous extensive collection effort. The exotic species Osmia taurus Smith is newly recorded for the state, reflecting general expansion of this species (e.g., to New York City, JSA unpublished). Until recently, O. taurus was best known from Mid-Atlantic states, where it was probably introduced when the similar exotic species O. cornifrons (Radoszkowski) was intentionally established by USDA scientists (Batra 1979). The Michigan specimen was collected with a long series of the closely related O. cornifrons, highlighting the risk of missing closely related species if material is not examined by expert taxonomists. THE BEES OF MICHIGAN


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Our total of verified bee species is comparable to those for other states in the northern Midwest and Northeastern US (Fig. 3), such as Massachusetts (377; Goldstein & Ascher, 2016); Wisconsin (391 Wolf & Ascher 2009; updated by Scott et al. 2011), Pennsylvania (371; Donovall & vanEngelsdorp 2010), Connecticut (349; Zarrillo et al. 2016), and New York (447; Ascher et al. 2014). Indiana, directly to the south of western Michigan has a reported 420 bee species (Jean 2010). No published estimate exists for Ohio, to the south of eastern Michigan other than the records available from Mitchell (1960, 1962). JSA’s online compilation (Ascher & Pickering 2017) records only 295 species, suggesting that the state remains undersampled. The province of Ontario to the north and east of Michigan likely has a similar number of bee species; certainly exceeding 400 according to Packer et al. (2007) and C. Sheffield (pers. comm.), although Ascher & Pickering (2017) record only 364 as verified. The relatively high number for Michigan is evidently a reflection of the existing collection effort in a range of habitat types and locations, combined with its unique biogeographical context. The contribution of species that are characteristic of the Great Plains, the northeeastern United States, and the boreal region is unique among states. The number of species recorded from each of Michigans’ counties varies considerably (Fig. 1). Ingham and Van Buren Counties both have 234 species. The former is the home of MSU and, until recently, the lead author, whereas the latter has been thoroughly sampled in recent years by RI’s lab group. In contrast, Luce County in the UP has the lowest total of a mere 30 bee species currently known. Such low species counts certainly reflect undersampling rather than true bee species richness. By contrast, five counties in the UP that received considerable sampling effort in the early 1990’s have more than 3 times that amount, including Dickinson County with 130 species. Counties with any level of systematic sampling, even if for a short time, typically have over 100 verified species records, but 57 of Michigan’s 83 counties do not reach that amount. Ingham County is largely unremarkable in terms of geographical location or landscape diversity so there is every reason to suspect that other counties will also have more than 200 species present. Livingston County, immediately adjacent to Ingham, also has over 200 species, but only 140 species in common with Ingham County, suggesting that true species richness for both counties, and by extension those surrounding them, could easily exceed 300 species. Such a high number is plausible as it is comparable to that found in Robertson’s (1929) exhaustive surveys of Carlinville, Illinois, and the 308 species reported from Tompkins County in central New York State (JSA, unpublished; most records are supported by specimens in the AEC database).

FIGURE 3. Relative bee richness of states in the Northeast and Midwestern United States with recently published records of species numbers. See text for sources.


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The composition of the bee community is fairly typical for the region (Giles & Ascher 2006; Goldstein & Ascher 2016). Species richness of families are ordered from largest to smallest: Halictidae (123), Apidae (117), Andrenidae (102), Megachilidae (87), Colletidae (35) and Melittidae (one). A number of bee genera that are known from neighbouring states are noticeably absent from Michigan, including Melitoma, Ptilothrix (Apidae: Emphorini), Melecta, Xeromelecta (Apidae: Melectini), and Melitta (Melittidae). Our analysis included documentation of the traits of the bee community, which can be useful for future ecological studies. Approximately 60% of species are solitary (including communal species and incipiently social members of the Xylocopinae), more than 22% are cleptoparasites or social parasites and nearly 18% are known or inferred to be eusocial. Due to taxonomic difficulties, predominantly in the genus Nomada, among which undescribed species exist even in well-studied areas of eastern North America (see Goldstein & Ascher, 2016) and a lack of direct observations of social behavior in most halictid bees, these percentages are likely to change. Lasioglossum display a wide diversity of social behaviors, but the social status of most species has not been studied in sufficient detail, if at all, so must be inferred based on phylogenetic patterns available for exemplar taxa (Danforth et al. 2003; Gibbs et al. 2012b). A number of reversals to solitary behavior have been reported in Lasioglossum (Danforth et al. 2003), so inferences of social behavior may prove to be incorrect in a minority of cases as additional biological data and improved phylogenetic reconstruction become available. The majority of nest-building species (70%) dig underground burrows. The remainder nest in pre-existing tunnels (12%), stems (8%), wood (2%), abandoned snail shells (Osmia conjuncta Cresson), or construct hives in larger cavities (5%) or on exterior surfaces or rocks or plant stems (three species). Nesting substrates of Osmia are sufficiently diverse (Cane et al. 2007) that the nesting habits for eight Michigan species are uncertain because of a lack of published information. Cavity and stem nesting species often bring foreign materials into the nest to construct cells. These include cut leaf pieces or masticated leaf pulp (43 species), plant resins (seven species), plant hairs (four species) or mud (four species). Some species use multiple materials, such as Osmia bucephala Cresson, which mixes wood fibers with leaf pulp in cell construction (Krombein 1967). There has been widespread concern about the status and trends of wild bee populations (Cane & Tepedino 2001; Goulson et al. 2008), and reports of global declines despite awareness that for many regions of the world there is insufficient taxonomic and distributional information to accurately and comprehensively assess bee faunas. Berenbaum et al. (2007) and Gonzalez et al. (2013) highlighted the need for more thorough monitoring across the United States, but recognized that such studies are hampered by insufficient taxonomic and distributional resources for bees. Bartomeus et al. (2013a) examined historical changes of bees in the Northeast including a large proportion of species found in Michigan. Fifty-one of these were found to be significantly declining in relative abundance, 78 were stable, and 56 were increasing in relative abundance. Although statistical evaluation was possible for the majority of regional bee species, 166 species included in that study were insufficiently represented in historical databases at the time to assess the temporal trends in populations. The poorly represented species likely include many that are genuinely rare and deserving of highest priority for conservation action, but others were underrepresented due to difficulties in identification. The quality and completeness of our checklist and the ability to assess conservation status comprehensively would be much improved if recent revisions were available for certain taxa, especially Nomada and Sphecodes, genera for which existing species delimitations and identification criteria are unreliable. For many Nomada and some species in other genera, identifications of material other than the primary type are speculative, rendering records for these species in Michigan and other states open to doubt. Based on our examination of some primary types, we elevate two species of Nomada to species status, however in the absence of a revision the range and status of these species remains uncertain. A new species of Triepeolus is described below. In addition, it was necessary to clarify the taxonomy of Andrena (Cnemidandrena) to address the status of nominal species not treated in the revision of this subgenus (Donovan 1977) but subsequently placed there. Even recent revisions may be incomplete, leaving room for additional study. For example, despite integrative taxonomic revisions of Lasioglossum (Dialictus) for Canada (adjoining Michigan) and for Eastern North America (Gibbs 2010b, 2011) challenges remain for fully resolving the L. (Dialictus) in this region, due to uncertainty about some sex associations and about species limits in the L. viridatum species-group. This bee species checklist and the information on host associations and nesting provided here can support the development of bee conservation strategies. Efforts to expand the area of pollinator plantings that provide foraging and nesting resources for bees have focused on diverse seed mixes assuming they will support the greatest diversity of bees (Williams et al. 2015), and there has been significant investment to establish pollinator habitat in Michigan



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by federal, state, and private organizations. There is also great interest in developing sustainable practices for crop pollination, and this may include conservation of wild bees to provide pollination services (Garibaldi et al. 2014). Although honey bees are the primary strategy used for crop pollination in large, commercial fields (Gibbs et al. 2016), wild bees can make important contributions (Isaacs & Kirk 2010). Highlighting their role in food production may also enhance support for broader adoption of practices to conserve bees in other landscapes. Revitalized integrative taxonomic research of bees, digitization of historical records, mobilization of citizen scientist data including images, and improved understanding of natural history are needed to understand bee diversity and promote bee conservation. Lacking advances in each of these four areas of study, it will be challenging to understand what species are present, when and how they have been impacted by human activity or how to predict their occurrence or habitat requirements across the landscape. Previous studies taking advantage of extensive databasing efforts in the northeastern US have documented a diversity of historical trends including declines, stability and increasing abundance of exotics (Bartomeus et al. 2013a), changes in phenology (Bartomeus et al. 2011, 2013b) and range extensions (Zarrillo et al. 2016). Although our data are not fully digitized, it is evident that some changes to the Michigan bee fauna have occurred over time. In spite of extensive sampling, no contemporary collections of Bombus affinis Cresson have been found, supporting its well-documented decline (Cameron et al. 2011). In addition, the large distinctive bee Dieunomia heteropoda heteropoda has only recently been found in Michigan at the limits of its range (Gibbs et al. 2014), suggesting a possible range extension. Most recently, a collection of Osmia cornifrons (Radoszkowski) was made in mid-February 2017, which is evidence of a dramatically earlier spring emergence for bees in the region. Additional sampling efforts are also needed to fill gaps in our basic understanding of bee distribution. Given the paucity of records for many counties in Michigan, despite a history of active collectors, renewed effort to sample areas of the state and identify the collections are needed. The same is true for other areas of the country and globally. With this study, we provide an updated and extensive resource for those interested in bees collected in Michigan or the Midwest region, whether for understanding which species are present or their ecology, conservation, or contributions to pollination of flowering wild and crop plants. Identification of 465 species of bees from Michigan is more than the richness reported from nearby states, suggesting a high proportion of the species pool has been reported here. However, with new state records becoming known as recently as 2017, continued sampling will undoubtedly result in additional discoveries and improved clarity of bee species distribution. We hope this report will stimulate additional interest in wild bees and their ecology across Michigan landscapes.

Bees of Michigan annotated checklist ANDRENIDAE Andreninae Andrenini Genus Andrena Fabricius Taxonomy: Mitchell 1960; LaBerge 1967, 1969, 1973, 1977, 1980, 1985, 1986, 1989; Ribble 1967, 1968, 1974; LaBerge & Bouseman 1970; LaBerge & Ribble 1972, 1975; Bouseman & LaBerge 1978. Behavior: Andrena are predominately solitary, although some species will form communal nests (Osgood 1989). Species are often narrowly polylectic or mesolectic, but many species are oligolectic, specializing on a particular floral host taxon (Linsley 1958; Robertson 1926).

Subgenus Andrena Fabricius s. s. Revision: LaBerge 1980.


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Andrena (Andrena) carolina Viereck 1909 County records: Alger, Allegan, Bay, Clare, Crawford, Dickinson, Jackson, Lake, Macomb, Mason, Midland, Montmorency, Muskegon, Oscoda, Ottawa, Presque Isle, Roscommon, Schoolcraft, Van Buren. Notes. Specialist on Vaccinium (Ericaceae). Andrena carolina is abundant in blueberry fields and an important native pollinator of the crop.

Andrena (Andrena) clarkella (Kirby 1802) County records: Clare, Crawford, Dickinson, Lake, Marquette, Ottawa, Van Buren. Notes. Holarctic. Specialist on Salix (Salicaceae).

Andrena (Andrena) frigida Smith 1853 County records: Allegan, Antrim, Clare, Clinton, Emmet, Genesee, Grand Traverse, Ingham, Isabella, Luce, Marquette, Midland, Missaukee, Newaygo, Oakland, Saginaw, Van Buren, Wayne. Notes. Specialist on Salix (Salicaceae). Nests in California were briefly described by MacSwain (1945, as rhodotricha Linsley).

Andrena (Andrena) mandibularis Robertson 1892 County records: Allegan, Berrien, Livingston, Oakland, Van Buren, Washtenaw, Wayne. Notes. Polylectic.

Andrena (Andrena) milwaukeensis Graenicher 1903 County records: Alger, Berrien, Houghton, Ingham, Leelanau, Mackinac, Marquette, Oceana, Ottawa.

Andrena (Andrena) rufosignata Cockerell 1902 County records: Antrim, Arenac, Benzie, Chippewa, Dickinson, Emmet, Grand Traverse, Isabella, Kalkaska, Keweenaw, Lake, Leelanau, Mackinac, Manistee, Montmorency, Oceana, Schoolcraft, Van Buren.

Andrena (Andrena) thaspii Graenicher 1903 County records: Alger, Antrim, Baraga, Benzie, Cheboygan, Chippewa, Dickinson, Emmet, Gratiot, Ingham, Iron, Lenawee, Macomb, Manistee, Marquette, Midland, Ontonagon, Van Buren, Washtenaw.

Andrena (Andrena) tridens Robertson 1902 County records: Ingham, Van Buren, Washtenaw. Notes. Polylectic, but may prefer Ribes (Grossulariaceae) (LaBerge 1980), making A. tridens a potentially beneficial wild pollinator of currants.



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Subgenus Callandrena Cockerell s. l. Revision: LaBerge (1967) Note. Callandrena is apparently polyphyletic with none of the Michigan species belonging to A. (Callandrena s. s.) (Larkin et al. 2006).

Andrena (Callandrena s. l.) aliciae Robertson 1891 (aliciae group) County records: Berrien, Ionia, Kalamazoo, Livingston, Oakland, Van Buren. Notes. Asteraceae specialist, with preference for Helianthus (Asteraceae) (LaBerge 1967).

Andrena (Callandrena s. l.) asteris Robertson 1891 (simplex group) County records: Bay, Crawford, Dickinson, Gladwin, Grand Traverse, Gratiot, Huron, Ingham, Isabella, Kalamazoo, Kent, Mackinac, Manistee, Mecosta, Midland, Missaukee, Montcalm, Montmorency, Otsego, Roscommon, Saginaw, Van Buren, Washtenaw, Wayne, Wexford. Notes. Specialist on Symphyotrichum and Solidago (Asteraceae) (LaBerge 1967).

Andrena (Callandrena s. l.) gardineri Cockerell 1906 (gardineri group) (New state record) (Fig. 4) County records: Van Buren. Notes. Andrena gardineri is a specialist on Packera (Asteraceae: Senecioneae) (Larkin et al. 2008). A series of A. gardineri was collected from Packera aurea (L.) Á. Löve & D. Löve (originally identified as Senecio jacobaea L.), found growing at the edge of a ditch adjacent to a commercial highbush blueberry farm. Subsequently, a single specimen collected in 2008 was identified from KCIC. In 2016, collections were also made from Packera aurea in two additional sites in Van Buren County, and the bee was found almost immediately in each location, suggesting it might be quite common where the host plant is found. Material examined. Kalamazoo Co.: Sand Creek Preserve, 10 May 2008, A.M. Fraser (1 ♀ KCIC); Van Buren Co.: Almena, 4.5 km NNE, 20 May 2014, Packera aurea, A. Adamczyk (2 ♀, 1 ♂ MSUC: RI); Almena, 4.5 km NNE, 21 May 2014, P. aurea, J. Gibbs (1 ♀ JBWM); Almena, 4.5 km NNE, 23 May 2014, P. aurea, J. Gibbs (2 ♀ JBWM); Almena, 4.5 km NNE, 18 May 2016, P. aurea, K. Odanaka (1 ♂ MSUC: RI); Almena, 3 km NW, 19 May 2016, P. aurea, J. Gibbs (1 ♀ JBWM); Gobles, 3 km NW, 19 May 2016, P. aurea, J. Gibbs (4♀ JBWM).

FIGURE 4. Andrena (Callandrena s.l.) gardineri Cockerell female on Packera. Photographed by JG in Van Buren County, Michigan.


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Andrena (Callandrena s. l.) helianthi Robertson 1891 (helianthi group) County records: Dickinson, Ingham, Ionia, Jackson, Kalamazoo, Kent, Lenawee, Shiawassee, Van Buren, Washtenaw. Notes. Specialist on Helianthus (Asteraceae) (LaBerge 1967).

Andrena (Callandrena s. l.) krigiana Robertson 1901 (krigiana group) County records: Livingston, Midland. Notes. Specialist on Krigia (Asteraceae) (LaBerge 1967).

Andrena (Callandrena s. l.) placata Mitchell 1960 (simplex group) County records: Allegan, Bay, Cass, Cheboygan, Clinton, Dickinson, Ingham, Ionia, Jackson, Kalamazoo, Kalkaska, Leelanau, Livingston, Midland, Monroe, Oakland, Oceana, Ogemaw, Ottawa, Van Buren, Washtenaw. Notes. Specialist on Solidago and Symphyotrichum (Asteraceae) (LaBerge 1967).

Andrena (Callandrena s. l.) rudbeckiae Robertson 1891 (melliventris group) County records: Allegan, Berrien, Cass, Ingham, Ionia, Kalamazoo, Livingston, Ottawa, Saginaw, Van Buren. Notes. Specialist on Rudbeckia and Ratibida (Asteraceae) (LaBerge 1967). Nesting biology and foraging described from Texas by Neff & Simpson (1997).

Andrena (Callandrena s. l.) simplex Smith 1853 (simplex group) County records: Allegan, Bay, Huron, Ingham, Isabella, Mecosta, Midland, Muskegon, Ogemaw, Otsego, Roscommon, Shiawassee, Washtenaw. Notes. Specialist on Solidago and Symphyotrichum (Asteraceae) (LaBerge 1967).

Subgenus Cnemidandrena Hedicke Revision: Donovan (1977).

Andrena (Cnemidandrena) canadensis Dalla Torre 1896 (nubecula group) County records: Alger, Bay, Benzie, Cass, Cheboygan, Crawford, Dickinson, Emmet, Gladwin, Gratiot, Ingham, Ionia, Iron, Kalkaska, Keweenaw, Lake, Lapeer, Livingston, Mackinac, Manistee, Marquette, Mecosta, Midland, Montcalm, Osceola, Oscoda, Roscommon, Saginaw, Schoolcraft, Van Buren, Wexford. Notes. Specialist on Solidago and Symphyotrichum (Asteraceae).

Andrena (Cnemidandrena) chromotricha Cockerell 1899 (chromotricha group) County records: Midland, Saginaw.



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Andrena (Cnemidandrena) hirticincta Provancher 1888 (hirticincta group) County records: Alger, Allegan, Bay, Benzie, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Crawford, Delta, Dickinson, Emmet, Gladwin, Gogebic, Gratiot, Huron, Ingham, Iron, Isabella, Jackson, Kalamazoo, Keweenaw, Leelanau, Livingston, Luce, Manistee, Marquette, Menominee, Midland, Missaukee, Montcalm, Montmorency, Oakland, Otsego, Ottawa, Saginaw, Schoolcraft, Van Buren, Washtenaw, Wayne, Wexford. Notes. Specialist on Solidago and Symphyotrichum (Asteraceae).

Andrena (Cnemidandrena) nubecula Smith 1853 (nubecula group) County records: Calhoun, Cass, Clinton, Dickinson, Ingham, Jackson, Kalamazoo, Lapeer, Lenawee, Livingston, Marquette, Midland, Newaygo, Van Buren, Washtenaw. Notes. Specialist on Solidago (Asteraceae).

Andrena (Cnemidandrena) parnassiae Cockerell 1902 (scutellinitens group) (New state record) (Fig. 5A) County records: Jackson. Notes. Andrena parnassiae has been overlooked and is therefore of particular taxonomic and conservation interest. The type female (NMNH) was collected by S. Graenicher in Milwaukee, Wisconsin on 14 September (perhaps 1901). Mitchell (1960) also recorded it from Michigan and Vermont. Andrena parnassiae was included in subgenus Leucandrena (Hurd 1979; Lanham 1949; Mitchell 1960) until LaBerge (1986) removed it to Cnemidandrena, so it was not included in Donovan’s (1977) revision of the latter subgenus. The record below was collected as part of a prairie-fen pollinator study (Fiedler et al. 2012) and subsequently identified by JG. The specimen was originally identified as A. runcinatae Cockerell, another poorly known species, by M. Arduser (see below; Gusenleitner et al. 2005). Andrena parnassiae can be distinguished from other members of the ‘scutellinitens’ group: clypeus with dark setae; labral process much broader than long, without median emargination; head distinctly wider than long (length/width = 0.75), following measurements of Donovan (1977); and mesoscutellum dull due to microsculpture. Material examined. Jackson Co.: Liberty-Grand River Fen, N42°05ʹ02ʹʹ, W84°27ʹ38ʹʹ, 31 Aug. 2009, A.K. Fiedler (1♀ MSUC).

Andrena (Cnemidandrena) peckhami Cockerell 1902 (chromotricha group) (New state record) (Fig. 5B) County records: Iron. Notes. The type female (NMNH) of A. peckhami was described in the same publication as A. parnassiae, and was also collected in Milwaukee, Wisconsin, but on 24 July (likely 1901). This species resembles A. parnassiae but can be recognized by the smoother more sparsely punctate medial area of the clypeus and convergent eyes above. It should be noted that the convergent eyes of A. peckhami and A. chromotricha (above) is subtle and requires careful examination. A single female was compared to a Minnesota specimen determined by Mitchell (MSUC) and images of the type (NMNH). Material examined. Iron Co.: T43N R35W Sec. 1, 19 Jul. 1983, R.L. Fischer (1 ♀ MSUC). Additional Material examined. ILLINOIS: Dupage Co.: 17 Sep. 2010 (National University of Sinapore); MINNESOTA: Lake of the Woods Co.: Baudette, 25 mi S. E., 11 Jul. 1952, R.L. Fischer (1 ♀ MSUC); Baudette, 9 Jul. 1952, R.L. Fischer (1 ♂ MSUC, W.E. LaBerge det.); VERMONT: Caledonia Co.: Burke Mt. St. Park, 20 Aug. 1956, Solidago, R.L. Fischer (1 ♀ MSUC—new state record).


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Andrena (Cnemidandrena) robervalensis Mitchell 1960 (scutellinitens group) (Fig. 5C) = Andrena (?Cnemidandrena) robervalensis Mitchell 1960: 175.

County records: Gladwin, Isabella. Notes. Andrena robervalensis was tentatively treated as a junior synonym of A. runcinatae Cockerell 1906 by Gusenleitner et al. (2005), based on JSA’s examination of a paratype of A. runcinatae at the AMNH. However, Cockerell (1906), in his original description of A. runcinatae, describes the clypeus as “shining, with sparse strong punctures, the anterior middle smooth, but no median ridge”. The holotype female of A. runcinatae, from Florissant, Colorado in Teller County “a little over 8000 feet” collected by S. A. Rohwer on 22 July 1906 and deposited at the California Academy of Sciences, matches this description and also has a highly polished mesoscutellum, similar to Donovan’s (1977) description of A. specularia Donovan. Donovan (1977) did not treat A. runcinatae in his revision of the subgenus Cnemidandrena. Hurd (1979) lists A. runcinatae in the subgenus Simandrena. The type of A. runcinatae was compared directly to a paratype of A. robervalensis (MSUC) and images of the holotype female (NMNH). Andrena robervalensis is noticeably duller than A. runcinatae on the clypeus (Fig. 5C, D) and mesoscutellum. A more thorough reexamination of Donovan’s entire ‘scutellinitens’ group in North America is required, i.e., the western species A. scutellinitens Viereck and A. specularia, the previously associated eastern species A. robervalensis, and and the newly associated A. runcinatae and A. parnassiae. Material examined. Gladwin Co.: T19N R2E Sec. 2, 4 Sep. 1959, R.L. Fischer (1 ♀ MSUC); Isabella Co.: (no locality), 9 Sep. 1950, R.R. Dreisbach (1 ♀ paratype MSUC).

Subgenus Conandrena Viereck Revision: LaBerge 1986.

Andrena (Conandrena) bradleyi Viereck 1907 County records: Allegan, Clare, Grand Traverse, Kent, Livingston, Mackinac, Midland, Missaukee, Montcalm, Montmorency, Newaygo, Oakland, Schoolcraft. Notes. Specialist on Vaccinium (Ericaceae) and a wild pollinator of highbush blueberry (LaBerge 1985; Gibbs et al. 2017).

Subgenus Euandrena Hedicke Revisions: LaBerge & Ribble (1975), LaBerge (1977).

Andrena (Euandrena) algida Smith 1853 County records: Allegan, Clare, Gratiot, Iosco, Isabella, Lake, Midland, Otsego, Ottawa, Van Buren. Notes. Polylectic, but may prefer Salix (Salicaceae).

Andrena (Euandrena) geranii Robertson 1891 County records: Allegan, Berrien, Clinton, Dickinson, Grand Traverse, Ingham, Isabella, Lapeer, Leelanau, Mackinac, Monroe, Osceola, Saginaw, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Specialist on Hydrophyllum (Hydrophyllaceae). THE BEES OF MICHIGAN


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FIGURE 5. Faces of female Andrena (Cnemidandrena) species. A) Andrena parnassiae Cockerell from Michigan. B) Andrena peckhami from Michigan. C) Andrena robervalensis paratype female from Michigan. D) Andrena runcinatae holotype female (CAS).

Andrena (Euandrena) nigrihirta (Ashmead 1890) = Andrena (Thysandrena) crenata Mitchell, 1960: 221 (Michigan holotype). Holotype. ♂ USA: Michigan: Arenac Co.: 30 May 1951, R.R. Dreisbach (NCSU). Synonymy by Laberge & Ribble (1975).

County records: Alcona, Allegan, Arenac, Baraga, Cheboygan, Dickinson, Eaton, Emmet, Ingham, Iron, Isabella, Jackson, Keweenaw, Mackinac, Midland, Newaygo, Ontonagon, Ottawa, Presque Isle, Schoolcraft. Notes. Apparently polylectic (LaBerge & Ribble 1975). Mitchell (1960) lists the crenata holotype in the Dreisbach collection, the name-bearing types of which were transferred to the NMNH. The holotype is currently listed in the NCSU type database.

Andrena (Euandrena) polemonii Robertson 1891 County records: None. Notes. This oligolectic species is known from neighbouring states. It is included based on Mitchell’s (1960) authority and records from the Chicago area within 100 km west of the southwest corner of Michigan (LaBerge


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1977; Pearson 1933). It visits Polemonium reptens L., which is restricted to the southern corners of Michigan, but also Geranium maculatum L. and Ranunculus hispidus Michx., which are more widespread in the LP (Robertson 1929; Voss & Reznicek 2012).

Subgenus Gonandrena Viereck Revision: LaBerge & Ribble (1972).

Andrena (Gonandrena) fragilis Smith 1853 County records: Alpena, Arenac, Barry, Bay, Berrien, Calhoun, Cheboygan, Chippewa, Clare, Clinton, Gratiot, Huron, Ingham, Isabella, Kent, Lake, Lapeer, Lenawee, Livingston, Macomb, Manistee, Midland, Missaukee, Montmorency, Oceana, Osceola, Presque Isle, Saginaw, Shiawassee, St. Joseph, Van Buren, Washtenaw. Notes. Specialist on Cornus (Cornaceae) (LaBerge & Ribble 1972).

Andrena (Gonandrena) integra Smith 1853 County records: Allegan, Arenac, Bay, Calhoun, Clinton, Dickinson, Eaton, Gratiot, Hillsdale, Ingham, Ionia, Isabella, Jackson, Lapeer, Lenawee, Livingston, Midland, Saginaw, Sanilac, St. Clair, St. Joseph, Van Buren. Notes. A specialist on Cornus (Cornaceae) (LaBerge & Ribble 1972), although A. integra and A. persimulata Viereck, below, were not included in a recent review of specialist bees of the Northeastern United States (Fowler 2016).

Andrena (Gonandrena) persimulata Viereck 1917 County records: Antrim, Berrien, Calhoun, Cheboygan, Chippewa, Clare, Delta, Dickinson, Livingston, Mackinac, Mason, Midland, Oakland, Osceola, Oscoda, St. Joseph, Tuscola, Van Buren. Notes. Specialist on Cornus (Cornaceae) (LaBerge & Ribble 1972).

Andrena (Gonandrena) platyparia Robertson 1895 = Andrena (Gonandrena) monroensis Mitchell, 1960: 233 (Michigan holotype; Fig. 6A). Holotype. ♀ USA: Michigan: Monroe Co.: 23 Jun. 1948, R.R. Dreisbach (NMNH: 75182). Synonymy by LaBerge & Ribble (1972).

County records: Allegan, Antrim, Calhoun, Clinton, Grand Traverse, Ingham, Kalamazoo, Kent, Lapeer, Lenawee, Livingston, Macomb, Mason, Midland, Monroe, Oakland, Osceola, St. Joseph, Van Buren. Notes. A relatively common specialist on Cornus (Cornaceae) (LaBerge & Ribble 1972).

Subgenus Holandrena Pérez Revision: LaBerge (1986).

Andrena (Holandrena) cressonii cressonii Robertson 1891 County records: Allegan, Alpena, Barry, Berrien, Calhoun, Clare, Clinton, Eaton, Ingham, Ionia, Iosco, Isabella,



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Jackson, Kalamazoo, Kalkaska, Livingston, Macomb, Manistee, Mecosta, Midland, Missaukee, Newaygo, Oakland, Oceana, Ottawa, Saginaw, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Polylectic. Nesting in Illinois described by Miliczky (1988).

Subgenus Iomelissa Robertson Revision: LaBerge (1986). Monotypic.

Andrena (Iomelissa) violae Robertson 1891 County records: Barry, Ingham, Jackson, Kalamazoo, St. Joseph, Van Buren, Washtenaw. Notes. Specialist on Viola (Violaceae). Brief description of nest observed in New Jersey by Smith (1901).

Subgenus Larandrena LaBerge Revision: Ribble 1967.

Andrena (Larandrena) miserabilis Cresson 1872 County records: Allegan, Antrim, Arenac, Barry, Bay, Bezie, Berrien, Branch, Calhoun, Clare, Clinton, Dickinson, Eaton, Genesee, Gladwin, Grand Traverse, Gratiot, Huron, Ingham, Ionia, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Lapeer, Leelanau, Livingston, Mackinac, Macomb, Manistee, Mason, Mecosta, Midland, Monroe, Montcalm, Montmorency, Newaygo, Oakland, Oceana, Ontonagon, Osceola, Oscoda, Otsego, Ottawa, Roscommon, Saginaw, Sanilac, Schoolcraft, Shiawassee, St. Clair, St. Joseph, Van Buren, Washtenaw, Wayne, Wexford. Notes. Nests have been described from Florida (Norden & Scarborough 1979) and Kansas (Michener & Rettenmeyer 1956; as bipunctata). One of the most abundant wild pollinators of apple and cherry in the state (Gibbs et al. 2017) and known to visit commercial plums and pears (Michener & Rettenmeyer 1956).

Subgenus Leucandrena Hedicke Revision: LaBerge (1987).

Andrena (Leucandrena) barbilabris (Kirby 1802) County records: Alger, Allegan, Alpena, Antrim, Baraga, Barry, Bay, Benzie, Berrien, Calhoun, Clare, Crawford, Dickinson, Grand Traverse, Ingham, Ionia, Kalamazoo, Kalkaska, Kent, Keweenaw, Lapeer, Leelanau, Mackinac, Manistee, Marquette, Midland, Oakland, Oceana, Osceola, Ottawa, Van Buren, Washtenaw, Wayne, Wexford. Notes. Holarctic. Abundant wild pollinator of Michigan tart cherry and apple orchards (Gibbs et al. 2017).

Andrena (Leucandrena) erythronii Robertson 1891 County records: Alger, Allegan, Arenac, Benzie, Chippewa, Clare, Dickinson, Houghton, Ingham, Kalkaska, Leelanau, Luce, Mackinac, Manistee, Oakland, Oceana, Osceola, Van Buren. Notes. Specialist on Erythronium (Liliaceae). Nesting in Kansas described by Michener & Rettenmeyer (1956).


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Subgenus Melandrena Pérez Revision: Bouseman & LaBerge (1979).

Andrena (Melandrena) barbara Bouseman & LaBerge 1979 County records: Livingston.

Andrena (Melandrena) carlini Cockerell 1901 County records: Alger, Allegan, Antrim, Baraga, Barry, Benzie, Berrien, Calhoun, Chippewa, Clinton, Dickinson, Grand Traverse, Ingham, Ionia, Isabella, Huron, Jackson, Kalamazoo, Kent, Lapeer, Leelanau, Livingston, Mackinac, Midland, Monroe, Montcalm, Newaygo, Oakland, Oceana, Osceola, Ottawa, Sanilac, Schoolcraft, Shiawassee, Van Buren, Washtenaw, Wayne. Notes. Nests from New Brunswick and Nova Scotia described by Schrader & LaBerge (1978) and Atwood (1933), respectively. Pollen provisions included apple, blueberry, cherry and strawberry (Schrader & LaBerge 1978). This vernal species is among the most abundant wild bees on apple, highbush blueberry and tart cherry in Michigan (Gibbs et al. 2016, 2017).

Andrena (Melandrena) commoda Smith 1879 County records: Allegan, Antrim, Bay, Berrien, Branch, Calhoun, Cass, Cheboygan, Crawford, Gladwin, Gratiot, Ingham, Isabella, Kalamazoo, Kent, Keweenaw, Lenawee, Livingston, Macomb, Manistee, Mecosta, Midland, Monroe, Montcalm, Oakland, Osceola, Ottawa, Saginaw, Shiawassee, Tuscola, Van Buren, Washtenaw, Wexford.

Andrena (Melandrena) confederata Viereck 1917 County records: Allegan. Notes. A single male specimen of this primarily southern species has been recorded from Michigan (Tuell et al. 2009), but it could not be relocated at MSUC.

Andrena (Melandrena) dunningi Cockerell 1898 County records: Benzie, Berrien, Genesee, Grand Traverse, Houghton, Ingham, Kalamazoo, Kent, Leelanau, Livingston, Oakland, Oceana, Osceola, Ottawa, Shiawassee, Van Buren, Washtenaw, Wayne. Notes. Nests were described by Johnson (1981, 1984) along with preferred pollen sources, which included cherry, and apple or pear (Johnson 1984).

Andrena (Melandrena) hilaris Smith 1853 County records: Allegan, Livingston, Midland, St. Joseph, Van Buren. Notes. Andrena hilaris was first recorded from Michigan by Mitchell (1960). No specimens from Michigan or neighboring states were examined by Bouseman & LaBerge (1978) and they refer to it as a relatively rare, southeastern species. Four male Dreisbach specimens identified as A. hilaris were found at MSUC, but upon examination by JG were found to be A. nivalis. Andrena hilaris does occur in Michigan, based on three male specimens collected from a single site in Allegan County (Tuell et al. 2009).



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Andrena (Melandrena) nivalis Smith 1853 County records: Alger, Alpena, Antrim, Baraga, Cheboygan, Chippewa, Clare, Delta, Dickinson, Emmet, Gladwin, Gogebic, Houghton, Ingham, Iron, Jackson, Kalamazoo, Kalkaska, Keweenaw, Leelanau, Livingston, Mackinac, Macomb, Manistee, Marquette, Missaukee, Montmorency, Muskegon, Newaygo, Oceana, Ontonagon, Ottawa, Roscommon, Schoolcraft. Notes. Nesting in Oregon described by Miliczky et al. (1990), where it was a pollinator of pears.

Andrena (Melandrena) pruni Robertson 1891 County records: Allegan, Berrien, Kalamazoo, Midland, St. Joseph, Van Buren. Notes. Underrepresented in historical collections from the eastern United States relative to its current prevalence (JSA, pers. obs.).

Andrena (Melandrena) regularis Malloch 1917 County records: Alger, Baraga, Cheboygan, Clare, Ingham, Iosco, Kalamazoo, Kalkaska, Mackinac, Missaukee, Osceola, Roscommon, Schoolcraft, Wexford. Notes. Nests from New Brunswick described by Schrader & LaBerge (1978), where pollen provisions included apple, blueberry, cherry and strawberry, but not found in recent surveys of Michigan apple, blueberry and cherry.

Andrena (Melandrena) vicina Smith 1853 County records: Alger, Allegan, Alpena, Antrim, Arenac, Barry, Benzie, Berrien, Branch, Calhoun, Cheboygan, Chippewa, Clare, Clinton, Crawford, Delta, Dickinson, Eaton, Gladwin, Grand Traverse, Gratiot, Houghton, Huron, Ingham, Ionia, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Lapeer, Leelanau, Lenaweee, Livingston, Mackinac, Manistee, Marquette, Mason, Mecosta, Midland, Missaukee, Monroe, Montcalm, Montmorency, Muskegon, Newaygo, Oakland, Oceana, Ogemaw, Osceola, Oscoda, Otsego, Ottawa, Roscommon, Saginaw, Schoolcraft, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. An important native pollinator of spring crops in Michigan including apple, tart cherry and highbush blueberry (Gibbs et al. 2016, 2017). Nests from Maine and Washington were described by Miliczky & Osgood (1995).

Subgenus Micrandrena Ashmead Revision: Ribble (1968).

Andrena (Micrandrena) illinoiensis Robertson 1891 (illinoiensis group) County records: Arenac, Clinton, Eaton, Gladwin, Ingham, Iosco, Isabella, Jackson, Kalkaska, Mecosta, Midland, Missaukee, Montcalm, Newaygo, Oakland, Osceola, Wexford. Notes. Strongly prefers Salix (Salicaceae) (Ribble 1968).


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Andrena (Micrandrena) melanochroa Cockerell 1898 (piperi group) County records: Allegan, Clinton, Crawford, Dickinson, Ingham, Kalkaska, Livingston, Marquette, Ontonagon, Otsego, Roscommon, Van Buren, Wexford. Notes. Polylectic, but prefers rosaceous plants in the genera Fragaria (strawberry) and Potentilla (Ribble 1968). This potential pollinator of strawberries can be collected in abundance from their flowers (Knerer & Atwood 1964a).

Andrena (Micrandrena) neonana Viereck 1917 (piperi group) County records: Allegan, Van Buren. Notes. First recorded from Michigan by Tuell et al. (2009) based on two female specimens. This is a notable range extension since Ribble (1986) recorded it only as far north as southern Illinois and Ohio. Subsequently, Jean (2010) recorded it from southern Indiana.

Andrena (Micrandrena) nigrae Robertson 1905 (illinoiensis group) County records: Allegan, Arenac, Ottawa. Notes. Strongly prefers Salix (Salicaceae) (Ribble 1968).

Andrena (Micrandrena) personata Robertson 1897 (piperi group) County records: Branch, Ingham, Shiawassee. Notes. Polylectic with some preference for Rosaceae, including Crataegus and Rubus (Ribble 1968). Nesting in Illinois described by Miliczky (1988).

Andrena (Micrandrena) salictaria Robertson 1905 (illinoiensis group) County records: Allegan, Antrim, Arenac, Cheboygan, Clinton, Gladwin, Grand Traverse, Ingham, Iosco, Isabella, Jackson, Kalamazoo, Kalkaska, Lake, Lapeer, Livingston, Midland, Missaukee, Montcalm, Newaygo, Oakland, Osceola, Ottawa, Saginaw, Washtenaw, Wexford. Notes. Specialist on Salix (Salicaceae) (Ribble 1968).

Andrena (Micrandrena) ziziae Robertson 1891 (piperi group) County records: Hillsdale, Ingham, Jackson, Midland, Monroe, Van Buren, Wayne. Notes. Specialist on Zizia (Apiaceae) and other members of the family (Ribble 1968).

Subgenus Parandrena Robertson Revision: LaBerge & Ribble (1972).

Andrena (Parandrena) andrenoides (Cresson 1878) County records: Arenac, Clinton, Huron, Ingham, Isabella, Mecosta, Midland, Newaygo, Oakland, Van Buren, Washtenaw. Notes. Specialist on Salix (Salicaceae). THE BEES OF MICHIGAN


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Andrena (Parandrena) wellesleyana Robertson 1897 County records: Allegan, Antrim, Calhoun, Clare, Clinton, Crawford, Gladwin, Grand Traverse, Ingham, Ionia, Isabella, Kalkaska, Kent, Lake, Marquette, Midland, Montcalm, Newaygo, Oakland, Osceola, Ottawa, Shiawassee, Wayne, Wexford. Notes. Visits Salix and Prunus (LaBerge & Ribble 1972).

Subgenus Plastandrena Hedicke Revision: LaBerge (1969).

Andrena (Plastandrena) crataegi Robertson 1893 (crataegi group) County records: Alcona, Alger, Allegan, Alpena, Antrim, Arenac, Baraga, Barry, Bay, Benzie, Berrien, Calhoun, Charlevoix, Cheboygan, Clare, Clinton, Crawford, Dickinson, Eaton, Emmet, Genesee, Gladwin, Grand Traverse, Gratiot, Houghton, Huron, Ingham, Ionia, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lake, Lapeer, Leelanau, Lenawee, Livingston, Mackinac, Macomb, Manistee, Marquette, Mason, Mecosta, Midland, Missaukee, Monroe, Montcalm, Montmorency, Newaygo, Oakland, Oceana, Ogemaw, Osceola, Oscoda, Otsego, Ottawa, Presque Isle, Roscommon, Saginaw, Sanilac, Schoolcraft, Shiawassee, St. Clair, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Communal nesting in Maine described by Osgood (1989). Andrena crataegi is one of the most abundant wild bees in apple and cherry orchards in Michigan (Gibbs et al. 2017), but the relatively late flight season of females, better matching the bloom of hawthorne, may reduce its value in some areas (Gardner & Ascher 2006).

Subgenus Ptilandrena Robertson Revision: LaBerge (1987).

Andrena (Ptilandrena) distans Provancher 1888 County records: Berrien, Gladwin, Ingham, Jackson, Kalamazoo, Kent, Lake, Livingston, Midland, Oakland, Saginaw, Shiawassee, St. Joseph, Van Buren, Washtenaw. Notes. Specialist on Geranium (Geraniaceae).

Andrena (Ptilandrena) erigeniae Robertson 1891 = Andrena (Leucandrena) bifurcata Mitchell, 1960: 211 (Michigan holotype; Fig. 6B). Holotype. ♂ USA: Michigan: Washtenaw Co.: Ann Arbor, 3 Apr. 1949, U.N. Lanham (NMNH: 75171). Synonymy by LaBerge (1987).

County records: Alger, Allegan, Barry, Berrien, Calhoun, Clare, Gratiot, Ingham, Jackson, Kalamazoo, Kent, Lake, Lenawee, Livingston, Midland, Montcalm, Montmorency, Ottawa, Saginaw, St. Joseph, Van Buren, Washtenaw, Wayne. Notes. A specialist of Claytonia, commonly seen in undisturbed woodlands. Nesting biology in Illinois described by Davis & LaBerge (1975).


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Subgenus Rhacandrena LaBerge Revision: LaBerge (1977).

Andrena (Rhacandrena) brevipalpis Cockerell 1930 County records: Alcona, Antrim, Arenac, Benzie, Berrien, Calhoun, Cheboygan, Eaton, Emmet, Genesee, Gladwin, Grand Traverse, Ingham, Iosco, Kalamazoo, Lapeer, Leelanau, Lenawee, Livingston, Manistee, Mason, Midland, Missaukee, Montmorency, Muskegon, Newaygo, Oakland, Oceana, Osceola, Otsego, Presque Isle, Roscommon, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Shows a strong preference for Rhus (sumac; Anacardiaceae) (LaBerge 1977).

Andrena (Rhacandrena) robertsonii Dalla Torre 1896 = Andrena (Gonandrena) dreisbachi Mitchell, 1960: 230 (Michigan holotype; Fig. 6C). Holotype. ♀ USA: Michigan: Roscommon Co.: 5 Jun. 1955, R.R. Dreisbach (NMNH: 75177). Synonymy by LaBerge (1977).

County records: Cheboygan, Ionia, Isabella, Lapeer, Lenawee, Livingston, Macomb, Missaukee, Newaygo, Osceola, Ottawa, Roscommon, Shiawassee, Van Buren, Washtenaw. Notes. Andrena robertsonii is challenging to distinguish from its apparent sister species A. brevipalpis Cockerell (LaBerge 1977), which also occurs in Michigan. Mitchell identified specimens of A. robtersonii at MSUC were found to be be A. brevipalpis upon reexamination by JG. It is known Mitchell (1960) misapplied the name leading him to redescribe true robertsonii as dreisbachi (LaBerge 1977).

Subgenus Scaphandrena Lanham Revision: Ribble (1974).

Andrena (Scaphandrena) arabis Robertson 1897 (scurra group) County records: Allegan, Ingham, Kalamazoo, Montcalm, Ottawa, Van Buren, Washtenaw. Notes. Specialist on Brassicaceae.

Subgenus Scrapteropsis Viereck Revision: LaBerge (1971).

Andrena (Scrapteropsis) alleghaniensis Viereck 1907 (alleghaniensis group) County records: Allegan, Antrim, Berrien, Branch, Clare, Grand Traverse, Ingham, Jackson, Kalamazoo, Leelanau, Livingston, Macomb, Manistee, Muskegon, Oakland, Ottawa, Saginaw, Schoolcraft, St. Joseph, Van Buren, Wexford. Notes. Nesting biology in New York described by Batra (1990a).

Andrena (Scrapteropsis) imitatrix Cresson 1872 (imitatrix group) County records: Antrim, Bay, Benzie, Berrien, Branch, Calhoun, Clare, Clinton, Crawford, Eaton, Gladwin, THE BEES OF MICHIGAN


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Grand Traverse, Ingham, Iosco, Jackson, Kalamazoo, Leelanau, Livingston, Manistee, Mecosta, Midland, Muskegon, Oceana, Osceola, Oscoda, Otsego, Ottawa, Saginaw, Shiawassee, Van Buren, Washtenaw, Wayne, Wexford. Notes. A relatively abundant wild bee in apple and tart cherry orchards in Michigan (Gibbs et al. 2017). Rau (1922) describes a nest in Missouri very briefly.

Andrena (Scrapteropsis) morrisonella Viereck 1917 (imitatrix group) County records: Allegan, Clinton, Genesee, Ionia, Isabella, Kalamazoo, Leelanau, Lenawee, Midland, Oceana, Osceola, Ottawa, Schoolcraft, Van Buren. Notes. This relatively late-flying species remains a challenge to distinguish from A. imitatrix.

Subgenus Simandrena Pérez Revision: LaBerge (1989).

Andrena (Simandrena) nasonii Robertson 1895 County records: Allegan, Barry, Berrien, Branch, Calhoun, Cass, Clare, Clinton, Eaton, Gladwin, Grand Traverse, Huron, Ingham, Jackson, Kalamazoo, Kent, Lake, Lapeer, Leelanau, Lenawee, Livingston, Macomb, Manistee, Midland, Monroe, Oceana, Osceola, Otsego, Ottawa, Saginaw, St. Joseph, Van Buren, Washtenaw. Notes. Andrena nasonii is a relatively abundant wild bee in apple and tart cherry orchards in Michigan (Gibbs et al. 2017).

Andrena (Simandrena) wheeleri Graenicher 1904 = Andrena (?Leucandrena) chippewaensis Mitchell, 1960: 212 (New Synonymy; Michigan holotype; Fig. 6D). Holotype. ♀ USA: Michigan: Chippewa Co.: 3 Jun. 1957, R. & K. Dreisbach (NMNH: 75173).

County records: Antrim, Baraga, Cheboygan, Chippewa, Dickinson, Gladwin, Keweenaw, Leelanau, Mackinac, Marquette, Wayne. Notes. Andrena chippewaensis was described based on two female specimens. The paratype, deposited at North Carolina State University and collected from New Brunswick, Canada was examined by JSA and determined to be A. wheeleri. Examination of images of the NMNH holotype confirms the synonymy. The basis for separation of A. chippewaensis from A. wheeleri in Mitchell’s (1960) key was the wide gena of the former (see couplet 104), but the holotype of A. chippewaensis does not seem to match Mitchell’s own description in this respect, having a relatively narrow gena. As a result, it should come to A. wheeleri in Mitchell’s key instead. A male specimen of A. wheeleri (MSUC) identified by Mitchell has identical collection information to the holotype of A. chippewaensis. LaBerge (1989) suggests it may be an important pollinator of blueberry, although it has not been collected in Michigan’s major highbush blueberry growing region in the southwest. It is also regularly collected from Rubus. Material examined. Antrim Co.: Glacial Hills, Eckhart, 12 Jun. 2016, Rubus (3 ♀ MSUC: RI, JBWM); Torch Lake, southeast, 12 Jun. 2016, Rubus (1 ♀ 2 ♂ MSUC: RI, JBWM); Chippewa Co.: (no locality) 3 Jun. 1957, R. & K. Dreisbach (1 ♂ MSUC); (no locality), 25 Jun. 1960, R. & K. Dreisbach (1 ♀ MSUC); Dickinson Co.: T43N R29W Sec. 14, 24 Jun. 1983, Rubus allegheniensis, M. Arduser (1 ♀ MSUC); T43N R30W Sec. 18, 4 Jun. 1983, M. Arduser (1 ♀ MSUC); Gladwin Co.: (no locality) 19 Jun. 1943, R.R. Dreisbach (1 ♀ MSUC); Keweenaw Co.: (no locality) 26 Jun. 1955, R.R. Dreisbach (1 ♀ MSUC); Copper Harbor, 18 Jun. 1957, R.W. Hodges (1 ♀ MSUC); Mackinac Co.: (no locality) 7 Jun. 1960, R. & K. Dreisbach (1 ♀ MSUC); T43N R12W Sec.22,29 May 1965, R.W. Husband (1 ♀ MSUC); Marquette Co.: (no locality) 20 Jun. 1955, R.R. Dreisbach (1 ♀ MSUC).


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FIGURE 6. Michigan holotype material at the National Museum of Natural History, Smithisonian Institution. A) Andrena (Gonandrena) monroensis Mitchell, face (= A. platyparia Robertson). B) Andrena (Leucandrena) bifurcata Mitchell, lateral habitus (= A. erigeniae Robertson). C) Andrena (Gonandrena) dreisbachi Mitchell, face (= A. robertsonii Dalla Torre). D) Andrena (?Leucandrena) chippewaensis Mitchell, dorsal habitus (= A. wheeleri Graenicher). Photographs provided with the permission of the National Museum of Natural History, Smithsonian Institution, 10th and Constitution Ave. N.W., Washington, DC 20560-0193. (http://www.nmnh.si.edu/).

Subgenus Taeniandrena Hedicke Revision: LaBerge (1989).

Andrena (Taeniandrena) wilkella (Kirby 1802) County records: Alcona, Alger, Allegan, Arenac, Baraga, Barry, Bay, Berrien, Branch, Calhoun, Cass, Cheboygan, Chippewa, Clinton, Crawford, Delta, Dickinson, Eaton, Emmet, Genesee, Gladwin, Gogebic, Grand Traverse, Gratiot, Hillsdale, Houghton, Huron, Ingham, Ionia, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lake, Lapeer, Leelanau, Lenawee, Livingston, Luce, Mackinac, Macomb, Manistee, Marquette, Mason, Mecosta, Menominee, Midland, Missaukee, Monroe, Montcalm, Montmorency, Newaygo, Oakland, Oceana, Ogemaw, Ontonagon, Osceola, Oscoda, Otsego, Ottawa, Roscommon, Saginaw, Sanilac, Schoolcraft, Shiawassee, St. Clair, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Exotic (Linsley 1958). This common species prefers plants in the Fabaceae (LaBerge 1989). Nests in Nova Scotia were described by Atwood (1933).



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Subgenus Thysandrena Lanham Revision: LaBerge (1977).

Andrena (Thysandrena) bisalicis Viereck 1908 County records: Alger, Allegan, Antrim, Baraga, Bay, Benzie, Clare, Gladwin, Grand Traverse, Houghton, Iosco, Kalkaska, Kent, Lapeer, Leelanau, Mackinac, Mecosta, Midland, Missaukee, Newaygo, Oceana, Osceola, Roscommon. Notes. Specialist on Salix (Salicaceae).

Andrena (Thysandrena) w-scripta Viereck 1904 County records: Benzie, Clare, Crawford, Dickinson, Gladwin, Grand Traverse, Ingham, Kalkaska, Keweenaw, Lake, Lapeer, Leelanau, Mackinac, Marquette, Mecosta, Midland, Montmorency, Oceana, Osceola, Otsego, Presque Isle, Saginaw, Schoolcraft, Van Buren, Wexford.

Subgenus Trachandrena Robertson Revision: LaBerge (1973).

Andrena (Trachandrena) ceanothi Viereck 1917 = Andrena (Trachandrena) compacta Mitchell, 1960: 181 (Michigan holotype; Fig 7A). Holotype. ♂ USA: Michigan: Roscommon Co.: 26 May 1957, R. & K. Dreisbach (NMNH: 75174). Synonymy by LaBerge (1973).

County records: Allegan, Antrim, Arenac, Clare, Crawford, Emmet, Grand Traverse, Gratiot, Houghton, Ionia, Iosco, Iron, Kalamazoo, Kalkaska, Lake, Livingston, Mecosta, Midland, Missaukee, Montcalm, Monroe, Montmorency, Muskegon, Newaygo, Oakland, Osceola, Oscoda, Otsego, Ottawa, Presque Isle, Roscommon, Saginaw, Shiawassee, St. Joseph, Van Buren, Wayne, Wexford. Notes. LaBerge (1973) records A. ceanothi as a polylectic species, although it was reported as a Vaccinium specialist by Goldstein & Ascher (2016) in an analysis lumping oligoleges and mesoleges. Nearly half its flower records are from Rubus (Rosaceae), but it does also show some preference for Vaccinium and may prove to be a valuable wild pollinator of berry crops.

Andrena (Trachandrena) forbesii Robertson 1891 County records: Allegan, Antrim, Bay, Benzie, Berrien, Branch, Calhoun, Clinton, Crawford, Dickinson, Eaton, Gladwin, Grand Traverse, Gratiot, Houghton, Ingham, Ionia, Isabella, Kalamazoo, Kalkaska, Kent, Lake, Lapeer, Leelanau, Livingston, Mackinac, Macomb, Manistee, Marquette, Mason, Mecosta, Midland, Montcalm, Muskegon, Oakland, Oceana, Ogemaw, Osceola, Ottawa, Roscommon, Saginaw, St. Joseph, Van Buren, Washtenaw, Wayne, Wexford. Notes. Common wild pollinator of Michigan tart cherry and apple orchards (Gibbs et al. 2017).


GIBBS ET AL.

Andrena (Trachandrena) heraclei Robertson 1897 County records: Allegan. Notes. Andrena heraclei, originally described from Carlinville, Illinois, has been recorded previously from southern New England and New York, where it is rarely collected (JSA, unpublished), to Kansas (LaBerge 1973). These new records extend the northern range of the species in the Midwest. A male of A. heraclei was collected in a blueberry field, but misidentified as the similar species A. spiraeana Robertson (Tuell et al. 2009). Females of A. heraclei were collected at the same site nine years later (Gibbs et al. 2016). The females of A. heraclei and A. spiraeana are readily distinguished by the punctation of metasomal tergum 1 (LaBerge 1973). Material examined. Allegan Co.: Lee Twp., (3.6 mi. ESE Pullman), 25 May 2004 (1 ♀ MSUC: RI); Pullman, 5 km E, 26 May 2013 on Vaccinium corymbosum (2 ♀ MSUC: RI); Ottawa Co.: Holland, 4 mi. WNW, 17 May 2005 (1 ♂ MSUC: RI).

Andrena (Trachandrena) hippotes Robertson 1895 = Andrena (Trachandrena) arenakensis Mitchell, 1960: 177 (Michigan holotype; Fig 7B). Holotype. ♀ USA: Michigan: Arenac Co.: 30 May 1951, R.R. Dreisbach (NMNH: 75168). Stylopized. Synonymy by LaBerge (1973).

County records: Allegan, Alpena, Arenac, Benzie, Berrien, Calhoun, Clare, Clinton, Dickinson, Emmet, Genesee, Gladwin, Grand Traverse, Ingham, Iosco, Isabella, Kalamazoo, Kalkaska, Kent, Lake, Lapeer, Leelanau, Lenawee, Livingston, Macomb, Manistee, Midland, Montmorency, Oceana, Osceola, Ottawa, Saginaw, Sanilac, Van Buren, Washtenaw, Wexford. Notes. A common spring-flying species and a frequent visitor of tart cherry orchards (Gibbs et al. 2017).

Andrena (Trachandrena) mariae Robertson 1891 County records: Allegan, Clare, Clinton, Dickinson, Eaton, Gladwin, Grand Traverse, Ingham, Kalamazoo, Kalkaska, Lake, Leelanau, Lenawee, Livingston, Mecosta, Midland, Otsego, Van Buren, Washtenaw. Notes. Specialist on Salix (Salicaceae).

Andrena (Trachandrena) miranda Smith 1879 County records: Alger, Antrim, Arenac, Baraga, Bay, Branch, Calhoun, Cheboygan, Clinton, Crawford, Delta, Dickinson, Eaton, Emmet, Gladwin, Grand Traverse, Gratiot, Hillsdale, Ingham, Iosco, Iron, Kalamazoo, Kent, Keweenaw, Lake, Leelanau, Lenawee, Livingston, Luce, Macomb, Mecosta, Midland, Missaukee, Montcalm, Oakland, Ogemaw, Otsego, Ottawa, Presque Isle, Roscommon, Saginaw, Schoolcraft, St. Joseph, Van Buren, Washtenaw.

Andrena (Trachandrena) nuda Robertson 1891 County records: Allegan, Berrien, Calhoun, Eaton, Gladwin, Ingham, Jackson, Kalamazoo, Livingston, Monroe, Ottawa, St. Joseph, Van Buren.

Andrena (Trachandrena) rehni Viereck 1907 County records: Allegan, Eaton, Ottawa, Washtenaw. Notes. Rare in most regional bee collections, but may be overlooked due to identification difficulties.



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Andrena (Trachandrena) rugosa Cockerell 1906 County records: Allegan, Antrim, Benzie, Berrien, Dickinson, Eaton, Grand Traverse, Gratiot, Ingham, Kalamazoo, Leelanau, Livingston, Manistee, Montcalm, Muskegon, Newaygo, Oceana, Ottawa, St. Joseph, Van Buren, Washtenaw, Wexford. Notes. A relatively common vernal species, frequently collected on commercial apple, highbush blueberry and tart cherry flowers (Gibbs et al. 2016, 2017).

Andrena (Trachandrena) sigmundi Cockerell 1902 County records: Alger, Allegan, Antrim, Clare, Clinton, Crawford, Dickinson, Gladwin, Grand Traverse, Ingham, Iosco, Isabella, Lake, Livingston, Mackinac, Midland, Montmorency, Otsego, Roscommon, Sanilac, Schoolcraft. Notes. Specialist on Salix (Salicaceae) (LaBerge 1973).

Andrena (Trachandrena) spiraeana Robertson 1895 County records: Allegan, Ingham, Kalamazoo, Ottawa, Van Buren. Notes. See notes for A. heraclei above.

Andrena (Trachandrena) virginiana Mitchell 1960 County records: Branch, Clare, Delta, Ingham, Keweenaw, Livingston, Macomb, Mason, Midland, Ottawa. Notes. Polylectic, but may prefer to visit Ceanothus (Rhamnaceae) (LaBerge 1973).

Subgenus Tylandrena LaBerge Revision: LaBerge & Bouseman (1970).

Andrena (Tylandrena) erythrogaster (Ashmead 1890) County records: Allegan, Antrim, Arenac, Clare, Clinton, Eaton, Gladwin, Gratiot, Ingham, Ionia, Iosco, Isabella, Kalkaska, Lake, Mecosta, Midland, Montcalm, Newaygo, Oakland, Osceola, Saginaw, Schoolcraft, St. Clair, Van Buren, Washtenaw, Wayne. Notes. A specialist on Salix. See Miliczky (1988) for notes on nesting biology from central Illinois.

Andrena (Tylandrena) perplexa Smith 1853 County records: Allegan, Berrien, Calhoun, Cass, Clinton, Eaton, Genesee, Gladwin, Ingham, Ionia, Jackson, Kalamazoo, Kent, Lapeer, Livingston, Midland, Newaygo, Oceana, Ottawa, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw. Notes. Nesting biology and immatures studied by Stephen (1966a, b) in Oregon and nests from Washington D.C. described by Parker & Böving (1924).

Andrena (Tylandrena) wilmattae Cockerell 1906 = Andrena (Bythandrena) acra Mitchell, 1960: 111 (Michigan holotype; Fig 7C).


GIBBS ET AL.

Holotype. ♂ USA: Michigan: Kalamazoo Co.: Gull Lake Biol. Sta. (KBS), 9 Jun. 1956, on Crataegus, R.L. Fischer (NMNH: 75167). Synonymy by LaBerge & Bouseman (1970).

County records: Branch, Kalamazoo, Lake, Washtenaw. Notes. LaBerge & Bouseman (1970) state that this large spring-flying bee is rare in collections but the number of available Michigan records and recent collection of a long series from Illinois (Griffin et al. 2017) suggest that it can be locally numerous. Material examined. Branch Co.: (no locality) 1 Jun. 1941, R.R. Dreisbach (1 ♂ MSUC); Bronson, 14 Jun. 1959, R.L. Fischer (1 ♀ MSUC); Kalamazoo Co.: Gull Lake Biol. Sta. (KBS), 9 Jun. 1956 (3 ♂ paratypes (R.W. Hodges), 1 ♀ MSUC (R.L. Fischer)); Lake Co.: (no locality) 17 Apr. 1955, R.R. Dreisbach (1 ♂ MSUC).

Panurginae Calliopsini Genus Calliopsis Smith Taxonomy: Mitchell (1960); Shinn (1967). Behavior: Calliopsis are solitary and either polylectic or oligolectic.

Subgenus Calliopsis Smith s. s. Callopsis (Calliopsis) andreniformis Smith 1853 County records: Allegan, Baraga, Barry, Bay, Berrien, Calhoun, Cheboygan, Dickinson, Eaton, Genesee, Gogebic, Gratiot, Hillsdale, Houghton, Huron, Ingham, Kalamazoo, Kent, Keweenaw, Lake, Lenawee, Livingston, Manistee, Marquette, Mason, Missaukee, Montmorency, Oakland, Ontonagon, Otsego, Shiawassee, Van Buren, Washtenaw. Notes. Nests from Kansas, Texas, Tennessee, and New Jersey described by Shinn (1967) and (Rozen 1967). Notes on Iowa nests described by Ainslie (1937). This species is considered polyletic, but it is usually found in association with Fabaceae and may be an effective pollinator of alfalfa and cultivated clovers (Shinn 1967).

Subgenus Verbenapis Cockerell & Atkins Callopsis (Verbenapis) nebraskensis Crawford 1902 County records: Ionia, Lapeer, Livingston, St. Joseph. Notes. Specialist on Verbena (Verbenaceae) (Shinn 1967). Notes on nest sites in Missouri by Rau & Rau (1916).

Perditini Genus Perdita Smith Taxonomy: Mitchell (1960); Timberlake (1954, 1958, 1960, 1968).



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Subgenus Cockerellia Ashmead Revision: Timberlake (1954) Biology. Specialists on Asteraceae.

Perdita (Cockerellia) albipennis Cresson 1868 pallidipennis Graenicher 1910 County records: Allegan, Barry, Berrien, Kalamazoo. Notes. Specialist on Helianthus (Asteraceae). Nest from Kansas described by Danforth (1989). Wolf & Ascher (2009) expressed doubt regarding the status of this subspecies, and suggested it may be a subspecies of P. bequaerti Viereck (see below). If correct, then only a single species of Perdita (Cockerellia) would be present in Michigan.

Perdita (Cockerellia) bequaerti Viereck 1917 indianensis Cockerell 1922 County records: Allegan, Barry, Berrien, Cass, Gladwin, Ionia, Kent, Leelanau, Livingston, Van Buren. Notes. Specialist on Helianthus (Asteraceae).

Subgenus Perdita Smith s. s. Taxonomy: Timberlake (1958, 1960, 1968)

Perdita (Perdita) gerhardi gerhardi Viereck 1904 (octomaculata group) County records: Allegan, Barry, Cass, Newaygo, Van Buren. Notes. Specialist on Monarda punctata L. (Lamiaceae). Large numbers have been observed on dunes near Lake Michigan. Nesting in central Illinois described by Miliczky (1991).

Perdita (Perdita) halictoides Smith 1853 (halictoides group) County records: Barry, Berrien, Clinton, Ingham, Kalamazoo, Midland, Oscoda, Saginaw, St. Joseph, Van Buren, Washtenaw. Notes. Specialist on Physalis (Solanaceae). Nest from Florida described by Eickwort (1977).

Perdita (Perdita) maculigera Cockerell 1896 maculipennis Graenicher 1914 (octomaculata group) County records: Alcona, Arenac, Delta, Gratiot, Hillsdale, Midland, Newaygo, Saginaw, Tuscola, Washtenaw. Notes. Specialist on Salix (Salicaceae), especially sandbar willow along streams and rivers. Nesting biology in Kansas described by Michener & Ordway (1963).

Perdita (Perdita) octomaculata octomaculata (Say 1824) (octomaculata group) County records: Allegan, Arenac, Barry, Bay, Berrien, Clare, Clinton, Gladwin, Ingham, Isabella, Kalamazoo, Livingston, Midland, Oakland, Ottawa, Shiawassee, Van Buren, Washtenaw, Wayne. Notes. Specialist on Solidago (Asteraceae). Nests from New York described by Eickwort (1977).


GIBBS ET AL.

Perdita (Perdita) swenki Crawford 1915 (octomaculata group) County records: Alger, Cheboygan, Dickinson, Kalkaska, Leelanau, Mecosta, Van Buren. Notes. Asteraceae specialist.

FIGURE 7. Michigan holotype material at the National Museum of Natural History, Smithisonian Institution. A) Andrena (Trachandrena) compacta Mitchell, dorsal habitus (= A. ceanothi Viereck). B) Andrena (Trachandrena) arenakensis Mitchell, lateral habitus (= A. hippotes Robertson). C) Andrena (Bythandrena) acra Mitchell, lateral habitus (= A. wilmattae (Cockerell)). D) Bombus ternarius var. expallidus Cockerell, lateral habitus (subspecies not recognized). Photographs provided with the permission of the National Museum of Natural History, Smithsonian Institution, 10th and Constitution Ave. N.W., Washington, DC 20560-0193. (http://www.nmnh.si.edu/).

Protandrenini Genus Protandrena Cockerell Taxonomy: Timberlake (1976).

Protandrena bancrofti Dunning 1897 (New state record) (Fig. 8A) County records: Kalamazoo, St. Joseph. Notes. A prairie-associated species at the northeastern edge of its range. A recent record of this species from



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New Hampshire (Tucker & Rehan 2016) is almost certainly erroneous as there are no verified records east of Michigan and Indiana. Brief notes on nests in Indiana are available (Chandler 1962). Material examined. Kalamazoo Co.: Gull Lake Biol. Sta. (KBS), 9 Jul. 1959 (1 ♂ MSUC); St. Joseph Co.: White Pigeon, 14 Jun. 1959 (1 ♂ MSUC).

Protandrena cockerelli Dunning 1897 (Fig. 8B) County records: Berrien, Ionia, Newaygo. Notes. A small number of males and a single female of this species have been recently collected, notably all from Asclepias verticillata L (Asclepiadaceae). Mitchell (1960) recorded this species from Michigan as Psaenythia (Protandrena) mexicanorum (type locality: [Ciudad] Juárez, Chihuahua, Mexico], with cockerelli (type locality: Topeka, Kansas) as a synonym. Timberlake (1955) treated P. cockerelli, as a subspecies of mexicanorum. A female from Newaygo County identified by Mitchell (MSUC) has facial and pronotal maculations consistent with P. mexicanorum, but Hurd (1979) recorded the Michigan record as Protandrena cockerelli instead. Based on the geographic distribution of Protandrena species (Timberlake 1976), it is unlikely that P. mexicanorum sensu stricto would occur in Michigan, as the closest currently accepted P. mexicanorum record is from West Texas (Timberlake 1976). Thus, we attribute Hurd’s (1979) records of mexicanorum from North Dakota, Nebraska, and Colorado to outdated identifications or taxonomic concepts (Chandler 1962). To further add to confusion about the true distribution of these bees, Timberlake (1976) recorded P. cockerelli from New Jersey, but this record seems implausible and likely involves mislabeled or misinterpreted material from much further west in North America (perhaps from New Mexico). Material examined. Berrien Co.: Southwest Michigan Research and Education Center, 42.084 -86.3535, 14 Jul. 2016, Asclepias verticillata (1 ♂ MSUC: RI); Ionia Co.: Clarksville Research Center, 42.8708 -85.2544, 11 Aug. 2015, Asclepias verticillata L., L. Rowe (1 ♂ MSUC: RI); Clarksville Research Center, 42.8708 -85.2544, 3 Aug. 2016, Asclepias verticillata L., L. Rowe (1 ♂ MSUC: RI); Clarksville Research Center, 42.8708 -85.2544, 8 Aug. 2016, Asclepias verticillata L., L. Rowe (1 ♀ MSUC: RI); Newaygo Co.: (no locality), 13 May 1956, R. & K. Dreisbach (1 ♀ MSUC).

FIGURE 8. Protandrena from Michigan. A) Oblique habitus of male Protandrena bancrofti Dunning. B) Face of female Protandrena cockerelli Dunning from Michigan.

Genus Pseudopanurgus Cockerell Taxonomy: Mitchell (1960); Timberlake (1967, 1973); see also Scott et al. (2011).


GIBBS ET AL.

Pseudopanurgus aestivalis (Provancher 1882) (rudbeckiae group) County records: Allegan, Cheboygan, Iosco, Kalkaska, Lake, Leelanau, Livingston, Manistee, Oscoda, Van Buren. Notes. Visits Symphyotrichum and Solidago (Hurd 1979). Prior to Sheffield & Perron (2014), this species was commonly referred to as P. nebrascensis (Crawford).

Pseudopanurgus albitarsis (Cresson 1872) (rudbeckiae group) (New state record) (Fig. 9) County records: Berrien. Notes. A member of the rudbeckiae species group (i.e. Pterosarus Timberlake) often associated with prairies, but extends east locally to North Carolina (Mitchell, 1960) and Virginia (Droege, pers. comm., 2015) and north in the Midwestern United States to Wisconsin (Droege, pers. comm., 2007) in addition to Michigan. Not reported from the Mid-Atlantic or Northeastern United States north of Virginia other than a state record for Connecticut by Mitchell (1960) that we cannot confirm and consider unlikely. Material examined. Berrien Co.: Southwest Michigan Research and Education Center, 42.084 -88.3535, 23 Aug. 2016, Coreopsis tripteris (1 ♂ MSUC: RI).

FIGURE 9. Oblique habitus of male Pseudopanurgus albitarsis (Cockerell).



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Pseudopanurgus andrenoides (Smith 1853) (rudbeckiae group) County records: Bay, Clinton, Jackson, Kalamazoo, Midland. Notes. Specialist on Solidago (Asteraceae).

Pseudopanurgus compositarum (Robertson 1893) (rudbeckiae group) (New state record) County records: Ingham, Kalamazoo. Notes. A relatively scarce species at its northern range limits in the region. Like other members of the rudbeckiae group it is a specialist on Asteraceae. Michigan records in Zarrillo et al. (2016) are based on the specimens reported here. Material examined. Ingham Co.: Mason, 6 Sep. 2013, Symphyotrichum laeve (1 ♀), Solidago (1 ♀) (MSUC: RI); Kalamazoo Co.: Kalamazoo Nature Center, 5 Sep. 2013, J. Gibbs (JBWM).

Pseudopanurgus labrosiformis (Robertson 1898) (rudbeckiae group) (New state record) County records: Leelanau. Notes. An Astereaceae specialist recorded from Sleeping Bear Dunes by S. Droege (pers. comm.)

Pseudopanurgus rugosus (Robertson 1895) (aethiops group) (New state record) County records: Jackson, Kalamazoo. Notes. The only eastern member of the aethiops species group, i.e. Pseudopanurgus sensu stricto, is another example of a species occurring further north in the Midwestern states than in the Atlantic states, where it is recorded north only to Maryland. It is also a specialist on Asteraceae. Nesting biology in Illinois was described by Miliczky (1991). Material examined. Kalamazoo Co.: Gull Lake (W.K. Kellogg) Biol. Sta., 19 Aug. 1960 (1 ♀ MSUC); Sand Creek Preserve, N42.3496, W85.7393, 1 Aug. 2016, Coreopsis tripteris, S. Naranjo (1 ♂ MSUC: RI).

APIDAE Apinae Anthophorini Genus Anthophora Latreille Taxonomy. Mitchell (1962); Brooks (1983).

Subgenus Clisodon Patton Anthophora (Clisodon) terminalis Cresson 1869 County records: Alger, Allegan, Antrim, Baraga, Barry, Berrien, Cass, Cheboygan, Clinton, Crawford, Dickinson, Genesee, Gladwin, Gogebic, Ingham, Ionia, Iosco, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lapeer, Leelanau, Livingston, Mackinac, Manistee, Marquette, Mecosta, Menominee, Midland, Monroe, Montcalm, Oakland, Oceana, Oscoda, Ottawa, Presque Isle, Roscommon, Saginaw, Shiawassee, Van Buren, Washtenaw, Wayne.


GIBBS ET AL.

Notes. Trap nesting in sumac stems from Wisconsin described by Medler (1964b), Anthophora terminalis nests in wood, not just in stems.

Subgenus Melea Sandhouse Revision: Brooks (1983).

Anthophora (Melea) abrupta Say 1837 (bomboides group) County records: Berrien, Eaton, Ingham, Keweenaw, Livingston, Mackinac, Manistee, Mecosta, Monroe, Montcalm. Notes. Nests described by Norden (1984) and Rau (1926, 1929) from Maryland and Missouri, respectively.

Anthophora (Melea) bomboides Kirby 1837 (bomboides group) County records: Barry, Cheboygan, Dickinson, Emmet, Houghton, Huron, Ingham, Jackson, Keweenaw, Mackinac, Manistee, Montcalm, Saginaw, Washtenaw, Wayne. Notes. Nests of the Pacific Coast color form stanfordiana Cockerell were described by Nininger (1920) and by Brooks (1983).

Subgenus Mystacanthophora Brooks Anthophora (Mystacanthophora) walshii Cresson 1869 (montana group) County records: Allegan, Kalamazoo, Leelanau, Livingston, Newaygo, Shiawassee. Notes. Nesting notes in Kansas by Cane (1991). This species is notably rare and restricted in range in the Northeastern and Mid-Atlantic United States (Goldstein & Ascher 2016).

Genus Habropoda Smith Habropoda laboriosa (Fabricius 1804) (New state record) County records: Cass, Huron, Ingham, Livingston, Manistee, Midland, Oakland, Washtenaw. Notes. An important blueberry pollinator in southern parts of its range.

Apini Genus Apis Linnaeus Apis (Apis) mellifera Linnaeus 1758 County records: Alcona, Alger, Allegan, Alpena, Antrim, Arenac, Baraga, Barry, Bay, Benzie, Berrien, Branch, Calhoun, Cass, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Crawford, Delta, Dickinson, Eaton, Emmet, Genesee, Gladwin, Gogebic, Grand Traverse, Gratiot, Hillsdale, Houghton, Huron, Ingham, Ionia, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lake, Lapeer, Leelanau, Lenawee, Livingston, Luce, Mackinac, Macomb, Manistee, Marquette, Mason, Mecosta, Menominee, Midland, Missaukee, Monroe, Montcalm, Montmorency, Muskegon, Newaygo, Oakland, Oceana, Ogemaw, Ontonagon, Osceola, Oscoda, THE BEES OF MICHIGAN


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Otsego, Ottawa, Presque Isle, Roscommon, Saginaw, Sanilac, Schoolcraft, Shiawassee, St. Clair, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. This non-native managed species undoubtedly occurs in all counties. It is often the primary pollinator of commercial fruit and vegetable crops, but it may also have negative ecological impacts on native bees (Cane & Tepedino 2016; Lindström et al. 2016).

Bombini Genus Bombus Latreille Taxonomy: Milliron (1971, 1973a, b); Mitchell (1962); Laverty & Harder (1988); Williams et al. (2008, 2014). Biology. Bumble bees are primarily eusocial with annual colonies, with the exception of social parasites, which in the continental United States are entirely within the subgenus Psithyrus (Heinrich 2004; Husband et al. 1980; Michener 1974; Plowright & Laverty 1984). Nests are commonly made in abandoned rodent nests underground (Husband et al. 1980), but other similar cavities may be used, including walls of houses, and some species typically nest above ground (Richards 1978).

Subgenus Bombias Robertson Biology. Subgenus Bombias is unique among bumble bees in the isolation of eggs within the nest (Michener 1974).

Bombus (Bombias) auricomus (Robertson 1903) County records: Allegan, Arenac, Barry, Bay, Benzie, Berrien, Calhoun, Cass, Clinton, Eaton, Genesee, Gratiot, Hillsdale, Huron, Ingham, Ionia, Iosco, Isabella, Jackson, Kalamazoo, Kent, Lake, Lapeer, Lenawee, Livingston, Macomb, Manistee, Midland, Monroe, Montcalm, Muskegon, Newaygo, Oakland, Oceana, Ottawa, Saginaw, Sanilac, Shiawassee, St. Joseph, Van Buren, Washtenaw, Wayne.

Subgenus Bombus Latreille s. s. Bombus (Bombus) affinis Cresson 1863 County records: Alcona, Allegan, Alpena, Barry, Benzie, Berrien, Branch, Calhoun, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Emmet, Genesee, Gladwin, Grand Traverse, Gratiot, Hillsdale, Ingham, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Lapeer, Lenawee, Livingston, Macomb, Manistee, Marquette, Mason, Mecosta, Midland, Monroe, Montcalm, Montmorency, Newaygo, Oakland, Oceana, Ogemaw, Osceola, Ottawa, Saginaw, Sanilac, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. The US Fish and Wildlife Service has approved this species for endangered status, after being sued for failing to act on a petition to list the species in an appropriate time frame. Although described as ‘uncommon’ in Michigan by Husband et al. (1980), Bombus affinis is relatively well represented in the LP historically as represented by specimens deposited at MSUC. Bombus affinis has suffered a catastrophic decline in population size (Giles & Ascher 2006; Cameron et al. 2011). No specimens are known from Michigan since 2000, but with extensive sampling effort (e.g., Cameron et al. 2011), a small number of recent specimens (since 2001) have been collected, photographed (see http://bugguide.net and http://bumblebeewatch.org), or observed in Ontario, Illinois, Indiana, Iowa, Maryland, Minnesota, Ohio, Virginia, and Wisconsin. A specimen collected in 2013 from Toledo, Lucas Co., Ohio, less than 13 km south of the Michigan border (http://bumblebeewatch.org), suggests that this species may persist in southeastern Michigan.


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Bombus (Bombus) terricola Kirby 1837 County records: Alcona, Alger, Allegan, Alpena, Antrim, Arenac, Baraga, Bay, Benzie, Cass, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Crawford, Delta, Dickinson, Emmet, Gladwin, Gogebic, Grand Traverse, Gratiot, Hillsdale, Houghton, Huron, Ingham, Ionia, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lake, Leelanau, Lenawee, Livingston, Luce, Mackinac, Macomb, Manistee, Marquette, Mason, Mecosta, Menominee, Midland, Missaukee, Montcalm, Montmorency, Muskegon, Newaygo, Oakland, Oceana, Ogemaw, Ontonagon, Osceola, Oscoda, Otsego, Presque Isle, Roscommon, Saginaw, Schoolcraft, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Bombus terricola has undergone large contractions in its US distribution (Cameron et al. 2011) but with evidence of a recent partial recovery in some areas. Historically, the primarily northern species has been recorded throughout Michigan, but only rarely in the SLP (Husband et al. 1980). Recent collections examined are few and restricted to the NLP in Antrim, Leelanau, and Oceana Counties and Keweenaw County in the UP.

Subgenus Cullumanobombus Vogt Bombus (Cullumanobombus) fraternus (Smith 1854) (robustus group) Included based on the the authority of Franklin (1912), Frison (Milliron 1939), and Mitchell (1962), but no specimens of this primarily southern species have been examined.

Bombus (Cullumanobombus) griseocollis (DeGeer 1773) (griseocollis group) County records: Alcona, Allegan, Alpena, Arenac, Barry, Bay, Benzie, Berrien, Branch, Calhoun, Cass, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Crawford, Delta, Dickinson, Eaton, Emmet, Genesee, Gladwin, Grand Traverse, Gratiot, Hillsdale, Huron, Ingham, Ionia, Iosco, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Lapeer, Leelanau, Lenawee, Livingston, Mackinac, Macomb, Manistee, Mason, Mecosta, Midland, Missaukee, Monroe, Montcalm, Montmorency, Muskegon, Newaygo, Oakland, Oceana, Ogemaw, Osceola, Oscoda, Otsego, Ottawa, Roscommon, Saginaw, Sanilac, Shiawassee, St. Clair, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Nests are made at the surface (Plath 1927).

Bombus (Cullumanobombus) rufocinctus Cresson 1863 (rufocinctus group) County records: Alger, Alpena, Baraga, Bay, Charlevoix, Cheboygan, Chippewa, Clare, Emmet, Gladwin, Gogebic, Grand Traverse, Gratiot, Huron, Ingham, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lapeer, Livingston, Luce, Mackinac, Manistee, Mason, Mecosta, Menominee, Midland, Missaukee, Montcalm, Montmorency, Muskegon, Oakland, Ogemaw, Osceola, Otsego, Presque Isle, Saginaw, Shiawassee, Tuscola, Washtenaw, Wayne, Wexford. Notes. Nests are made both underground and at the surface. A highly variable species, uncommonly seen in the state.

Subgenus Psithyrus Lepeletier Biology. All members of this subgenus are social parasites of other bumble bees.

Bombus (Psithyrus) ashtoni (Cresson 1864) (bohemicus group) County records: Alcona, Alger, Allegan, Alpena, Arenac, Baraga, Barry, Bay, Benzie, Charlevoix, Cheboygan,



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Chippewa, Clare, Clinton, Crawford, Delta, Dickinson, Emmet, Gladwin, Gogebic, Grand Traverse, Gratiot, Houghton, Huron, Ingham, Iosco, Iron, Isabella, Kalamazoo, Kalkaska, Keweenaw, Lake, Leelanau, Lenawee, Livingston, Luce, Mackinac, Manistee, Marquette, Mason, Mecosta, Menominee, Midland, Missaukee, Montmorency, Oakland, Ogemaw, Ontonagon, Osceola, Oscoda, Otsego, Presque Isle, Roscommon, Schoolcraft, Shiawassee, Washtenaw, Wexford. Notes. Bombus ashtoni is a social parasite of B. affinis and B. terricola (Plath 1922, 1927). The severe declines in B. affinis and range contractions to the north of B. terricola (Cameron et al. 2011) imply even greater declines in B. ashtoni, decline of which was first noted in the late 1990s at Ithaca, Tompkins Co., New York by JSA (pers. obs.). To our knowledge, no specimen has been collected in Michigan since at least 1993 (OSUC record), but there has been a lack of recent collections in the UP and NLP where B. terricola is still present. Williams et al. (2014) treat B. ashtoni as a junior synonym of B. bohemicus Seidl, 1838, a relatively common species in the Palaearctic region. There are subtle differences between the two in male genitalia (Williams 1991), but DNA sequence data show few differences (Cameron et al. 2007; Williams et al. 2014). Because of this synonymy, Hatfield et al. (2016) list this species as data deficient due to the need for a global analysis, although the North American populations are recognized as critically endangered. Lack of recent records in both specimen and image databases (e.g., http:// bugguide.net) validates concern about the status of North American populations of this bee.

Bombus (Psithyrus) citrinus (Smith 1854) (citrinus group) County records: Alger, Allegan, Alpena, Antrim, Arenac, Barry, Bay, Berrien, Calhoun, Cass, Cheboygan, Chippewa, Clinton, Delta, Dickinson, Emmet, Genesee, Gladwin, Grand Traverse, Gratiot, Hillsdale, Huron, Ingham, Ionia, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lake, Lapeer, Lenawee, Livingston, Luce, Mackinac, Macomb, Marquette, Mecosta, Menominee, Midland, Missaukee, Monroe, Montcalm, Muskegon, Newaygo, Oakland, Oceana, Osceola, Oscoda, Otsego, Ottawa, Presque Isle, Roscommon, Saginaw, Schoolcraft, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Social parasite of B. impatiens, B. bimaculatus and B. vagans.

Bombus (Psithyrus) fernaldae (Franklin 1911) (sylvestris group) County records: Alger, Alpena, Baraga, Charlevoix, Cheboygan, Chippewa, Gogebic, Huron, Ingham, Kalamazoo, Keweenaw, Livingston, Luce, Marquette, Schoolcraft. Notes. Social parasite of B. rufocinctus. No recent specimens examined, although it still occurs in the eastern United States. Williams et al. (2014) treat B. fernaldae as a junior synonym of B. flavidus Eversmann.

Bombus (Psithyrus) insularis (Smith 1861) (citrinus group) County records: Alger, Alpena, Baraga, Charlevoix, Cheboygan, Chippewa, Crawford, Delta, Emmet, Ingham, Kalamazoo, Luce, Mackinac, Marquette, Menominee, Schoolcraft. Notes. Social parasite of B. fervidus, B. rufocinctus, B. ternarius and B. terricola. No recent specimens examined from Michigan, and the species has evidently declined more generally in its limited (northern) historical range in the eastern United States, although it remains numerous further north in Canada and further west in North America.

Subgenus Pyrobombus Dalla Torre Bombus (Pyrobombus) bimaculatus Cresson 1863 (lapponicus group) County records: Allegan, Alpena, Antrim, Arenac, Barry, Berrien, Branch, Calhoun, Cass, Cheboygan,


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Chippewa, Clare, Eaton, Emmet, Gladwin, Grand Traverse, Gratiot, Hillsdale, Huron, Ingham, Ionia, Iosco, Jackson, Kalamazoo, Kent, Leelanau, Lenawee, Livingston, Mackinac, Macomb, Manistee, Mason, Mecosta, Midland, Missaukee, Monroe, Montcalm, Oakland, Oceana, Oscoda, Ottawa, Presque Isle, Roscommon, Saginaw, Schoolcraft, Shiawassee, St. Clair, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford.

Bombus (Pyrobombus) frigidus Smith 1854 (pratorum group) County records: None. Notes. This is an alpine/boreal species recorded from Michigan based on misidentified specimens, but it may nevertheless occur in the state. Milliron (1939) included specimens from Isle Royale identified by Frison as B. frigidus var. couperi. These were re-examined by JG (MSUC, INHS) and were determined to be B. sandersoni, which is consistent with the couperi epithet being a junior synonym of B. sandersoni. Wolf & Ascher (2009) record this species from Oneida County, Wisconsin, which is directly south of Michigan’s Upper Peninsula. Laverty & Harder (1988) also map this species to the Thunder Bay area of Ontario, very near Isle Royale. The distribution of the species is such that the UP is likely part of its southern range in the east (Williams et al. 2014).

Bombus (Pyrobombus) impatiens Cresson 1863 (lapponicus group) County records: Alcona, Allegan, Alpena, Antrim, Barry, Bay, Benzie, Berrien, Branch, Calhoun, Cass, Charlevoix, Cheboygan, Clare, Clinton, Crawford, Eaton, Emmet, Genesee, Gladwin, Grand Traverse, Gratiot, Hillsdale, Huron, Ingham, Ionia, Iosco, Isabella, Jackson, Kalamazoo, Kent, Lake, Lapeer, Leelanau, Lenawee, Livingston, Mackinac, Macomb, Manistee, Mason, Mecosta, Menominee, Midland, Missaukee, Monroe, Montcalm, Muskegon, Newaygo, Oakland, Oceana, Osceola, Ottawa, Saginaw, Sanilac, St. Clair, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Bombus impatiens is often the most common wild pollinator of agricultural crops in Michigan. It is abundant as queens and workers in spring crops such as apple, blueberry and tart cherry (Gibbs et al. 2016; Russo et al. 2015) and the workers visit field crops such as cucumber and pumpkin (Artz & Nault 2011; Petersen et al. 2013; Smith et al. 2013). It nests underground (Plath 1922). Commercially produced colonies of this species are available, with a production facility located in Michigan.

Bombus (Pyrobombus) perplexus Cresson 1863 (hypnorum group) County records: Alcona, Alger, Allegan, Alpena, Antrim, Baraga, Barry, Benzie, Calhoun, Charlevoix, Cheboygan, Chippewa, Clare, Delta, Dickinson, Emmet, Gladwin, Gogebic, Gratiot, Hillsdale, Houghton, Huron, Ingham, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kent, Keweenaw, Lapeer, Leelanau, Lenawee, Livingston, Luce, Mackinac, Manistee, Marquette, Mason, Mecosta, Missaukee, Montcalm, Montmorency, Oakland, Oceana, Ontonagon, Osceola, Oscoda, Ottawa, Saginaw, Schoolcraft, Shiawassee, St. Clair, Van Buren, Washtenaw. Notes. Nests at surface and in hollow of trees and logs.

Bombus (Pyrobombus) sandersoni Franklin 1913 (pratorum group) County records: Alger, Baraga, Cheboygan, Chippewa, Keweenaw, Marquette.

Bombus (Pyrobombus) ternarius Say 1837 (lapponicus group) = Bombus ternarius var. expallidus Cockerell 1916: 9. (Michigan holotype; Fig. 7D) Holotype. ♀ USA: Michigan: Schoolcraft Co.: Floodwood, 20 Jul. 1915, F.M. Gaige (NMNH: 40085).



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County records: Alcona, Alger, Alpena, Arenac, Baraga, Benzie, Charlevoix, Cheboygan, Chippewa, Crawford, Delta, Dickinson, Emmet, Gladwin, Gogebic, Grand Traverse, Gratiot, Houghton, Huron, Ingham, Iosco, Iron, Kalkaska, Kent, Keweenaw, Leelanau, Luce, Mackinac, Marquette, Mason, Mecosta, Menominee, Midland, Missaukee, Montcalm, Montmorency, Oakland, Oceana, Ogemaw, Ontonagon, Oscoda, Otsego, Presque Isle, Roscommon, Saginaw, Schoolcraft, Shiawassee, Van Buren, Wayne, Wexford. Notes. The holotype of B. t. expallidus differs from most B. ternarius in having metasomal terga 2 and 3 entirely yellow (Cockerell 1916; Williams et al. 2014). Bumble bees routinely vary in color due to both genetic variation and wear, and it is not currently recognized as a valid taxon.

Bombus (Pyrobombus) vagans vagans Smith 1854 (vagans group) County records: Alcona, Alger, Allegan, Alpena, Antrim, Arenac, Baraga, Barry, Bay, Benzie, Berrien, Branch, Calhoun, Cass, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Crawford, Delta, Dickinson, Eaton, Emmet, Genesee, Gladwin, Gogebic, Grand Traverse, Gratiot, Hillsdale, Huron, Ingham, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lake, Lapeer, Leelanau, Lenawee, Livingston, Luce, Mackinac, Macomb, Manistee, Marquette, Mason, Mecosta, Menominee, Midland, Missaukee, Monroe, Montcalm, Montmorency, Muskegon, Newaygo, Oakland, Oceana, Ogemaw, Ontonagon, Osceola, Oscoda, Otsego, Ottawa, Presque Isle, Roscommon, Saginaw, Schoolcraft, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Nests underground and at surface (Plath 1922).

Subgenus Subterraneobombus Vogt Bombus (Subterraneobombus) borealis Kirby 1837 County records: Alcona, Alger, Alpena, Arenac, Baraga, Barry, Bay, Benzie, Berrien, Calhoun, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Delta, Dickinson, Eaton, Emmet, Grand Traverse, Houghton, Huron, Ingham, Iosco, Iron, Isabella, Jackson, Kalamazoo, Keweenaw, Lake, Leelanau, Lenawee, Livingston, Mackinac, Manistee, Marquette, Mason, Mecosta, Menominee, Midland, Missaukee, Montmorency, Oakland, Oceana, Ontonagon, Osceola, Otsego, Ottawa, Presque Isle, Saginaw, Schoolcraft, St. Clair, Van Buren, Washtenaw, Wexford.

Subgenus Thoracobombus Dalla Torre Bombus (Thoracobombus) fervidus (Fabricius 1798) (pensylvanicus group) County records: Alcona, Alger, Allegan, Alpena, Arenac, Barry, Bay, Benzie, Berrien, Branch, Calhoun, Cass, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Crawford, Delta, Dickinson, Eaton, Emmet, Genesee, Gladwin, Gogebic, Grand Traverse, Gratiot, Hillsdale, Huron, Ingham, Ionia, Iosco, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Lapeer, Leelanau, Lenawee, Livingston, Luce, Mackinac, Macomb, Manistee, Mason, Mecosta, Menominee, Midland, Missaukee, Monroe, Montcalm, Muskegon, Newaygo, Oakland, Oceana, Ogemaw, Ontonagon, Osceola, Otsego, Ottawa, Presque Isle, Roscommon, Saginaw, Sanilac, Schoolcraft, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Nest locations may be above or below ground (Plath 1922).

Bombus (Thoracobombus) pensylvanicus (DeGeer 1773) (pensylvanicus group) County records: Alger, Allegan, Alpena, Arenac, Barry, Bay, Benzie, Berrien, Branch, Calhoun, Cass,


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Cheboygan, Chippewa, Clare, Clinton, Eaton, Gladwin, Gogebic, Hillsdale, Huron, Ingham, Ionia, Iosco, Isabella, Jackson, Kalamazoo, Kent, Lapeer, Lenawee, Livingston, Macomb, Manistee, Mason, Mecosta, Midland, Monroe, Muskegon, Newaygo, Oakland, Oceana, Otsego, Saginaw, Sanilac, Shiawassee, St. Clair, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne. Notes. No recent specimens examined. Bombus pensylvanicus is evidently a declining species in the northern extent of its historical range (Grixti et al. 2009), but is still commonly seen in the south and western part of its range (Cameron et al. 2011; many recent records at http://bugguide.net/node/view/65631).

Eucerini Genus Eucera Scopoli Subgenus Synhalonia Patton Revision: Timberlake (1969)

Eucera (Synhalonia) atriventris (Smith 1854) County records: Allegan, Berrien, Cass, St. Joseph, Van Buren. Notes. First recorded from Michigan by Tuell et al. (2009) based on a single specimen. Dozens of specimens captured in blue vane traps during spring in the southwest have been subsequently examined (Gibbs et al. 2017).

Eucera (Synhalonia) dubitata (Cresson 1878) (New state record) County records: Cass. Notes. This species could be easily mistaken for E. atriventris, which is relatively abundant in the southwestern part of the state. Females can be distinguished from E. atriventris by the paler hind tibial scopa and the highly polished integument of the mesoscutum. Timberlake (1969) recorded E. dubitata from both Carlinville, Illinois and Columbus, Ohio. Material examined. Cass Co.: Edward Lowe Foundation, 5 km NE Cassopolis, N41.956 W85.982, 6 Jun. 2013, Trifolium incarnatum (1 ♀ JBWM); Edward Lowe Foundation, 5 km NE Cassopolis, N41.954 W85.988, 2 Jun. 2014, Lupinus perennis (1 ♀ MSUC: RI).

Eucera (Synhalonia) hamata (Bradley 1942) County records: Allegan, Barry, Berrien, Calhoun, Cass, Clinton, Ionia, Jackson, Kalamazoo, Lapeer, Livingston, Montcalm, Oakland, Oceana, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne. Notes. Nests in central Illinois described by Miliczky (1985 as Tetralonia). This species may be expanding its range northward in some areas (see Zarrillo et al. 2016).

Genus Florilegus Robertson Subgenus Florilegus Robertson s. l. Florilegus (Florilegus) condignus (Cresson 1878) County records: Barry, Kalamazoo. Notes. These records are at the northern limit of the range of F. condignus, which extends into South America. THE BEES OF MICHIGAN


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Nesting biology in Nebraska described by LaBerge & Ribble (1966a). Where abundant, it has been recorded as an important pollinator of alfalfa.

Genus Melissodes Latreille Taxonomy: LaBerge (1956a, b, 1961); Mitchell (1962).

Subgenus Apomelissodes LaBerge Revision: LaBerge (1956b).

Melissodes (Apomelissodes) apicatus Lovell & Cockerell 1906 County records: Allegan, Missaukee, Tuscola, Wexford. Notes. Specialist on Pontederia (Pontederiaceae), which grows on lake and river margins. First recorded from Michigan by Tuell et al. (2009). Its occurrence in Wexford County is a notable northern extension of its range.

Subgenus Eumelissodes LaBerge Revision: LaBerge (1961).

Melissodes (Eumelissodes) agilis Cresson 1878 County records: Allegan, Barry, Berrien, Cass, Grand Traverse, Eaton, Huron, Ingham, Jackson, Kalamazoo, Kent, Lenawee, Macomb, Manistee, Midland, Montcalm, Muskegon, Oakland, Ottawa, Saginaw, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne. Notes. Specialist on Helianthus (Asteraceae). Ground nest in Missouri recorded by Rau (1922).

Melissodes (Eumelissodes) denticulatus Smith 1854 County records: Allegan, Ingham, Kalamazoo, Kent, Keweenaw, Newaygo, Saginaw, Van Buren, Washtenaw, Wayne. Notes. Specialist on Vernonia (Asteraceae).

Melissodes (Eumelissodes) dentiventris Smith 1854 County records: Barry, Hillsdale, Kalamazoo, Livingston. Notes. Specialist on Asteraceae, particularly Helianthus and Chrysopsis.

Melissodes (Eumelissodes) druriellus (Kirby 1802) County records: Allegan, Barry, Benzie, Calhoun, Cheboygan, Clinton, Dickinson, Gladwin, Gogebic, Gratiot, Hillsdale, Huron, Ingham, Ionia, Iron, Jackson, Kalamazoo, Kent, Keweenaw, Leelanau, Livingston, Luce, Manistee, Mecosta, Menominee, Midland, Muskegon, Newaygo, Oakland, Oceana, Osceola, Otsego, Ottawa, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Specialist on Asteraceae. Nesting in Arkansas and Wyoming described by Cameron et al. (1996) and Clement (1973), respectively (as rustica).


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Melissodes (Eumelissodes) illatus Lovell & Cockerell 1906 County records: Alger, Baraga, Bay, Cheboygan, Delta, Dickinson, Emmet, Keweenaw, Leelanau, Mackinac, Marquette, Midland, Montcalm, Oakland, Oceana, Ogemaw, Ontonagon. Notes. Specialist on Solidago and Symphyotrichum (Asteraceae).

Melissodes (Eumelissodes) niveus Robertson 1895 County records: Livingston. Notes. First recorded by Evans (1986). Specialist on Asteraceae.

Melissodes (Eumelissodes) subillatus LaBerge 1961 Melissodes (Eumelissodes) subillata, LaBerge, 1961: 568. (Michigan holotype) Holotype. ♀ USA: Michigan: Washtenaw Co.: Ann Arbor, 17 Jun. 1952, L. H. Shinners on Gaillardia aristata (SEMC).

County records: Arenac, Barry, Berrien, Cheboygan, Clinton, Eaton, Houghton, Ingham, Kalamazoo, Leelanau, Livingston, Midland, Montcalm, Oakland, Oceana, Ottawa, Saginaw, Van Buren, Washtenaw. Notes. This Asteraceae specialist is relatively common in the LP. In addition to the holotype, LaBerge (1961) lists the following Michigan locations in the original description: Arenac Co., Cheboygan Co.: Cheboygan, Douglas Lake, Clinton Co., Eaton Co., Ingham Co.: East Lansing, Eaton Co., Midland Co., Oakland Co., and Oceana Co.. It is easily confused with other Melissodes species, in particular M. illatus Lovell and Cockerell and M. denticulatus Smith.

Melissodes (Eumelissodes) tinctus LaBerge 1961 County records: Dickinson, Ionia, Lake, Leelanau, Livingston, Ottawa, Van Buren. Notes. Specialist on Asteraceae.

Melissodes (Eumelissodes) trinodis Robertson 1901 County records: Allegan, Cass, Emmet, Kalamazoo, Saginaw, Van Buren, Wayne. Notes. Specialist on Asteraceae. Nests in the ground (Graenicher 1905).

Melissodes (Eumelissodes) wheeleri Cockerell 1906 County records: Alger, Leelanau, Livingston, Roscommon. Notes. Specialist on Asteraceae. Specimens from Pictured Rocks National Park and Sleeping Bear Dunes National Lakeshore from PWRC identified by Karen Wright.

Subgenus Heliomelissodes LaBerge Revision: LaBerge (1956b).

Melissodes (Heliomelissodes) desponsus Smith 1854 County records: Alger, Allegan, Arenac, Barry, Benzie, Cass, Cheboygan, Delta, Dickinson, Eaton, Emmet,



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Gladwin, Gogebic, Gratiot, Huron, Ingham, Ionia, Iosco, Kalamazoo, Kent, Lapeer, Livingston, Macomb, Mason, Mecosta, Midland, Montcalm, Montmorency, Oakland, Oceana, Osceola, Otsego, Ottawa, Saginaw, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne. Notes. Specialist on Cirsium (Asteraceae).

Subgenus Melissodes Latreille s. s. Revision: LaBerge (1956a).

Melissodes (Melissodes) bimaculatus bimaculatus (Lepeletier 1825) County records: Allegan, Barry, Berrien, Calhoun, Cass, Clinton, Eaton, Huron, Ingham, Jackson, Kalamazoo, Kent, Lenawee, Livingston, Marquette, Midland, Montcalm, Muskegon, Newaygo, Oakland, Ottawa, Saginaw, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne. Notes. A very broadly polylectic species, known to even visit corn (Terrell & Batra 1984) and a potentially valuable pollinator of summer crops such as cucumber. Ashmead (1894) records M. bimaculatus nesting in an open field, with the entrance under a stone.

Melissodes (Melissodes) communis communis Cresson 1878 County records: Allegan, Barry, Clinton, Ionia, Leelanau, Livingston, Montcalm, Ottawa. Notes. Broadly polylectic.

Genus Peponapis Robertson Revision: Hurd & Linsley (1964).

Subgenus Peponapis Robertson s. l. Peponapis (Peponapis) pruinosa (Say 1837) County records: Barry, Cass, Eaton, Ingham, Isabella, Jackson, Midland, Montcalm, Oakland, Oceana, Saginaw, Shiawassee, St. Joseph, Tuscola, Washtenaw, Wayne. Notes. A cucurbit specialist. This is an important native pollinator of cucurbit crops in Michigan (Quinn 2015). Nests from California and Rhode Island were described by Hurd et al. (1974) and Mathewson (1968), respectively. Population genetics of the species in North America were investigated by López-Uribe et al. (2016).

Genus Svastra Holmberg Subgenus Epimelissodes Ashmead Revision: LaBerge (1956a).

Svastra (Epimelissodes) obliqua obliqua (Say 1837) (obliqua group) County records: Alger, Berrien, Cass, Isabella, Mackinac, St. Joseph. Notes. Nesting biology in Florida described by Rozen (1964). More county records are suggested by the map


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attributed to this species in Hurd & Linsley (1980), but this map evidently pertains to Melissodes agilis due to an error, i.e. fig. 9 in that paper pertains to M. agilis whereas their fig. 10 pertains to S. obliqua. Based on the distribution of counties, it may occur throughout the state, but it is very uncommon.

Nomadinae Ammobatoidini Genus Holcopasites Ashmead Taxonomy: Mitchell (1962); Hurd & Linsley (1972).

Holcopasites calliopsidis calliopsidis (Linsley 1943) (heliopsis group) County records: Gogebic, Hillsdale, Houghton, Ingham, Ionia, Kalamazoo, Livingston, Monroe, St. Joseph, Van Buren. Notes. Cleptoparasite of Calliopsis andreniformis (Ainslie 1937; Hurd & Linsley 1972; Rozen 1966, 1967; Shinn 1967; Swenk 1907) and possibly Pseudopanurgus (Linsley et al. 1956).

Epeolini Genus Epeolus Latreille Taxonomy: Mitchell (1962); Brumley (1965); Onuferko (2017). Behavior: Cleptoparasites. All known hosts of Epeolus are in the genus Colletes (Michener 2007).

Epeolus ainsliei Crawford 1932 (New state record) County records: Livingston. Notes. Possibly a cleptoparasite of Colletes susannae Swenk. The two were recently collected together in relative abundance in Birds Hill Provincial Park, Manitoba, and have been found together in the past (Wolf & Ascher 2009). However, Colletes susannae is not known to occur in Michigan and the host plant of this bee, Dalea purpurea Vent., is rare and possibly adventive in the state (Voss & Reznicek 2012). Colletes americanus Cresson has also been collected in association with E. ainsliei (Wolf & Ascher 2009). The closest previous record of the prairie-associated E. ainsliei was from Wisconsin (Wolf & Ascher 2009). Material examined. Livingston Co.: E. S. George Reserve, 19 Jul. 1960, F.C. Evans (1 ♀ UMMZ), 2 Aug. 1960, U.N. Lanham (3♀ UMMZ).

Epeolus americanus (Cresson 1878) = Epeolus lanhami Mitchell, 1962: 450. (Michigan holotype). Synonymy by Onuferko (2017). Holotype. ♀ USA: Michigan: Washtenaw Co.: near Saline, 26 Jun. 1954, U.N. Lanham (UCMC).

County records: Kalkaska, Keweenaw, Livingston, Midland, Washtenaw. Notes. Epeolus americanus has a primarily western distribution, but occurs in parts of eastern Canada and Michigan where it is uncommonly collected (Brumley 1965; Onuferko 2017). Material examined. Kalkaska Co.: (no locality) 19 Jun.?49 (illegible), R.R. Dreisbach (1 ♂ E. lanhami paratype); Keweenaw Co.: Isle Royale, 3 Jul. 1957, Trifolium repens, R.W. Hodges (1 ♀ MSUC).



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Additional material. Washtenaw Co.: near Saline, 26 Jun. 1954, U.N. Lanham (UCMC).

Epeolus autumnalis Robertson 1902 County records: None. Notes. Recorded from Michigan by Mitchell (1962). The record is supported by the regional distribution of the species and its likely host, C. compactus compactus Cresson (see Ascher et al. 2014)

Epeolus bifasciatus Cresson 1864 County records: Berrien, Clinton, Ingham, Kalamazoo, Livingston, Manistee, Midland, Oakland, Shiawassee, Washtenaw. Notes. A cleptoparasite of Colletes latitarsis Robertson (see Brumley 1965).

Epeolus canadensis Mitchell 1962 County records: Cheboygan, Livingston. Notes. Considered by JSA (unpublished) to be a cleptoparasite of C. kincaidii Cockerell based on historical (but not recent) co-occurrence of these species at a single site in Tompkins County, New York.

Epeolus interruptus Robertson 1900 County records: Cheboygan, Kalamazoo, Lake, Livingston, Midland. Note. A possible cleptoparasite of Colletes aestivalis Patton according to Brumley (1965), but this host is very rare in Michigan (and elsewhere) with no recent records from the state suggesting that an associated parasite would be of conservation concern.

Epeolus lectoides Robertson 1901 County records: Cass, Ionia, Kalamazoo, Kalkaska, Livingston, Shiawassee, St. Joseph, Van Buren, Washtenaw. Note. Its presumed host is Colletes nudus Robertson (see Ascher et al. 2014).

Epeolus minimus (Robertson 1902) County records: Alpena, Cheboygan. Notes. Graenicher (1906) recorded E. minimus as a cuckoo of Colletes kincaidii (as C. eulophi Robertson).

Epeolus pusillus Cresson 1864 County records: Livingston, Mecosta, Midland, Oakland, Washtenaw, Wayne. Notes. Reported to be a cleptoparasite of Colletes compactus (see Rozen & Favreau 1968), but the small body size of E. pusillus and its limited occurrence in certain localities (e.g., Tompkins County, New York) suggest an alternative or additional association with C. americanus (see Ascher et al. 2014).


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Epeolus scutellaris Say 1824 County records: Alger, Bay, Cheboygan, Emmet, Ingham, Kalamazoo, Kalkaska, Livingston, Menominee, Midland, Washtenaw. Notes. Presumably a cleptoparasite of Colletes simulans armatus Patton (Ascher et al. 2014).

Genus Triepeolus Robertson Revision: Rightmyer (2008). Biology: Triepeolus are cleptoparasites, primarily of the tribe Eucerini.

Triepeolus concavus (Cresson 1878) County records: Ingham. Notes. Cleptoparasite of Svastra obliqua obliqua. Single specimen at MSUC with "Ag. Coll." label. Recorded from Michigan by Rightmyer (2008).

Triepeolus cressonii (Robertson 1897) County records: Livingston.

Triepeolus donatus (Smith 1854) County records: Gladwin, Huron, Ingham, Kalamazoo, Livingston, Menominee, Shiawassee. Notes. Cleptoparasite of Melissodes desponsus (see Rightmyer 2008).

Triepeolus eliseae Rightmyer, new species (verbesinae group) (Figs 10, 11) Diagnosis. This is the only species of the Triepeolus verbesinae (Cockerell) species group found in eastern North America. Females are known by the circular pseudopygidial area (Fig. 10A) and strongly downturned S5 (Fig. 10B), both characters that are diagnostic for the species group as a whole in North America (Rightmyer 2008: 131). In addition, T. eliseae is one of a few Triepeolus species in which the banding of hairs on the body are extremely pale yellow to white (sometimes appearing grey) (Fig. 10C). Among those very pale yellow species, females and males of T. eliseae are known from other eastern North American species by the lack of a preoccipital carina on the dorsal margin of the head, short erect hairs on the mesepisternum (such hairs 0.5 OD or less in length) (Fig. 10D), absence of a midline on the clypeus (Fig. 10E), and short, rounded axillar spines (Fig. 10F). Males of T. eliseae might be confused with T. obliteratus but can be separated by the pygidial plate, which is very narrow (ca. 1 OD in width) and parallel-sided in T. eliseae, and wider and more triangular in appearance in T. obliteratus, and by the axillar spines, which are somewhat rounded apically and do not reach the scutellar midpoint in T. eliseae (reaching or exceeding scutellar midpoint in T. obliteratus). Description. Length ca. 8–9 mm; ITW 1.6–1.9 mm. Integument black, orange on basal half of mandible, tegula, and legs, sometimes orange on apical margin of labrum, outer F1, and pronotal lobe; dorsal aspect with bands of setae very pale yellow, almost white, relatively narrow on metasoma (Fig. 10C). Clypeus convex in profile, lacking midline and larger punctures, asetose or sparsely covered with white setae (especially males). Mesepisternum lacking erect, simple setae or with sparse, short, erect, simple setae, with punctures nearly contiguous, but separated by up to one or two puncture diameters in a few places ventrally, these areas somewhat elevated, weakly tuberculate in appearance; with dense, white, branched setae on dorsal third, anterior margin, and



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anterior surface of mesepisternum, ventrally mostly asetose. Paramedian bands distinct (some females) or joined laterally to diffuse pale setae on anterior margin of scutum (some females and all males). Scutellum with moderate biconvexities; axillar spines triangular; sometimes rounded apically, almost reaching or reaching midpoint of scutellum (Fig. 10F). T1 interspace widely ovate; T2 with lateral bands forming acute angle with apical transverse band of pale setae (Fig. 10C). Female: Pseudopygidial area strongly circular (Fig. 10A); S5 strongly downcurved (Fig. 10B); S2–S4 with white apical bands of setae. Male: Pygidial plate narrow, lateral margins nearly parallelsided, lacking or with weak basal transverse ridge; S3–S4 with brown apical fringes (slightly white laterally on S4); S2–S3 with white apical bands of setae (medially extending past apical margin of S3).

FIGURE 10. Triepeolus eliseae Rightmyer, new species. Female from Michigan A) pseudopygidial area. B) S5 strongly downturned. C) Dorsal habitus. D) Mesepisternum in lateral view. E) Face. F) Oblique dorsal view of mesosoma.


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FIGURE 11. Triepeolus eliseae Rightmyer, new species. Female from Grand Rapids, Michigan photographed by Kim Phillips, 23 Jul. 2014. A) Oblique lateral view. B) Oblique posterior dorsal view. Photographs used with permission.

Comments. This species was referred to as Triepeolus n. sp. 1 in the key to males and females of the United States east of the Mississippi River and Eastern Canada (Rightmyer 2008: 26); it is also known as Triepeolus sp. 101 in the online guide Discover Life (http://discoverlife.org). JSA has examined additional specimens of what he considers to be this species from Maritime Canada (S. Javorek collection) and has identified images taken in New Brunswick as this taxon under the name “Triepeolus new species in the verbesinae group”: http://bugguide.net/ node/view/826110/bgimage. An additional Michigan record was photographed in Grand Rapids, Michigan (Fig. 11) (see also: http://bugguide.net/node/view/967055/bgimage). Distribution. CANADA: Ontario; USA: Indiana, Maryland, Michigan, Minnesota, New Jersey, and Wisconsin. Floral Records. Heliopsis sp., Solidago sp., Symphyotrichum sp. Seasonal Records. June 27 to August 27. Holotype.—"17 mi. SE Kenora, Ontario CAN. [Canada] VIII-7-1979 J. R. Powers, Collr. // M. G. Rightmyer Triepeolus Database Specimen No. 1190 // Holotype. ♀ Triepeolus eliseae Rightmyer" (CAS). Paratypes. CANADA: ONTARIO: 15 mi. SE Kenora, 4 Aug. 1979, J. R. Powers (1♀ CAS), 17 mi. SE Kenora, 7 Aug. 1979, J. R. Powers (1♀ CAS); USA: INDIANA: Jasper Co.: Prairie Border, 25 Jul. 2003, Heliopsis sp., R. P. Jean (1♀ RJPC); MARYLAND: Prince George's Co.: North Tract Patuxent, 27–28 Jun. 2012, S. Droege lab, pan trap (1♀ NMNH); MICHIGAN: Alger Co.: (no locality), 31 Aug. 1946, R.R. Dreisbach (1 ♀ MSUC); Dickinson Co.: T42N R29W Sec. 13, 14 Aug. 1983, R.L. Fischer (1 ♀ MSUC); T43N R29W Sec. 14, 18 Aug. 1983, Solidago, A. R. Porter (1 ♀ MSUC); T43N R29W Sec. 14, 16 Aug. 1983, Aster, A. R. Porter (1 ♀ MSUC); Kalkaska Co.: (no locality), 21 Jul. 1966, window pane trap, L. F. Wilson (2 ♀ MSUC); Leelanau Co.: 5 Jul. 1941, R.R. Dreisbach (1♂ NMNH); Missaukee Co.: Lake City, 3 Aug. 1948, D. Bray (1 ♀ MSUC). MINNESOTA: [St. Louis Co.]: Eagles Nest, 10 Aug. 1959, W. V. Balduf (1♀ INHS); NEW JERSEY: Union Co.: Westfield, 1956, C. Ferguson (1♀ OSAC), 8 Jul. 1956, C. Ferguson (1♀ BBSL); WISCONSIN: Bayfield Co.: Salmo, 12 Aug. 1919, Solidago, W. L. McAtee (1♂ NMNH), Douglas Co.: Wascott, 21 Jul. 1910, Timberlake (1♂ NMNH). Etymology. This species is named in honor of Mary Elise G. Y. Gee, daughter of M. G. Rightmyer. County records: Alger, Dickinson, Leelanau, Kalkaska, Kent, Missaukee.



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Triepeolus helianthi (Robertson 1897) (New state record) County records: Wayne. Material examined. Wayne Co.: Detroit, 14 Aug. 1929, G. Steyskal (1 ♀ MSUC).

Triepeolus lunatus (Say 1824) County records: Ingham, Ottawa, Shiawassee, Washtenaw, Wayne. Notes. Likely a cleptoparasite of Melissodes bimaculatus (see Rightmyer 2008).

Triepeolus michiganensis Mitchell 1962 (Michigan holotype; Fig. 12A) Triepeolus michiganensis Mitchell, 1962: 473. Holotype. ♀ USA: Michigan: Shiawassee Co.: 3 Sep.1950, R.R. Dreisbach (NMNH: 75244)

County records: Shiawassee. Notes. This cleptoparasite is quite rare. JSA observed this species at a nest site of a Melissodes (Eumelissodes) in Ithaca, New York (Rightmyer 2008). The host record was originally documented as M. denticulatus, but the identification needs to be rechecked as it more likely pertains to M. subillatus. Both Melissodes species are relatively common in southern Michigan.

Triepeolus pectoralis (Robertson 1897) County records: Berrien, Cheboygan, Gogebic, Huron, Ingham. Notes. Cleptoparasite of Melissodes druriellus (see Rightmyer 2008).

Triepeolus remigatus (Fabricius 1804) County records: Berrien, Ingham, Shiawassee. Notes. Cleptoparasite of Peponapis pruinosa.

Triepeolus simplex Robertson 1903 (simplex group) County records: Kalamazoo, Kalkaska, Leelanau, Livingston, Saginaw.

Nomadini Genus Nomada Scopoli Taxonomy: Alexander & Schwarz (1994); Broemeling & Moalif (1988); Droege et al. (2010); Evans (1972); Mitchell (1962); Schwarz & Gusenleitner (2004). Nomada is in serious need of revision, especially the species-rich ruficornis group, which may be paraphyletic and includes a particularly challenging subgroup of species with bidentate mandibles. Nomada is the most problematic genus for the state and the following list is likely to be changed substantially following taxonomic revision of the genus. Biology. All Nomada are cleptoparasites. Hosts are primarily within the genus Andrena (the known or


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suspected host of species in the ruficornis and vincta groups), but bees in other families are also be attacked including halictid (e.g., Agapostemon by species in the erigeronis and vegana groups) and apid (e.g., Eucera by species in the superba group) bees.

Nomada alpha alpha Cockerell 1905 (ruficornis group) (New state record) (not N. sphaerogaster var. α Cockerell, 1905b, see below) County records: Iron, Marquette. Notes. This species described from Fort Collins, Larimer Co., Colorado and its two valid subspecies N. alpha dialpha Cockerell 1921 and N. alpha paralpha Cockerell 1921 (both from Walden, Colorado) are currently only known from that state (Cockerell 1905a, 1921; Swenk 1912; Hurd 1979). The close proximity of the type localities (within approximately 100 km) for all the subspecies, including two from near Walden, Jackson Co., Colorado, does not fit the standard concept of subspecies as being geographically separated populations. There seems to be a close affinity with N. obliterata Cresson, although that species is unusual for having only two submarginal cells and differs from N. alpha in several respects, such as clypeal sculpture, hind tibial setae, and color. As is the case for many Nomada species as currently recognized, N. alpha and its putative subspecies requires additional study. Note that N. alpha was described by Cockerell (1905a) with the epithet alpha spelled in letters whereas the infrasubspecific name of N. sphaerogaster described by Cockerell (1905b) in that same year from Wisconsin was designated originally with a symbol α. We regard the former as valid and the latter as an unavailable name (see N. sphaerogaster below). Material examined. Iron Co.: T44N R131W Sec. 13, 1 Jun. 1983, M. Arduser (1 ♀ MSUC); Marquette Co.: Huron Mountain Club, 19 Jun. 1983, D.C.L. Gosling (1 ♀ UMMZ).

Nomada armatella Cockerell 1903 (ruficornis group) County records: Kent, Livingston, Midland, Muskegon, Ottawa, Van Buren. Notes. Cleptoparasite of Andrena regularis and A. vicina (Miliczky & Osgood 1995). Additional material. Ag. Coll. Mich., C.F.B., Collection C.F. Baker (1 ♀ NMNH) (S. Droege, pers. comm.).

Nomada articulata Smith 1854 (erigeronis group) County records: Allegan, Arenac, Barry, Bay, Berrien, Calhoun, Cheboygan, Clinton, Eaton, Hillsdale, Ingham, Ionia, Isabella, Kalamazoo, Lake, Lapeer, Leelanau, Lenawee, Livingston, Oceana, Ottawa, Sanilac, Shiawassee, Tuscola, Van Buren, Washtenaw. Notes. Cleptoparasite of Agapostemon virescens and A. sericeus (Abrams & Eickwort 1980, 1981).

Nomada australis Mitchell 1962 (erigeronis group) County records: Barry, Ingham, Kent, Ottawa.

Nomada autumnalis Mitchell 1962 (ruficornis group) (Michigan holotype; Fig. 12B) Nomada (?Heminomada) autumnalis Mitchell, 1962: 382. Holotype. ♂ USA: Michigan: Oakland Co.: 23 Sep. 1923, A.W. Andrews (NMNH: 75212).

County records: Montmorency, Oakland. Notes. Species delimitation and identification of bees within the genus Nomada are notoriously challenging



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and the ruficornis group, which may be paraphyletic (Alexander 1994) and includes the majority of eastern species (Alexander & Schwarz 1994), is in need of revision. The taxonomy of this species, which is nominally endemic to Michigan, remains in doubt.

Nomada banksi Cockerell 1907 (ruficornis group) County records: Gladwin, Isabella, Menominee, Midland. Notes. Possibly a cleptoparasite of Nomada asteris (see Ascher et al., 2014).

Nomada bella Cresson 1863 (ruficornis group: bidentate mandible) County records: Jackson, Lapeer, Monroe, Ottawa.

Nomada besseyi Swenk 1913 (vincta group) County records: Kalamazoo. Notes. According to S. Droege, N. besseyi may be a junior synonym of N. graenicheri (Cockerell 1905b), a species known from relatively few specimens from Wisconsin and Ontario (Wolf & Ascher 2009) and recently reported from Minnesota (S. Droege, pers. comm.). Specimens from Michigan identified as N. besseyi in a revision (Broemeling & Moalif 1988) closely match images of the N. graenicheri holotype deposited at the NMNH. Broemeling & Moalif (1988) did not treat N. graenicheri in their revision of this group (as subgenus Pachynomada) and it was misplaced in the ruficornis species group by Alexander & Schwarz (1994). Members of this species group are cleptoparasites of Andrena (Callandrena s. l.) (Broemeling & Moalif 1988) Material examined. Kalamazoo Co.: Gull Lake Biol. Sta. (KBS), 6 Aug. 1959, R.L. Fischer (1 ♀ 1 ♂ MSUC): Gull Lake Biol. Sta. (KBS), 14 Aug.1965, R.L. Fischer (2 ♀ 1 ♂ MSUC). Additional material. Kalamazoo Co.: Gull Lake Biol. Sta. (KBS), 27 Jul. 1963, G.C. Eickwort (BBSL): Gull Lake Biol. Sta. (KBS), 12 Aug.1965, R.L. Fischer (BBSL).

Nomada bethunei Cockerell 1903 (ruficornis group) County records: Kalamazoo, Midland, Wayne.

Nomada binotata (Robertson 1903) (ruficornis group: bidentate mandible) (New status, new state record) Gnathias ovatus form binotatus Robertson 1903: 175 [female lectotype, INHS]

County records: Allegan, Dickinson, Lake, Livingston, Macomb, Midland, Montcalm, Montmorency, Newaygo, Oakland, Ottawa, Wexford. Notes. This species was originally described as Gnathias ovatus form binotatus. The name has been infrequently cited as a synonym of the nominal species (Alexander & Schwarz 1994; Hurd 1979; Mitchell 1962; Rodeck 1951). After examining lectotypes at INHS for this and the nominal species, JG considers that N. binotata deserves to be treated as distinct from N. ovata. Nomada binotata has a distinctive dense patch of 12 or more modified setae on the metatibial apex, but N. ovata has only four. A morphospecies recorded by Goldstein & Ascher (2016) from Massachusetts as “Nomada [Gnathias] sp. ["multispine"]” may prove to correspond with N. binotata, in which case it is likely widespread across northeastern North America. To clarify the status of this and other morphospecies in this species group, i.e. Gnathias sensu Mitchell (1962), revisionary study is required, to clarify, e.g., the status of the four “forms” of N. ovata described by Robertson (1903) [as forms of Gnathias ovatus


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with masculine specific epithets], octomaculata, plena, sexmaculata, and unicolor, in addition to the two reinstated here. Material examined. Allegan Co.: (no locality), 18 May 1959, R. & K. Dreisbach (1 ♀ MSUC); Dickinson Co.: Channing, T43N R30W sec. 18, 10 Jun. 1983, Malaise trap, M. Arduser (1 ♀ MSUC): Channing, T43N R30W sec. 18, 12 Jun. 1983, Malaise trap, M. Arduser (1 ♀ MSUC): Lake Co.: (no locality), 6 May 1945, R.R. Dreisbach (1 ♂ MSUC); Big Star Lake, 21–27 May 1975, Malaise trap, T.A. Bowling (1 ♀ MSUC); Big Star Lake, 8–10 Jun 1975, Malaise trap, T.A. Bowling (1 ♀ MSUC); Big Star Lake, 21–24 Jun 1974, Malaise trap, T.A. Bowling (1 ♀ MSUC); Big Star Lake, 29 Jun-1 Jul. 1974, Malaise trap, T.A. Bowling (1 ♀ MSUC); Branch, 28 May-4 Jun. 1975, Malaise trap, T.A. Bowling (1 ♀ MSUC); Branch, 15–17 Jun. 1975, Malaise trap, T.A. Bowling (1 ♀ MSUC); Livingston Co.: (no locality), 6 Jun. 1943, R.R. Dreisbach (1 ♀ MSUC); Macomb Co.: 9 mile Rd. & Woodard Ave., 10 May 1914, W.S. McAlpine (1 ♀ MSUC); Midland Co.: (no locality), R.R. Dreisbach, 12–21 Jun. 1961 (1 ♀ MSUC); Montcalm Co.: Flat River Game Area, 14 May 1955, R.W. Hodges (2 ♀ MSUC); Montmorency Co.: (no locality), 1–8 Jun. 1966, window pane trap, P.C. Kennedy (1 ♀ MSUC); (no locality), 15– 22 Jun. 1966, window pane trap, P.C. Kennedy (1 ♀ MSUC); Atlanta, 23 May 1957, R.L. Fischer (1 ♀ MSUC); Newaygo Co.: 5 mi. N Newaygo, 22–30 May 1975, Malaise trap, T.A. Bowling (1 ♀ MSUC); Ottawa Co.: 1 mi. W Harlem, 25 May 2005, J.K. Tuell et al. (1 ♀ MSUC:RI); Roscommon Co.: (no locality), 21 May 1950, R.R. Dreisbach (1 ♀ MSUC): Wexford Co.: Cadillac, 11 Jun. 1983, R.L. Fischer (1 ♀ MSUC); T24N R9W, sec. 16, 15 Jun. 1965, J.H. Shaddy (3 ♀ MSUC); T24N R9W, sec. 16, 23–30 Jun. 1965, J.H. Shaddy (3 ♀ MSUC); T24N R9W, sec. 22, 15 Jun. 1965, J.H. Shaddy (7 ♀ MSUC); T24N R9W, sec. 22, 23–30 Jun. 1965, J.H. Shaddy (7 ♀ MSUC).

Nomada composita Mitchell 1962 (ruficornis group) County records: Benzie, Chippewa, Grand Traverse, Lake, Ottawa, Presque Isle, Schoolcraft, Van Buren.

Nomada cressonii Robertson 1893 (ruficornis group) County records: Allegan, Antrim, Kent, Leelanau, Livingston, Missaukee, Oakland, Ottawa, Van Buren, Washtenaw, Wexford. Notes. Cleptoparasite of Andrena crataegi, A. vicina and A. regularis (Miliczky & Osgood 1995; Osgood 1989).

Nomada cuneata (Robertson 1903) (ruficornis group: bidentate mandible) County records: Allegan, Antrim, Berrien, Cass, Charlevoix, Delta, Gogebic, Isabella, Kalkaska, Mackinac, Marquette, Ontonagon, Ottawa, Van Buren. Notes. A cleptoparasite of Andrena vicina (Miliczky & Osgood 1995). Nomada cuneata is a large dark red bee with two long modified setae on the hind tibial apex. One is long and white and the second is thick and red. This is reminiscent of putative N. bella and N. maculata, but the setae are much closer to each other. Additional study is needed of this group to clarify species boundaries and Michigan records. Rodeck (1931, 1951) treated this name as a subspecies of N. lepida (see entry below).Robertson (1903) described four forms of Gnathias cuneatus: decemnotatus, octonotatus, sexnotatus and quadrisignatus, that have been treated as junior synonyms (Alexander & Schwarz 1994; Rodeck 1951) but that warrant further scrutiny. Material examined. Van Buren Co.: 1 km SW Breedsville, 42.3406 -86.0819, 16 May 2016, Barbarea vulgaris, J. Gibbs, K. Gundersen (1 ♀ MSUC:RI); 2.5 km NW Breedsville, 42.3576 -86.0982, 16 May 2015, Cerastium vulgatum, J. Brokaw, S. Way (1 ♀ MSUC:RI); 3.5 km NW Gobles, 42.3824 -85.9147, 18 May 2015, Cerastium vulgatum (1 ♀ MSUC:RI); 4.5 km W Grand Junction, 42.4006 -86.126, 22 May 2015, Barbarea vulgaris, J. Gibbs (1 ♀ MSUC:RI).



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Nomada decepta Mitchell 1962 (ruficornis group) County records: Midland.

Nomada dentariae (Robertson 1903) (ruficornis group) County records: Clare. Notes. Recorded by Mitchell (1962). The veracity of this record is uncertain.

Nomada denticulata Robertson 1902 (ruficornis group) County records: Allegan, Barry, Ingham, Monroe, Van Buren, Washtenaw.

Nomada depressa Cresson 1863 (ruficornis group) County records: Jackson, Shiawassee, Washtenaw.

FIGURE 12. Michigan holotype material at the National Museum of Natural History, Smithisonian Institution (not to scale). A) Triepeolus michiganensis Mitchell, dorsal habitus. B) Nomada (Xanthidium) autumnalis Mitchell, dorsal habitus. C) Nomada dreisbachi Mitchell, dorsal habiuts. D). Nomada sobrina Mitchell, dorsal habitus. Photographs provided with the permission of the National Museum of Natural History, Smithsonian Institution, 10th and Constitution Ave. N.W., Washington, DC 20560-0193. (http://www.nmnh.si.edu/).


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Nomada dreisbachi Mitchell 1962 (ruficornis group) (Michigan holotype; Fig. 12C) Nomada dreisbachi Mitchell, 1962: 414. Holotype. ♂ USA: Michigan: Newaygo Co.: 13 May 1956, R. & K. Dreisbach (NMNH: 75218).

County records: Kent, Midland, Newaygo, Osceola. Notes. This uncommon nominal species described from the male only is very similar to the common N. cressonii, differing in the more shallow medial impression of the mesoscutellum (Mitchell 1962). Nomada dreisbachi may be a junior synonym of N. cressonii. Material examined. Midland Co.: (no locality) 4 May 1945, R.R. Dreisbach (1 ♂ paratype MSUC); Osceola Co.: (no locality) 3 May 1952, R.R. Dreisbach (1 ♂ paratype MSUC).

Nomada erigeronis Robertson 1897 (erigeronis group) (New state record) County records: Ottawa. Notes. The size and sculpturing of the male specimens examined are consistent with the female lectotype (INHS). Material examined. Ottawa Co.: Marne, 2 km S. 43.0172 -85.8381, 2 Jul. 2015 on Rudbeckia hirta, J. Brokaw (2 ♂ MSUC: RI).

Nomada fervida Smith 1854 (vegana group) County records: Benzie, Berrien, Huron, Livingston, Midland, Newaygo.

Nomada gracilis Cresson 1863 (ruficornis group) County records: Chippewa.

Nomada illinoensis Robertson 1900 (ruficornis group) County records: Antrim, Missaukee, Oakland, Ottawa, Van Buren, Washtenaw.

Nomada imbricata Smith 1854 (ruficornis group) County records: Berrien, Calhoun, Clinton, Ingham, Iron, Kent, Livingston, Midland, Newaygo, Oceana, Van Buren, Washtenaw. Notes. Cleptoparasite of Andrena regularis (Miliczky & Osgood 1995). Nomada imbricata has been observed at nest sites of A. dunningi in Winnipeg, Manitoba (JG, pers. obs.) and at Ithaca, New York (JSA, pers. obs.).

Nomada lehighensis Cockerell 1903 (ruficornis group) County records: Marquette.



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Nomada lepida Cresson 1863 (ruficornis group: bidentate mandible) County records: Lake, Livingston, Manistee, St. Clair, Van Buren, Washtenaw, Wexford. Notes. Scott et al. (2011) noted that type material of this species from Colorado and Illinois may not be conspecific and could not confirm presence in Colorado of material matching N. lepida from the eastern United States as described by Mitchell (1962) and identified by that author. They therefore removed this species from the fully confirmed Colorado state list. However, the male lectotype in the ANSP designated by Cresson (1916) was collected in Pike’s Peak, Colorado, making that occurrence correct by definition. The type series of lepida may prove to have been composite, and this would explain failure by Scott et al. (2011) to confirm non-type lepida from Colorado that match this species sensu Mitchell (1962) and as identified by subsequent bee specialists working on the eastern fauna. If lepida sensu Mitchell (1962) do diverge from the Colorado lectotype, then populations from the eastern United States, including Michigan, may require description as a new species unless another name is found to be applicable.

Nomada luteoloides Robertson 1895 (ruficornis group) = Nomada (Heminomada) colorata Mitchell, 1962: 386. (Michigan holotype) Holotype. ♀ USA: Michigan: Sanilac Co.: 7 Aug. 1926 (NMNH). Synonymy by Schwarz & Gusenleitner (2004).

County records: Allegan, Antrim, Calhoun, Clinton, Dickinson, Eaton, Ingham, Iron, Kalamazoo, Kent, Leelanau, Lenawee, Livingston, Midland, Oakland, Sanilac, St. Clair, Van Buren, Washtenaw. Notes. Likely a cleptoparasite of Andrena carlini (Goldstein & Ascher 2016). See Table 1 and above.

Nomada maculata Cresson 1863 (ruficornis group: bidentate mandible) County records: Alger, Allegan, Antrim, Clinton, Grand Traverse, Ingham, Kalamazoo, Kent, Leelanau, Livingston, Montmorency, Oakland, Ottawa, Roscommon, St. Clair, Van Buren, Washtenaw, Wayne.

Nomada obliterata Cresson 1863 (ruficornis group) County records: Allegan, Clinton, Kalamazoo, Oakland, Washtenaw, Wayne. Notes. Miliczky (1988) and JSA at Ithaca, New York (pers. obs.) record this as a cleptoparasite of Andrena erythrogaster.

Nomada ochlerata Mitchell 1962 (ruficornis group) Michigan holotype) Nomada (?Heminomada) ochlerata Mitchell, 1962: 396. Holotype. ♂ USA: Michigan: Washtenaw Co.: Ann Arbor, 9 May 1935, I.J. Cantrall (UMMZ).

County records: Washtenaw. Notes. The taxonomy and identification of N. ochlerata requires additional study. This species, described from the male only, may prove to be associated with one described from females only, such as N. xanthura Cockerell (see also other poorly understood brown-spined forms such as N. mendica Mitchell, described from the male, and N. composita described from the female; these should all be compared with N. gracilis Cresson, now known from both sexes following association of N. inepta Mitchell as a synonym).

Nomada ovata (Robertson 1903) (ruficornis group: bidentate mandible) County records: Arenac, Berrien, Dickinson, Gogebic, Iron, Livingston, Mackinac, Ontonagon, Ottawa, Van Buren.


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Nomada parva Robertson 1900 (ruficornis group) County records: Ogemaw, Washtenaw.

Nomada perplexa Cresson 1863 (ruficornis group: bidentate mandible) County records: Allegan, Bay, Mackinac, Manistee, Midland, Oakland, Van Buren, Washtenaw.

Nomada placida Cresson 1863 (roberjeotiana group) (Fig. 13) County records: Berrien, Van Buren, Washtenaw. Notes. Rodeck (1949) in his revision of the subgenus Callinomada documented this species from the midAtlantic and Midwest, including records from Carlinville, Illinois and Lafayette, Indiana. Mitchell (1962) subsequently recorded N. placida from Michigan. This late summer-flying Nomada has been recently collected in the state. Hosts should be autumnal, composite-specialist Andrena. Material examined. Berrien Co.: SWMREC, 21 Aug. 2015, Solidago juncea (2 ♂ MSUC: RI); SWMREC, 27 Aug. 2015, Oligoneuron rigidum (1 ♂ MSUC: RI); SWMREC, 1 Sep. 2015, Oligoneuron rigidum (1 ♂ MSUC: RI); Tuscola Co.: (no locality), 4 Aug. 1946, R.R. Dreisbach (1 ♀ MSUC); Washtenaw Co.: Lodi Twsp., 21 Aug. 2016 (http://bugguide.net/node/view/1279545).

FIGURE 13. Nomada placida Cresson female from Michigan, lateral habitus.



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Nomada pygmaea Cresson 1863 (ruficornis group) County records: Allegan, Antrim, Ingham, Keweenaw, Livingston, Ottawa, Van Buren. Notes. Cleptoparasite of Andrena regularis (Miliczky & Osgood 1995).

Nomada quadrimaculata (Robertson 1903) (ruficornis group: bidentate mandible) (New status, new state record) Gnathias ovatus var. quadrimaculatus Robertson 1903: 175 [female lectotype, INHS].

County records: Alpena, Antrim, Clare, Gladwin, Oakland, Van Buren. Notes. This species was originally described as Gnathias ovatus var. quadrimaculatus, but has not been in common usage, having been typically synonymized with the nominal species. JG makes this tentative determination based on examination of the lectotype. It differs from N. ovata in part by the more numerous five or six specialized setae on the apex of the hind tibia, but it is possible this character varies and is not reliable for species delimitation. Other putative forms of N. ovata, described by Robertson and listed above, differ in the number of these setae, in addition to their varying color patterns. Material examined. Alpena Co.: (no locality), 2 Jun. 1951, R.R. Dreisbach (1 ♀ MSUC): Antrim Co.: Glacial Hills, Vandermark entrance, 44.987 -85.2417, 12 Jun. 2016, Rubus, K. Odanata, J. Brokaw, S. Naranjo (1 ♀ MSUC); Glacial Hills, Vandermark entrance, 44.987 -85.2417, 12 Jun. 2016, J. Gibbs (1 ♀ JBWM); Glacial Hills, Eckhardt entrance, 44.9906 -85.292, 12 Jun. 2016, Rubus, K. Odanata, J. Brokaw, S. Naranjo (2 ♀ JBWM); Clare Co.: (no locality), 20 Jun. 1950, R.R. Dreisbach (1 ♀ MSUC); Clinton Co.: Bath, 19 May 1956, R.L. Fischer (1 ♀ MSUC): Gladwin Co.: (no locality), 10–16 Jun. 1951, R.R. Dreisbach (1 ♀ MSUC); Mackinac Co.: (no locality), 7 Jun. 1960, R. & K. Dreisbach (1 ♀ MSUC): Oakland Co.: (no locality), 12 Aug. 1923 [illegible] (1 ♀ MSUC); Van Buren Co.: 3 mi. N Friendsville, 21 May 2005, J.K. Tuell et al. (1 ♀ MSUC: RI).

Nomada rubicunda Olivier 1811 (erigeronis group) County records: Barry, Chippewa, Midland, Newaygo, Ottawa. Notes. The localized distribution of this bee to sandy sites suggests an association with Agapostemon splendens (Lepeletier) (JSA, pers. obs.).

Nomada sayi Robertson 1893 (ruficornis group) County records: Barry, Livingston, Van Buren.

Nomada sobrina Mitchell 1962 (ruficornis group) Nomada sobrina Mitchell, 1962: 431. (Michigan holotype; Fig. 12D) Holotype. ♂ USA: Michigan: Macomb Co.: 28 May 1949, R.R. Dreisbach (NMNH: 75225).

County records: Macomb. Notes. The taxonomy and identification of this species requires additional study. The extensive red on the mesoscutum and yellow markings on the mesoscutellum and metasoma are reminiscent of Midwestern N. bethunei Cockerell (Droege et al. 2010).


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Nomada sphaerogaster Cockerell 1903 (ruficornis group) (New state record) (Fig. 14) =Nomada sphaerogaster var. α Cockerell 1905b (not N. alpha Cockerell 1905a)

FIGURE 14. Nomada sphaerogaster Cockerell female from Michigan. A) Dorsal habitus. B) Face. C) Lateral habitus. D) Hind tibia showing apical setae.

County records: Allegan, Clinton, Ingham, Van Buren. Notes. This poorly known species was described from a female collected Apr. 22 in Riverton, Burlington Co., New Jersey (Cockerell 1903a), to the west of the New Jersey Pine Barrens. Cockerell (1905b) subsequently described var. α from Milwaukee, Wisconsin (NMNH), although he evidently did not intend it to be a specific epithet since he appended to the descripton of the variety: “If the difference should prove constant in a long series, it ought to have a distinct name”. To our knowledge, it has not been used as a valid species group name and is therefore unvailable (ICZN art. 45.6.4). The variety α was excluded from catalogues by Hurd (1979), Alexander & Schwarz (1994), and Zuparko (2017). Furthermore, if it were valid it would be a primary homonym of N. alpha Cockerell, 1905a (see above). Nomada sphaerogaster has not been recognized subsequently from Wisconsin (Wolf & Ascher, 2009) or from elsewhere in the Midwest. Michigan specimens were collected at the edge of a clearing in a moderately damp woodland, with sandy soil, surrounded by suburban development. The Michigan specimens are quite distinctive among vernal Nomada as both sexes are black with complete yellow bands on T2-T5 (Fig. 14A), in combination with wide heads with distinct darkened facial setae (Fig. 14B). The Michigan specimen closely matches non-type specimens of N. sphaerogaster from New Jersey in the Museum of Comparative Zoology (S. Droege, det.). The holotype apparently differs from the Michigan specimens in having bands on T3 and T4



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narrowly interrupted. Cockerell (1905b) states in the description of sphaerogaster var. α that the bands are “narrowed or slightly interrupted in the middle”. The female has distinctive dark setae on the face and distinctly separated dark setae on the apex of the metatibia. Other spring Nomada, such as N. luteoloides, are superficially similar in the complete banding on the metasoma but differ in having a narrower head, lacking dark facial setae, and the apical setae of the metatibia are thicker and tightly compressed. Nomada sphaerogaster been collected during mid-April in central and southwest Michigan. Material examined. Allegan Co.: 0.8 mi. N Fillmore, blueberry field, 16 Apr. 2005, J. Tuell et al. (1 ♀ MSUC: RI); 1.3 mi. NNE Glenn, blueberry field, 24 Apr. 2006, Tuell et al. (1 ♀ MSUC: RI); Clinton Co.: Bath, 6 Apr. 1963, G.C. Eickwort (1 ♂ MSUC); Ingham Co.: Harrison Meadows/Abbot Road Park, East Lansing, N42.7645 W84.7945, 19 Apr. 2014, on Salix, J. Gibbs (1 ♀ JBWM); Harrison Meadows/Abbot Road Park, East Lansing, N42.7645 W84.7945, 20 Apr. 2014, on Salix, J. Gibbs (1 ♂ JBWM); Harrison Meadows Park/Abbot Road, East Lansing, N42.7645 W84.7945, 60 Apr. 2014, on Salix, J. Gibbs (1 ♂ JBWM), Van Buren Co.: 2.6 mi. ESE Breedsville, blueberry field, 16 Apr. 2005, J. Tuell et al. (1 ♀ MSUC: RI). Additional Material examined. New Jersey: Morris Co.: Great Swamp National Wildlife Refuge, (no date), J. Bequaert (1 ♀ Museum of Comparative Zoology, photographs).

Nomada superba Cresson 1863 (superba group) (New state record) (Fig. 15) County records: Allegan, Berrien, Kalamazoo. Notes. Newly recorded from Michigan based on two females. Mitchell (1962) records this species from Illinois, Indiana and Ohio and Evans (1972) records it from Wisconsin. Wolf & Ascher (2009) list Eucera (Synhalonia) as the host, which is consistent with other members of the superba group (Alexander 1991). Material examined. Allegan Co.: Dorr, 20 May 2010 (http://bugguide.net/node/view/1024536); Berrien Co.: SWMREC, 24 Jun. 2015, Lotus corniculatus (1 ♀ MSUC: RI); Kalamazoo Co.: Gull Lake Biol. Sta. (KBS), 20 Jun. 1968, ex. rotary trap set in field of oats, W.G. Ruesink (1 ♀ MSUC).

FIGURE 15. Nomada superba Cresson female from Michigan, lateral habitus.


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Nomada tiftonensis Cockerell 1903 (vegana group) County records: Barry, Dickinson, Ionia, Kalamazoo, Muskegon, Van Buren.

Nomada vegana Cockerell 1903 (vegana group) (New state record) County records: Allegan, Cass. Notes. Some taxonomic problems associated with members of the vegana species group were resolved by Droege et al. (2010). Nomada vegana was not treated in that paper, but Droege et al. (2010) provide useful information for comparison to other members of the group that occur in Michigan, including N. fervida (and its midwestern dark color form N. wisconsinensis Graenicher, previously considered as a distinct species, e.g., by Wolf & Ascher, 2009) and N. tiftonensis (treated as a junior synonym of N. heiligbrodtii Cresson by Mitchell, 1962 and Hurd, 1979, and also considered as a junior synonym of N. texana Cresson in earlier drafts of Ascher & Pickering, 2017). Material examined. Allegan Co.: 3 mi. ESE Pullman, N42.441 W86.082, 8–10 Jul. 2014, emergence trap (1 ♂ MSUC); Cass Co.: T5S R15W Sec. 23, 24 Jul. 1994, Malaise trap, W.P. Westrate (1 ♀ UMMZ); T5S R15W Sec. 23, 24 Jul. 1995, Malaise trap, W.P. Westrate (1 ♀ UMMZ); T5S R15W Sec. 23, 29 Jul. 1995, Malaise trap, W.P. Westrate (1 ♀ UMMZ); T5S R15W Sec. 23, 30 Jul. 1994, Malaise trap, W.P. Westrate (4♀ UMMZ); T5S R15W Sec. 23, 9 Aug. 1995, Malaise trap, W.P. Westrate (1 ♀ UMMZ).

Nomada vicina Cresson 1863 (ruficornis group) County records: Kalkaska, Livingston, Marquette, Washtenaw. Notes. Likely a cleptoparasite of Andrena hirticincta (see Ascher et al. 2014).

Nomada vincta Say 1837 (vincta group) County records: Barry, Kalamazoo, Livingston, Montcalm, Wayne.

Osirini Genus Epeoloides Giraud Taxonomy: Mitchell (1962).

Epeoloides pilosulus (Cresson 1878) County records: Berrien, Midland, Van Buren, Wayne. Notes. This is a cleptoparasite of multiple Macropis species likely including M. nuda (Cockerell) (Sheffield et al. 2004) and M. ciliata Patton (Wagner & Ascher 2008), which in turn are specialists on Lysimachia (Myrsinaceae, formerly Primulaceae) (Cane et al. 1983). Observations were lacking from its entire range for many decades (Michener 2000), but it has been rediscovered very locally in Canada (Sheffield et al. 2004) and in the United States in New England (Wagner & Ascher 2008) and New York (http://bugguide.net/node/view/954741). Of the four specimens examined from Michigan, none was collected in the last 70 years. Material examined. Berrien Co.: Paw Paw Lake, 9 Jul. 1906, E. Liljeblad (1 ♂ UMMZ); Midland Co.: (no locality), 23 Jul. 1928, R.R. Dreisbach (1 ♀ MSUC); (no locality), 24 Jun. 1944, R.R. Dreisbach (1 ♂ MSUC); Wayne Co.: Detroit, 18 Jul. 1937, G. Steyskal (1 ♀ MSUC).



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Xylocopinae Ceratinini Genus Ceratina Latreille Subgenus Zadontomerus Taxonomy: Mitchell (1962); Daly (1976); Rehan & Richards (2008); Rehan & Sheffield (2011).

Ceratina (Zadontomerus) calcarata Robertson 1900 County records: Alger, Allegan, Antrim, Barry, Bay, Berrien, Branch, Cass, Charlevoix, Cheboygan, Clinton, Eaton, Emmet, Gogebic, Grand Traverse, Gratiot, Huron, Ingham, Ionia, Iosco, Iron, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lake, Lapeer, Leelanau, Lenawee, Livingston, Macomb, Mason, Mecosta, Missaukee, Montcalm, Newaygo, Oakland, Oceana, Osceola, Ottawa, Saginaw, Sanilac, Shiawassee, St. Clair, Tuscola, Van Buren, Washtenaw, Wayne. Notes. Nesting biology and behavior described by Rehan & Richards (2010, 2013) and Kislow (1976) in Ontario and Georgia, respectively. This was the second most abundant wild bee netted from sour cherry flowers in commercial orchards (Gibbs et al. 2017).

Ceratina (Zadontomerus) dupla Say 1837 County records: Allegan, Barry, Bay, Berrien, Calhoun, Cass, Clinton, Eaton, Huron, Ingham, Jackson, Kalamazoo, Lake, Lapeer, Leelanau, Lenawee, Livingston, Mecosta, Newaygo, Ottawa, Saginaw, Van Buren, Washtenaw. Notes. Daly (1973) revised the Ceratina of America north of Mexico, which included examination of Michigan material. Due to problems with female identification, no doubt confounded by the presence of the then unknown C. mikmaqi Rehan & Sheffield, only males were identified in Daly’s revision. Specimens identified as C. dupla at MSUC were reexamined by JG. Of the 104 males identified by Daly as C. dupla only 12 were in fact that species. The remaining specimens were C. mikmaqi. Although most historical records of Ceratina dupla were misidentified, the species as currently definied is confirmed for 20 counties in the Lower Peninsula (see checklist entry below). Nesting in Ontario described by Vickruck et al. (2011).

Ceratina (Zadontomerus) mikmaqi Rehan & Sheffield 2011 County records: Allegan, Antrim, Barry, Berrien, Branch, Calhoun, Cass, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Delta, Dickinson, Eaton, Gratiot, Hillsdale, Huron, Ingham, Ionia, Jackson, Kalamazoo, Kalkaska, Lake, Lapeer, Leelanau, Lenawee, Marquette, Mecosta, Montcalm, Montmorency, Muskegon, Newaygo, Oakland, Oceana, Osceola, Ottawa, Roscommon, Saginaw, Shiawassee, St. Clair, St. Joseph, Van Buren, Washtenaw, Wayne, Wexford. Notes. Ceratina mikmaqi was described recently based on material from Nova Scotia, Ontario, Maryland, Nebraska, New York, Wisconsin and Kentucky (Rehan & Sheffield 2011). Since then it has proven to be a common and widely distributed species (Zarrillo et al. 2016). The first published record for Michigan came several years after the original description (Carson et al. 2016). Males of C. mikmaqi are very similar to C. dupla and would have been treated as this species in earlier works (e.g., Daly 1973, see above). Males differ from C. dupla by the sparse mesoscutal punctation, ecarinate hind tibia at ventral midlength, and wider T7 apical lamella. Females are similar to C. calcarata in the sparse mesoscutal punctation, but can be distinguished from that species by the sparse


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pubescence of the metasomal sterna. We also reconfirm the presence of C. calcarata and C. dupla in Michigan. The males of C. calcarata are distinctive, so Daly’s identifications of these remain valid. As Ceratina mikmaqi has proven to be so widespread and abundant, we do not list specific records, but a list of counties is presented below.Nesting in Ontario described by Vickruck et al. (2011).

Ceratina (Zadontomerus) strenua Smith 1879 County records: Allegan, Barry, Berrien, Calhoun, Cass, Clinton, Eaton, Huron, Ionia, Jackson, Kalamazoo, Lapeer, Lenawee, Menominee, Monroe, Montcalm, Muskegon, Ottawa, Saginaw, St. Clair, St. Joseph, Tuscola, Van Buren, Washtenaw, Wexford. Notes. Nest in Georgia described by Kislow (1976).

Xylocopini Genus Xylocopa Latreille Subgenus Xylocopoides Michener Taxonomy: Hurd (1961); Mitchell (1962) Behavior: Carpenter bees excavate nests in hardwood. Extra-limital species may use other substrates.

Xylocopa (Xylocopoides) virginica virginica (Linnaeus 1771) County records: Allegan, Barry, Berrien, Branch, Calhoun, Cass, Clare, Clinton, Eaton, Hillsdale, Ingham, Ionia, Jackson, Kalamazoo, Kent, Lake, Lapeer, Leelanau, Lenawee, Livingston, Macomb, Muskegon, Newaygo, Oakland, Oceana, Saginaw, Sanilac, Shiawassee, Tuscola, Van Buren, Washtenaw, Wayne. Notes. Carpenter bees are frequently found in man-made structures, but only occasionally are discovered in natural sites (Howard 1892). Xylocopa virginica has both single and multi-female nests. Social nests are unusual in that daughters may occupy nests without functioning as traditional workers (Richards 2011).

COLLETIDAE Colletinae Genus Colletes Latreille Taxonomy: Mitchell (1960); Stephen (1954). Behavior: Solitary ground-nesters known to line their nests with glandular secretions that harden into a cellophane-like material. Species may be floral specialists or generalists.

Colletes aberrans Cockerell 1897 (americanus group) County records: Berrien, Ingham, Washtenaw. Notes. A prairie associated species occurring widely in the Great Plains states, with the southern tier of Michigan counties appearing to be the eastern limit of its range. Material examined. Berrien Co.: Warren Dunes State Park, 16 Jul. 1982, M. & A. O’Brien (1 ♀ UMMZ); Washtenaw Co.: Pinckney State Recreation Area, 11 Jul. 1981, M. & A. O’Brien (1 ♀ UMMZ). Additional material. Ingham Co.: East Lansing, 29 Jul. 1947, R. Fischer (SEMC). THE BEES OF MICHIGAN


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Colletes aestivalis Patton 1879 (aestivalis group) (New state record) (Fig. 16A) County records: Monroe, Wayne. Notes. A male from Monroe Co. was identified by T.B. Mitchell, although he did not include Michigan in the species’ range in his 1960 revision. Stephen (1954) lists Carlinville, Illinois as the westernmost locality for C. aestivalis, whereas the closely related species C. andrewsi Cockerell occurs as far east as Wisconsin (Wolf & Ascher, 2009). Both are oligolectic on Heuchera spp. (Saxifragaceae). Identification of the two specimens from southeast Michigan reported below as aestivalis was confirmed by JG based on the relative length of the hind basitarsus (Stephen 1954) and reference to comparative material of that species from more eastern localities. There are very few recent records of C. aestivalis Patton and none from Michigan or, to our knowledge, from anywhere in the Northeastern United States. We therefore regard it as being of conservation concern. Material examined. Monroe Co.: (no locality), 12 Jun. 1957, R & K. Dreisbach (1 ♂ MSUC); Wayne Co.: (no locality) 30 May 1959, R & K. Dreisbach, (1 ♂ MSUC).

Colletes americanus Cresson 1868 (americanus group) County records: Alger, Allegan, Bay, Benzie, Cheboygan, Clare, Clinton, Crawford, Emmet, Gladwin, Grand Traverse, Huron, Ingham, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Leelanau, Livingston, Manistee, Mecosta, Midland, Montmorency, Oakland, Oscoda, Roscommon, Saginaw, St. Joseph, Van Buren, Wexford. Notes. Visits late-season Asteraceae.

Colletes banksi Swenk 1908 (hyalinus group) County records: Huron, Iosco, Lake, Leelanau. Notes. Specialist of Ilex (Aquifoliaceae). Michigan is at the northwestern edge of its range.

Colletes brevicornis Robertson 1897 (willistoni group) County records: Crawford, Midland, Oscoda. Notes. Recorded from many flowers, but Robertson records it as a specialist of Triodanis perfoliata (L.) Nieuwl. (as Specularia).

Colletes compactus compactus Cresson 1868 (compactus group) County records: Bay, Benzie, Cheboygan, Clinton, Huron, Ingham, Kalamazoo, Livingston, Manistee, Midland, Montcalm, Oakland, Van Buren, Washtenaw. Notes. Nests described by Rozen & Favreau (1968) in New York. Specialist on Symphyotrichum and Solidago (Asteraceae).

Colletes consors Cresson 1868 mesocopus Swenk 1907 (consors group) County records: Alcona, Alger, Cheboygan, Dickinson, Iosco, Kalkaska, Keweenaw, Otsego. Notes. A northern bee of limited occurrence in the eastern United States. The species is Holarctic, with the nominate form restricted to the Palearctic.


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Colletes hyalinus Provancher 1888 (hyalinus group) County records: Baraga, Charlevoix, Cheboygan, Chippewa, Clare, Dickinson, Iron, Kalkaska, Keweenaw, Lake, Mackinac, Manistee, Missaukee, Montmorency, Ontonagon, Osceola, Otsego, Presque Isle, Wexford.

Colletes impunctatus Nylander 1852 lacustris Swenk 1906 (impunctatus group) County records: Baraga, Cheboygan, Gladwin, Keweenaw, Marquette, Montmorency, Ontonagon.

Colletes inaequalis Say 1837 (inaequalis group) County records: Alger, Allegan, Barry, Benzie, Berrien, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Crawford, Dickinson, Emmet, Gladwin, Grand Traverse, Houghton, Ingham, Iosco, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Leelanau, Livingston, Luce, Mackinac, Manistee, Marquette, Midland, Missaukee, Montcalm, Montmorency, Newaygo, Oceana, Ogemaw, Osceola, Otsego, Ottawa, Roscommon, Schoolcraft, Shiawassee, Van Buren, Washtenaw, Wayne, Wexford. Notes. Nests described from New Jersey, Kansas and Maryland by Smith (1901; as compacta), Stephen (1954), and Batra (1980), respectively. Population structure and nest suitability studies in New York by LópezUribe et al. (2015). An abundant spring bee and important wild pollinator of spring crops including apple, blueberry and cherry (Gibbs et al. 2016, 2017).

Colletes kincaidii Cockerell 1898 (simulans group) County records: Alcona, Alpena, Antrim, Barry, Bay, Calhoun, Charlevoix, Cheboygan, Clare, Clinton, Delta, Dickinson, Eaton, Emmet, Grand Traverse, Gratiot, Huron, Ingham, Iosco, Iron, Kalamazoo, Kalkaska, Keweenaw, Lake, Leelanau, Lenawee, Livingston, Manistee, Mason, Mecosta, Menominee, Midland, Missaukee, Muskegon, Newaygo, Oakland, Oceana, Ontonagon, Otsego, Presque Isle, Roscommon, Saginaw, Van Buren, Wayne, Wexford. Notes. Nesting of Californian populations described by Torchio et al. (1988)

Colletes latitarsis Robertson 1891 (latitarsis group) County records: Allegan, Berrien, Cheboygan, Huron, Ingham, Ionia, Kalamazoo, Kalkaska, Livingston, Midland, Oakland, Otsego, St. Joseph, Van Buren, Washtenaw. Notes. Specialist on Physalis (Solanaceae).

Colletes mandibularis Smith 1853 (americanus group) County records: Leelanau.

Colletes nudus Robertson 1898 (nudus group) County records: Benzie, Cheboygan, Kalamazoo, Lake, Leelanau, Livingston, Manistee, Mason, Montmorency, Muskegon, Otsego, St. Joseph, Van Buren.



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Colletes simulans Cresson 1868 armatus Patton 1879 (simulans group) County records: Alcona, Allegan, Barry, Bay, Benzie, Cheboygan, Chippewa, Clare, Clinton, Crawford, Dickinson, Emmet, Gladwin, Gratiot, Hillsdale, Huron, Ingham, Iosco, Isabella, Jackson, Kalkaska, Kent, Keweenaw, Lake, Leelanau, Lenawee, Livingston, Luce, Mackinac, Macomb, Manistee, Marquette, Mason, Mecosta, Menominee, Midland, Missaukee, Montcalm, Oakland, Oceana, Ogemaw, Ontonagon, Osceola, Ottawa, Roscommon, Saginaw, Tuscola, Washtenaw, Wexford. Notes. Specialist on late-season Asteraceae.

Colletes solidaginis Swenk 1906 (americanus group) County records: Allegan, Arenac, Barry, Bay, Chippewa, Gladwin, Kalamazoo, Lake, Livingston, Mecosta, Midland, Montcalm, Muskegon, Newaygo, Oceana, Washtenaw. Notes. Specialist on Solidago (Wagner & Ascher 2014)

Colletes thoracicus Smith 1853 (inaequalis group) County records: Allegan, Berrien, Ingham, Muskegon, Ottawa, Van Buren. Notes. Nests described by Batra (1980), Cane (1991), and Parker & Böving (1924) from Maryland, New York, and Washington, D.C., respectively.

Colletes validus Cresson 1868 (inaequalis group) County records: Allegan, Barry, Berrien, Chippewa, Crawford, Emmet, Iosco, Jackson, Kent, Lake, Livingston, Mackinac, Marquette, Midland, Missaukee, Montcalm, Montmorency, Newaygo, Oscoda, Roscommon, Saginaw, Schoolcraft, Van Buren, Wexford. Notes. Due to its specialization on Vaccinium, this is a potentially valuable pollinator of blueberry. Nests described by Batra (1980) and Rajotte (1979) from Maryland and Connecticut, respectively.

Colletes willistoni Robertson 1891 (willistoni group) County records: Cheboygan, Iron, Isabella, Kalamazoo, Lake, Leelanau, Otsego. Notes. Physalis specialist.

Hylaeinae Genus Hylaeus Fabricius Taxonomy: Mitchell (1960); Snelling (1966b, 1968, 1970). Biology. Hylaeus nest in pre-existing cavities including plant stems and abandoned sweat bee nests (Barrows 1975; Krombein 1967; Scott 1994).

Subgenus Cephalylaeus Michener Revision: Snelling (1968).


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Hylaeus (Cephalylaeus) basalis (Smith 1853) County records: Alger, Cheboygan, Chippewa, Dickinson, Gogebic, Huron, Isabella, Keweenaw, Leelanau, Luce, Marquette, Mecosta, Midland, Missaukee, Oscoda, Sanilac, Schoolcraft, Wayne, Wexford. Notes. Biological information available from trap nests set out by Fye (1965) and Scott (1994, 1996) in Ontario and Michigan, respectively. There seem to be few recent records from the Northeastern United States, although the species is still routinely found further north and west.

Subgenus Hylaeus Fabricius s. s. Revision: Snelling (1970).

Hylaeus (Hylaeus) annulatus (Linnaeus 1758) (annulatus group) County records: Alger, Allegan, Bay, Berrien, Cass, Charlevoix, Chippewa, Clare, Delta, Dickinson, Emmet, Gladwin, Gogebic, Ingham, Iron, Jackson, Keweenaw, Marquette, Mecosta, Midland, Montmorency, Ontonagon, Oscoda, Ottawa, Roscommon, Saginaw, Sanilac, Schoolcraft, Tuscola, Van Buren, Washtenaw. Notes. Biological information, as Hylaeus ellipticus, available from trap nests set out in Ontario and Michigan by Fye (1965) and Scott (1994, 1996), respectively.

Hylaeus (Hylaeus) fedorica (Cockerell 1909) (mesillae group) County records: Baraga, Hillsdale, Ionia, Tuscola.

Hylaeus (Hylaeus) leptocephalus (Morawitz 1871["1870"]) (leptocephalus group) County records: Arenac, Gladwin, Huron, Ingham, Manistee, Midland, Montcalm, Saginaw, Washtenaw, Wayne, Wexford. Notes. Holarctic. Nesting biology in a Utah greenhouse study described by Torchio (1984) and occupancy in abandoned halictid ground nests by Barrows (1975), both as H. bisinuatus.

Hylaeus (Hylaeus) mesillae (Cockerell 1896) cressonii (Cockerell 1907) (mesillae group) County records: Allegan, Antrim, Arenac, Barry, Bay, Berrien, Branch, Calhoun, Cass, Charlevoix, Cheboygan, Clare, Clinton, Crawford, Dickinson, Gladwin, Grand Traverse, Gratiot, Huron, Ingham, Ionia, Iosco, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Leelanau, Livingston, Macomb, Manistee, Marquette, Mecosta, Midland, Montcalm, Montmorency, Muskegon, Newaygo, Oakland, Oceana, Otsego, Ottawa, Saginaw, Shiawassee, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Stem nesting recorded by Rau (1922) and Hicks (1926) from Missouri and Colorado, respectively.

Hylaeus (Hylaeus) rudbeckiae (Cockerell & Casad 1895) (mesillae group) (New state record) County records: Leelanau, Keweenaw. Notes. A single male specimen from the northern LP identified by Roy Snelling and tentatively confirmed by JG is deposited at MSUC. A female record from PWRC remains uncertain due to the difficulty in distinguishing this sex from H. mesillae but is plausible given the boreal habitat present there. Although rudbeckiae is abundant in



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the western United States, it has only rarely been recorded from states east of the Rocky Mountains, where expected only from near the Canadian border, but identification difficulties may obscure its true status. A Wisconsin record (Mitchell, 1960) was not accepted by Wolf & Ascher (2009) but should be reevaluated as it may prove valid. Mitchell’s records from Connecticut (also reported by Hurd, 1979) are less likely to be correct, and we consider a possible New Jersey record (Hurd, 1979) to be implausible given that confirmed records in eastern North America are from boreal sites. Material examined. Leelanau Co.: Glenn Haven, 24 Aug. 1960, G.C. Eickwort (1 ♂ MSUC). Additional record. Keweenaw Co.: Isle Royale National Park, Washington Island, 47.8763 -89.2326, 16 Aug. 2013, S. Wilson (1 ♀ PWRC).

Hylaeus (Hylaeus) saniculae (Robertson 1896) (mesillae group) County records: Alpena, Bay, Cheboygan, Dickinson, Iosco, Iron, Kalkaska, Marquette, Wexford. Notes. Across its range generally scarce in collections but may be underrecorded due to identification difficulties.

Hylaeus (Hylaeus) verticalis (Cresson 1869) (verticalis group) County records: Alcona, Alger, Alpena, Antrim, Arenac, Bay, Calhoun, Cheboygan, Dickinson, Emmet, Genesee, Gogebic, Grand Traverse, Houghton, Iosco, Iron, Keweenaw, Manistee, Marquette, Midland, Montcalm, Newaygo, Osceola, Ottawa, Van Buren, Wayne. Notes. Biological information available from trap nests set out in Ontario and Michigan by Fye (1965) and Scott (1994, 1996), respectively.

Subgenus Metziella Michener Revision: Snelling (1968).

Hylaeus (Metziella) sparsus (Cresson 1869) County records: Dickinson, Ingham, Jackson, Midland, Washtenaw. Notes. This is a rarely collected species with a relatively broad distribution across eastern North America (Sellers & McCarthy 2015). Hylaeus sparsus has been collected from several plant species, but it has been suggested that it prefers flowers of the family Apiaceae (Sellers & McCarthy 2015) and a recent record from Brooklyn in New York City (3 Jun 2012, Prospect Park Wellhouse) was taken on that host family (JSA, unpublished). Material examined. Dickinson Co.: T43N R30W sec. 19, 20 Jun. 1985 on Ranunculus acris, V.A. Scott (1 ♂ UCMC); Ingham Co.: Okemos, Nancy Moore Park, 42.733 -84.421, 31 Apr. 2017 (1 ♂ TJWC); Jackson Co.: Waterloo SRA, 42.318–84.244, 29 Apr. 2017 (1 ♀ TJWC); Midland Co.: (no locality), 17 Jun. 1945, R.R. Dreisbach (1 ♀ MSUC); Washtenaw Co.: Ann Arbor, Boehnke House, N42.2615 W82.7696, 29 May 2014, C. Vadiya (1 ♀ UMMZ). A second female deposited at MSUC and collected in the 19th century has the following label information: “Ag. Coll. Mich., 6-26-88/76”, but the actual collection locality of the specimen is unknown.

Subgenus Paraprosopis Popov Revision: Snelling (1970).


GIBBS ET AL.

Hylaeus (Paraprosopis) floridanus (Robertson 1893) (wootoni group) County records: Montcalm. Notes. This uncommon species has a much wider distribution than the name suggests, as attested by its occurrence in Michigan, but it has been rarely detected for unknown reasons. Mitchell (1960) recorded H. floridanus from Michigan and Snelling (1970) states the range as Florida north to Maine and west to Minnesota. Material examined. Montcalm Co.: Belding, 4 Aug. 1973, E.R. Hoebeke (2 ♀ MSUC).

Subgenus Prosopis Fabricius Hylaeus (Prosopis) affinis (Smith 1853) County records: Alcona, Allegan, Antrim, Arenac, Barry, Bay, Berrien, Cass, Cheboygan, Clare, Clinton, Dickinson, Eaton, Emmet, Genesee, Gladwin, Huron, Ingham, Ionia, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kent, Lake, Lapeer, Leelanau, Lenawee, Livingston, Mackinac, Manistee, Marquette, Mecosta, Midland, Montcalm, Newaygo, Oakland, Osceola, Ottawa, Roscommon, Saginaw, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw. Notes. Females are difficult to distinguish from H. modestus, see below.

Hylaeus (Prosopis) gaigei (Cockerell 1916) (Michigan holotype) (Fig. 16B) Prosopis gaigei Cockerell 1916: 2. Holotype. ♀ Michigan, Schoolcraft Co.: Floodwood, 26 Jul.1915, F.M. Gaige (NMNH: 23311).

County records: Schoolcraft. Notes. Originally described from two specimens, the holotype has ‘29’ handwritten on the locality label. The specimen was collected as part of the UMMZ Bryant Walker expedition to Michigan’s Upper Peninsula. The collection locality refers to Floodwoods Road, on the Manistique River, 26 mi. northeast of Manistique (Rogers 1918).

FIGURE 16. Colletidae faces from Michigan (not to scale). A) Colletes aestivalis Patton male. B) Hylaeus gaigei (Cockerell) holotype female. Photograph of H. gaigei provided with the permission of the National Museum of Natural History, Smithsonian Institution, 10th and Constitution Ave. N.W., Washington, DC 20560-0193. (http://www.nmnh.si.edu/).



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Snelling (1970) transferred H. gaigei to the subgenus H. (Prosopis) and suggested it may be a synonym of H. modestus. A photograph of the type on the NMNH Department of Entomology website is distinguishable from the standard H. modestus female by its small facial maculations. However, coloration characters of this sort can be unreliable. It is already challenging to distinguish females of H. modestus from related species (Zarrillo et al. 2016), so resolving the identity of H. gaigei requires a dedicated taxonomic study. Several females of H. (Prosopis) collected in Michigan’s Upper Peninsula also seem to have similarly reduced maculations. We provide information on these specimens to facilitate future study of this material. Material examined. Dickinson Co.: T43N R29W Sec. 14, 24 Jun. 1983, M. Arduser, Rubus allegheniensis (2 ♀ MSUC); T43N R30W Sec. 33, 30 Jul. 1983, M. Arduser, Cirsium arvense (1 ♀ MSUC); Channing, T43N R30W Sec. 18, 3 Jul. 1983, R.L. Fischer (1 ♀ MSUC); Gogebic Co.: Watersmeet, 19 Jul. 1983, M. Arduser (1 ♀ MSUC); Marquette Co.: Michigamme, 8 Aug. 1936, Sabrosky (1 ♀ MSUC).

Hylaeus (Prosopis) illinoisensis (Robertson 1896) County records: Alcona, Berrien, Kalamazoo, St. Joseph.

Hylaeus (Prosopis) modestus modestus Say 1837 County records: Allegan, Antrim, Barry, Berrien, Charlevoix, Cheboygan, Clare, Clinton, Ingham, Ionia, Kalamazoo, Kent, Mackinac, Marquette, Midland, Missaukee, Oakland, Ottawa, Shiawassee, St. Clair, Van Buren, Washtenaw, Wayne. Notes.Trap nests in New York described by Krombein (1967) and nests in Missouri sumac stems by Rau (1922, 1930).

Hylaeus (Prosopis) nelumbonis (Robertson 1890) County records: Allegan, Bay, Ingham, Monroe, Sanilac. Notes. This wetland specialist visits Nelumbo and Nymphaea (Hurd 1979), and was collected from Lake Lansing on Pontederia (JG, pers. obs.).

Subgenus Spatulariella Popov Taxonomy: Sheffield et al. (2011a).

Hylaeus (Spatulariella) hyalinatus Smith 1842 County records: Eaton, Ingham, Washtenaw, Wayne. Notes. Hylaeus hyalinatus is a recent introduction to the North American continent (Ascher 2001; Ascher et al. 2006; Tonietto & Ascher 2009). It is one of two members of Hylaeus (Spatulariella) in the Midwest and the only one currently known from Michigan, but see comments on Hylaeus punctatus (Brullé) in Appendix 1. Hylaeus hyalinatus is relatively common in urban centers in southern Michigan, including Detroit, Ann Arbor, Ypsilanti and Lansing.

HALICTIDAE Halictinae


GIBBS ET AL.

Augochlorini Genus Augochlora Smith Subgenus Augochlora Smith s. s. Augochlora (Augochlora) pura pura (Say 1837) County records: Alger, Allegan, Alpena, Antrim, Arenac, Baraga, Barry, Berrien, Calhoun, Cass, Charlevoix, Cheboygan, Chippewa, Clinton, Delta, Dickinson, Eaton, Emmet, Gogebic, Grand Traverse, Gratiot, Houghton, Huron, Ingham, Ionia, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Lapeer, Leelanau, Livingston, Luce, Mackinac, Manistee, Marquette, Mason, Mecosta, Midland, Missaukee, Monroe, Montcalm, Montmorency, Oakland, Oceana, Ontonagon, Osceola, Ottawa, Presque Isle, Saginaw, Shiawassee, St. Clair, St. Joseph, Van Buren, Washtenaw, Wayne, Wexford. Notes. A solitary species that nests in decaying wood as described in detail by Stockhammer (1966). Augochlora pura is a common pollinator of agricultural crops and is relatively abundant on highbush blueberry in spring (Gibbs et al. 2016, 2017).

Genus Augochlorella Sandhouse Taxonomy: Coelho (2004); Mitchell (1960); Ordway (1966a). Behavior: Ground-nesting primitively eusocial bees (Mueller 1996; Ordway 1966b; Packer 1990).

Augochlorella aurata (Smith 1853) (aurata group) County records: Alcona, Alger, Allegan, Alpena, Antrim, Arenac, Barry, Bay, Benzie, Berrien, Branch, Calhoun, Cass, Charlevoix, Cheboygan, Clare, Clinton, Crawford, Dickinson, Eaton, Emmet, Genesee, Gladwin, Grand Traverse, Gratiot, Hillsdale, Huron, Ingham, Ionia, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Lapeer, Leelanau, Lenawee, Livingston, Mackinac, Manistee, Marquette, Mason, Mecosta, Menominee, Midland, Missaukee, Monroe, Montcalm, Montmorency, Muskegon, Newaygo, Oakland, Oceana, Ogemaw, Osceola, Oscoda, Otsego, Ottawa, Presque Isle, Roscommon, Saginaw, Sanilac, Shiawassee, St. Clair, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Ground-nesting primitively eusocial species, based on colonies studied from Kansas and New York (Mueller 1996; Ordway 1966b; Sakagami & Michener 1962), as A. striata. However, the species makes solitary nests at high latitudes (Packer 1990) and may do so in the UP.

Augochlorella persimilis (Viereck 1910) (aurata group) County records: Lenawee, Washtenaw. Notes. Eusocial nesting in Kansas described by Ordway (1966b). This species is rare in Michigan and apparently limited to the southern part of the state.

Genus Augochloropsis Cockerell Taxonomy: Mitchell (1960).

Subgenus Paraugochloropsis Schrottky



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Augochloropsis (Paraugochloropsis) metallica (Fabricius 1793) fulgida (Smith 1853) County records: Allegan, Antrim, Barry, Branch, Cass, Cheboygan, Clinton, Hillsdale, Ingham, Ionia, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Livingston, Montmorency, Muskegon, Oakland, Osceola, Oscoda, Presque Isle, Roscommon, Shiawassee, Van Buren, Washtenaw, Wayne. Notes. A ground-nesting, putatively social species, based on excavation of a multi-female nest at Rose Lake State Wildlife Area, Shiawassee County, Michigan (Gibbs 2017).

Halictini Genus Agapostemon Guerin-Meneville Taxonomy: Mitchell (1960); Roberts (1972). Biology: Polylectic, ground-nesting species with solitary or communal nests (Abrams & Eickwort 1980, 1981; Eickwort 1981; LaBerge & Ribble 1966b; Roberts 1969).

Subgenus Agapostemon Guerin-Meneville s. s. Agapostemon (Agapostemon) sericeus (Förster 1771) (sericeus group) County records: Allegan, Alpena, Antrim, Arenac, Bay, Benzie, Berrien, Branch, Calhoun, Cass, Charlevoix, Cheboygan, Clare, Clinton, Eaton, Gladwin, Gogebic, Grand Traverse, Gratiot, Huron, Ingham, Iosco, Isabella, Jackson, Kalamazoo, Kent, Lake, Lapeer, Leelanau, Livingston, Manistee, Mecosta, Midland, Montcalm, Oakland, Oceana, Ogemaw, Osceola, Ottawa, Saginaw, Sanilac, Shiawassee, St. Joseph, Van Buren, Washtenaw, Wayne. Notes. Solitary, ground-nester (Eickwort 1981; LaBerge & Ribble 1966b; Roberts 1969).

Agapostemon (Agapostemon) splendens (Lepeletier 1841) (splendens group) County records: Alcona, Alger, Allegan, Alpena, Antrim, Arenac, Barry, Berrien, Charlevoix, Cheboygan, Clare, Emmet, Gladwin, Gratiot, Huron, Ingham, Iosco, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Lapeer, Leelanau, Lenawee, Livingston, Manistee, Mason, Mecosta, Midland, Missaukee, Montmorency, Muskegon, Newaygo, Oakland, Oceana, Ogemaw, Ottawa, Roscommon, Saginaw, St. Joseph, Van Buren, Washtenaw, Wexford. Notes. Solitary, ground-nester (Eickwort 1981; LaBerge & Ribble 1966b; Roberts 1969; Stevens 1921).

Agapostemon (Agapostemon) texanus Cresson 1872 (splendens group) County records: Alcona, Alger, Allegan, Baraga, Barry, Branch, Cass, Cheboygan, Clinton, Eaton, Emmet, Gratiot, Ingham, Ionia, Isabella, Jackson, Kalamazoo, Kalkaska, Keweenaw, Lake, Lapeer, Leelanau, Livingston, Mackinac, Manistee, Mecosta, Midland, Missaukee, Montcalm, Montmorency, Newaygo, Oakland, Oceana, Oscoda, Otsego, Ottawa, Presque Isle, Saginaw, Shiawassee, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Solitary, ground-nester (Eickwort 1981; Roberts 1969).

Agapostemon (Agapostemon) virescens (Fabricius 1775) (splendens group) County records: Allegan, Barry, Bay, Benzie, Berrien, Branch, Calhoun, Cass, Cheboygan, Clare, Clinton, Delta, Dickinson, Eaton, Genesee, Gratiot, Hillsdale, Huron, Ingham, Ionia, Jackson, Kalamazoo, Kalkaska, Kent, Lake,


GIBBS ET AL.

Lapeer, Leelanau, Livingston, Mackinac, Midland, Montcalm, Newaygo, Oakland, Oceana, Ottawa, Saginaw, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. A communal ground-nester whose nesting biology was studied in detail in New York (Abrams & Eickwort 1980, 1981).

Genus Halictus Latreille Revision: Mitchell (1960); Sandhouse (1941). Biology. Ground-nesting social species with solitary nesting in some contexts.

Subgenus Nealictus Pesenko Halictus (Nealictus) parallelus Say 1837 County records: Allegan, Barry, Berrien, Branch, Calhoun, Cass, Crawford, Dickinson, Eaton, Gratiot, Ingham, Jackson, Kalamazoo, Kent, Livingston, Macomb, Monroe, Montcalm, Oakland, Oceana, Ottawa, Saginaw, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw. Notes. Aspects of H. parallelus nesting have been documented outside of Michigan (Hungerford & Willlams 1912; Packard 1868; Sakagami & Michener 1962).

Subgenus Odontalictus Robertson Halictus (Odontalictus) ligatus Say 1837 County records: Alcona, Allegan, Antrim, Arenac, Barry, Bay, Berrien, Calhoun, Cass, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Eaton, Genesee, Gladwin, Grand Traverse, Gratiot, Hillsdale, Huron, Ingham, Ionia, Iosco, Isabella, Jackson, Kalamazoo, Kent, Lake, Lapeer, Leelanau, Lenawee, Livingston, Manistee, Mason, Mecosta, Midland, Montcalm, Muskegon, Newaygo, Oakland, Oceana, Osceola, Ottawa, Presque Isle, Saginaw, Shiawassee, St. Clair, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne. Notes. Social ground-nester. The nesting biology of H. ligatus has been well-studied (Cane 1991; Eickwort 1985; Hicks 1926; Michener & Bennett 1977; Packer 1986; Packer & Knerer 1986; Rau 1922; Rehan et al. 2013; Sakagami & Michener 1962) as has its relationship to its crypic sister species, H. poeyi Lepeletier (Carman & Packer 1996; Danforth et al. 1998, 1999; Packer 1999; Packer et al. 2016).

Subgenus Protohalictus Pesenko Halictus (Protohalictus) rubicundus (Christ 1791) County records: Alcona, Alger, Allegan, Alpena, Antrim, Baraga, Barry, Benzie, Berrien, Calhoun, Cass, Cheboygan, Chippewa, Clare, Clinton, Crawford, Delta, Dickinson, Eaton, Genesee, Gogebic, Grand Traverse, Gratiot, Houghton, Huron, Ingham, Ionia, Iosco, Iron, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lake, Lapeer, Leelanau, Lenawee, Livingston, Mackinac, Manistee, Marquette, Mason, Mecosta, Midland, Montcalm, Montmorency, Muskegon, Newaygo, Oakland, Oceana, Otsego, Ottawa, Roscommon, Saginaw, Sanilac, Schoolcraft, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Holarctic. Social ground-nester, but displays polyethism, reverting to solitary behavior at high latitudes and altitude (Eickwort et al. 1996; Field et al. 2010; Potts & Willmer 1997). The nesting of this widespread species has been documented across North America (Atwood 1933; Eickwort et al. 1996; Hicks 1926; Knerer & Atwood 1962; Soucy 2002; Yanega 1988).



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Subgenus Seladonia Robertson Halictus (Seladonia) confusus confusus Smith 1853 County records: Alcona, Allegan, Alpena, Antrim, Arenac, Baraga, Barry, Bay, Benzie, Berrien, Calhoun, Cass, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Crawford, Delta, Dickinson, Eaton, Emmet, Genesee, Gladwin, Grand Traverse, Gratiot, Hillsdale, Houghton, Huron, Ingham, Ionia, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lake, Lapeer, Leelanau, Lenawee, Livingston, Mackinac, Macomb, Manistee, Marquette, Mason, Mecosta, Menominee, Midland, Missaukee, Monroe, Montcalm, Montmorency, Muskegon, Newaygo, Oakland, Oceana, Ogemaw, Ontonagon, Osceola, Oscoda, Otsego, Ottawa, Roscommon, Saginaw, Sanilac, Schoolcraft, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Holarctic. A social ground-nester based on studies of southern Ontario populations (Knerer & Atwood 1962; Richards et al. 2010), although Nova Scotia populations were thought to be solitary (Atwood 1933).

Genus Lasioglossum Curtis Taxonomy: Gibbs (2010; 2011); Gibbs et al. (2013); Knerer & Atwood (1964b); McGinley (McGinley 1986, 2003); Mitchell (1960). Biology. Lasioglossum is extremely behaviorally diverse including solitary, communal, eusocial nesting in underground burrows or rotten logs, a range of floral host preferences, and multiple origins of cleptoparasitism (Danforth et al. 2003; Gibbs et al. 2012a, 2013; Michener 1974; Sakagami & Michener 1962; Yanega 1997).

Subgenus Dialictus Robertson Taxonomy: Gibbs (2010b, 2011); Mitchell (1960). Biology. Most species are expected to be primitively eusocial ground-nesters (Danforth et al. 2003; Gibbs et al. 2012b), but there are a number of exceptions (Knerer 1969; Packer 1994) and categorization of an entire species can be problematic (Wcislo 1997). Only a small fraction have had their nesting biology studied in any detail.

Lasioglossum (Dialictus) abanci (Crawford 1932) (viridatum group) County records: Clinton, Ingham, Osceola, Ottawa, Saginaw, Washtenaw, Wexford. Notes. A problematic species that requires additional study. Individuals in the north of its range show some minor differences from the type material and specimens collected near the type locality in the Appalachian Mountains of North Carolina.

Lasioglossum (Dialictus) achilleae (Mitchell 1960) County records: Allegan, Barry, Lake, Manistee, Mason. Notes. Uncommon throughout its range.

Lasioglossum (Dialictus) admirandum (Sandhouse 1924) (viridatum group) County records: Allegan, Barry, Calhoun, Cass, Clinton, Eaton, Gratiot, Huron, Ingham, Jackson, Kalamazoo, Lapeer, Manistee, Roscommon, Saginaw, St. Joseph, Tuscola, Van Buren, Washtenaw.


GIBBS ET AL.

Lasioglossum (Dialictus) albipenne (Robertson 1890) County records: Barry, Berrien, Cass, Cheboygan, Clinton, Dickinson, Eaton, Gratiot, Huron, Ingham, Jackson, Kalamazoo, Lapeer, Livingston, Manistee, Montcalm, Oceana, Roscommon, Saginaw, St. Joseph, Tuscola.

Lasioglossum (Dialictus) anomalum (Robertson 1892) County records: Allegan, Barry, Berrien, Calhoun, Cass, Clinton, Eaton, Gratiot, Huron, Ingham, Ionia, Jackson, Kalamazoo, Livingston, Mecosta, Midland, Montcalm, Newaygo, Oakland, Ottawa, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw,. Notes. The type species of the subgenus.

Lasioglossum (Dialictus) atwoodi Gibbs 2010 (viridatum group) County records: Allegan, Antrim, Van Buren.

Lasioglossum (Dialictus) bruneri (Crawford 1902) County records: Allegan, Barry, Calhoun, Cass, Clinton, Ingham, Jackson, Kalamazoo, Lapeer, Livingston, Midland, Montcalm, Oceana, Ottawa, Saginaw, St. Joseph, Van Buren, Washtenaw, Wayne. Notes. A common species in urban gardens of New York City (Matteson et al. 2008), so potentially an urbanassociated species.

Lasioglossum (Dialictus) callidum (Sandhouse 1924) County records: Kalamazoo. Notes. Past confusion with L. versatum (see Gibbs 2010).

Lasioglossum (Dialictus) cattellae (Ellis 1913) County records: Clinton, Ingham, Jackson, Kalamazoo, Lenawee, Mecosta, Van Buren, Washtenaw.

Lasioglossum (Dialictus) ceanothi (Mitchell 1960) County records: Mackinac. Notes. Rare. Record based on paratype.

Lasioglossum (Dialictus) coeruleum (Robertson 1893) County records: Allegan, Barry, Benzie, Berrien, Cheboygan, Clinton, Delta, Grand Traverse, Ingham, Jackson, Kalamazoo, Kalkaska, Kent, Lapeer, Leelanau, Livingston, Montcalm, Osceola, Ottawa, Saginaw, Shiawassee, St. Clair, Van Buren, Washtenaw. Notes. Wood-nesting and a social species based on detailed studies in Kansas (Stockhammer 1967). Nests have been recorded from elm (Stockhammer 1967) and maple (Knerer & Atwood 1962).



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Lasioglossum (Dialictus) coreopsis (Robertson 1902) County records: none. Notes. Michigan and Wisconsin are among the northern limits of its range. No Michigan specimens were examined, but the species was included based on Mitchell (1960). It is most likely to occur in the southwest near Lake Michigan.

Lasioglossum (Dialictus) cressonii (Robertson 1890) County records: Alcona, Alger, Allegan, Alpena, Antrim, Baraga, Barry, Benzie, Berrien, Calhoun, Cass, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Delta, Dickinson, Emmet, Gogebic, Grand Traverse, Gratiot, Hillsdale, Houghton, Huron, Ingham, Ionia, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lake, Lapeer, Leelanau, Lenawee, Livingston, Mackinac, Manistee, Marquette, Midland, Missaukee, Montcalm, Montmorency, Muskegon, Oakland, Oceana, Ogemaw, Ontonagon, Osceola, Oscoda, Otsego, Ottawa, Saginaw, St. Clair, St. Joseph, Van Buren, Washtenaw, Wexford. Notes. Recorded as a wood-nester based on a specimen collected in Maryland by Krombein and Moure (Mitchell 1960). It was never encountered during multiple excavations of logs in the Lansing area.

Lasioglossum (Dialictus) dreisbachi (Mitchell 1960) (viridatum group) Dialictus dreisbachi Mitchell, 1960: 391. (Michigan holotype) Holotype. ♀ USA: Michigan: Clare Co.: 24 May 1958 (R. & K. Dreisbach) (NMNH).

County records: Allegan, Antrim, Arenac, Charlevoix, Cheboygan, Clare, Clinton, Crawford, Eaton, Gladwin, Ionia, Iosco, Isabella, Kalkaska, Kent, Lake, Lapeer, Lenawee, Midland, Montcalm, Ottawa, Saginaw, Shiawassee, Van Buren, Washtenaw. Notes. This species appears to have a northern distribution, reaching its southernmost limits in states that border Canada. Its range remains uncertain due to insufficient sampling in boreal regions of Canada and northern states, and because bee specialists have not been able to distinguish this from other species in the taxonomically challenging Lasioglossum viridatum species group of Gibbs (2010b). Nevertheless, L. dreisbachi has unique mesepisternal sculpturing, which abruptly changes from coarsely rugose to smooth (Gibbs 2010b). Material examined. Allegan Co.: Trevor Nichols Res. Ctr. N42.6 W86.155, 5–16 Jun. 2015, Malaise trap, J. Gibbs & Y. Nozoe (1 ♀ JBWM); Lapeer Co.: Lapper SGA, Norway Lansing, N43.145, W83.353, 28 Jun. 2017, T. Wood (1 ♀ TJWC); Ottawa Co.: Marne, 2 km S, N43.0175 W85.8411, 4 Jun. 2015, Leucanthemum vulgare, S. Way (1 ♀ MSUC: RI); Van Buren Co.: Brown Lk., N42.0992 W86.1862, 12 May 2014, Prunus cerasus (1 ♀ MSUC: RI).

Lasioglossum (Dialictus) ellisiae (Sandhouse 1924) (tegulare group) County records: Allegan, Barry, Calhoun, Cass, Clinton, Eaton, Gratiot, Hillsdale, Ingham, Jackson, Kalamazoo, Lake, Leelanau, Mecosta, Montcalm, Oakland, Ottawa, Saginaw, St. Joseph, Tuscola, Van Buren, Washtenaw, Wexford. Notes. Prior to Gibbs (2009), L. ellisiae was included in L. tegulare.

Lasioglossum (Dialictus) ephialtum Gibbs 2010 (viridatum group) County records: Allegan, Cass, Genesee, Huron, Ingham, Jackson, Lapeer, Leelanau, Oceana, Saginaw, Washtenaw, Wayne. Notes. Most commonly collected in urban settings.


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Lasioglossum (Dialictus) fattigi (Mitchell 1960) (viridatum group) County records: Cass, Clinton. Notes. Rarely seen, but recorded by Tuell et al. (2009) based on JG determinations. Easily confused with L. paradmirandum.

Lasioglossum (Dialictus) floridanum (Robertson 1892) (pilosum group) (New state record) County records: Lake, Newaygo, Ottawa, Van Buren. Notes. Until Gibbs (2011), this was considered a subspecies of L. pilosum (Smith) that was restricted to the southeastern USA. Lasioglossum floridanum was found to range further north to overlap extensively with L. pilosum, including the northwest corner of Illinois (Gibbs 2011; Grundel et al. 2011). It was raised from subspecific status by Gibbs (2011) based on the morphological and molecular differences and sympatric distribution with L. pilosum. Material examined. Lake Co.: (no locality), 5 May 1945, R.R. Dreisbach (1 ♀ MSUC); Newaygo Co.: (no locality), 15 Jul. 1950, R.R. Dreisbach (1 ♀ MSUC); Van Buren Co.: Covert, 3 mi. SSE, 1 Jul. 2013, Hypochaeris radicata (1 ♀ MSUC: RI).

Lasioglossum (Dialictus) foveolatum (Robertson 1902) County records: Allegan, Barry, Berrien, Clinton, Jackson, Kalamazoo, Kalkaska, Kent, Livingston, St. Joseph, Washtenaw.

Lasioglossum (Dialictus) gotham Gibbs 2011 (New state record) County records: Ingham, Van Buren. Notes. Nesting biology described by Batra (1987) as laevissimum. Nests have been seen frequently near East Lansing, Michigan in soil at the base of fallen trees (JG, pers. obs.). Material examined. Ingham Co.: Fenner Nature Ctr., N42.709, W84.521, 4–24 Aug. 2015, lab reared from excavated nest (10 ♀ 6 ♂ JBWM); Van Buren Co.: 2 km SW Laurence, N42.2071, W86.0822, 6 May 2015, ex. Prunus cerasus (1 ♀ MSUC:RI).

Lasioglossum (Dialictus) hartii (Robertson 1892) County records: Barry, Saginaw. Notes. An apparent wetland specialist.

Lasioglossum (Dialictus) heterognathum (Mitchell 1960) County records: Allegan, Bay, Huron, Kalamazoo, Lake, Montcalm, Newaygo, Sanilac, Shiawassee, Van Buren, Washtenaw.

Lasioglossum (Dialictus) hitchensi Gibbs 2012 County records: Allegan, Barry, Bay, Berrien, Calhoun, Cass, Clinton, Eaton, Genesee, Gratiot, Huron, Ingham,



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Ionia, Jackson, Kalamazoo, Lapeer, Livingston, Manistee, Midland, Montcalm, Newaygo, Oakland, Ogemaw, Ottawa, Roscommon, Saginaw, Shiawassee, St. Clair, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne.

Lasioglossum (Dialictus) illinoense (Robertson 1892) County records: Allegan, Barry, Berrien, Calhoun, Cass, Clinton, Eaton, Genesee, Gratiot, Ingham, Ionia, Jackson, Kalamazoo, Kent, Leelanau, Lenawee, Livingston, Montcalm, Ottawa, Saginaw, Shiawassee, St. Joseph, Van Buren, Washtenaw.

Lasioglossum (Dialictus) imitatum (Smith 1853) County records: Allegan, Arenac, Barry, Bay, Benzie, Berrien, Branch, Calhoun, Cass, Clare, Clinton, Eaton, Emmet, Genesee, Grand Traverse, Gratiot, Hillsdale, Huron, Ingham, Ionia, Isabella, Jackson, Kalamazoo, Kent, Lake, Lapeer, Leelanau, Lenawee, Livingston, Manistee, Mecosta, Midland, Montcalm, Muskegon, Newaygo, Oakland, Oceana, Ogemaw, Ottawa, Saginaw, Sanilac, Shiawassee, St. Clair, St. Joseph, Van Buren, Washtenaw. Notes. A primitively eusocial ground-nester whose nesting biology was studied in detail in Kansas (Michener & Wille 1961).

Lasioglossum (Dialictus) laevissimum (Smith 1853) = Dialictus solidaginis Mitchell, 1960: 443. (Michigan holotype) Holotype. ♂ USA: Michigan: Marquette Co.: 20.vi.1955, R.R. Dreisbach (NCSU). Synonymy by Gibbs (2010b).

County records: Alger, Allegan, Antrim, Baraga, Barry, Cheboygan, Chippewa, Clinton, Delta, Dickinson, Gogebic, Houghton, Ingham, Iron, Kalamazoo, Keweenaw, Livingston, Luce, Mackinac, Marquette, Montcalm, Ontonagon, Otsego, Roscommon, Schoolcraft, St. Clair, Van Buren, Washtenaw, Wexford. Notes. A primitively euocial ground-nester based on studies from Alberta and Nova Scotia (Brittain 1933; Packer 1992; Packer et al. 1989a).

Lasioglossum (Dialictus) leucocomum (Lovell 1908) (pilosum group) = Dialictus otsegoensis Mitchell, 1960: 440. (Michigan holotype) Holotype. ♂ USA: Michigan: Ostego Co.: 7–24.vii.1955, R.R. Dreisbach (NMNH). Synonymy by Gibbs (2011).

County records: Allegan, Antrim, Barry, Benzie, Berrien, Calhoun, Cass, Cheboygan, Clinton, Crawford, Delta, Dickinson, Emmet, Genesee, Grand Traverse, Gratiot, Hillsdale, Huron, Ingham, Isabella, Jackson, Kalamazoo, Kalkaska, Lake, Lapeer, Leelanau, Livingston, Macomb, Manistee, Midland, Montcalm, Montmorency, Muskegon, Newaygo, Oakland, Oceana, Ogemaw, Osceola, Oscoda, Otsego, Ottawa, Saginaw, Sanilac, Schoolcraft, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw, Wexford. Notes. Until Gibbs (2010), L. leucocomum was included within L. pilosum.

Lasioglossum (Dialictus) lineatulum (Crawford 1906) Halictus lineatulus Crawford 1906: 5. Syntype ♀ USA, “Ag. Coll. Mich.”, 5 [4-11 Oct.] Oct. 1893, R.H. Wolcott (NMNH: 12071).

County records: Alcona, Allegan, Alpena, Antrim, Arenac, Barry, Bay, Benzie, Berrien, Branch, Calhoun, Cass, Cheboygan, Clare, Clinton, Crawford, Delta, Dickinson, Eaton, Emmet, Genesee, Gladwin, Gogebic, Grand


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Traverse, Gratiot, Huron, Ingham, Ionia, Iosco, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Lapeer, Leelanau, Livingston, Manistee, Mason, Mecosta, Midland, Missaukee, Montcalm, Montmorency, Muskegon, Newaygo, Oakland, Oceana, Osceola, Oscoda, Otsego, Ottawa, Roscommon, Saginaw, Sanilac, Shiawassee, St. Clair, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. A ground-nesting primitively eusocial species studied in detail in New York (Eickwort 1986). The type locality is uncertain. A number of specimens at MSUC were labeled as “Ag. Coll. Mich.” at a time when MSU was known as the “Agricultural College of the State of Michigan”. However, these labels may refer to the specimens as the property of the Agricultural College rather than the original collection locality. Careful records of accession were not maintained in the early management of MSUC. The type series is likely from Michigan since the collector, Robert H. Wolcott, studied at the University of Michigan until 1893 (Hungerford 1935), which coincides with the collection date. The species is common in the state and region.

Lasioglossum (Dialictus) lionotum (Sandhouse 1923) (cephalotes group) County records: Clare, Ingham, Isabella, Kalamazoo, Leelanau, Midland, Shiawassee, Van Buren. Notes. A social parasite of L. imitatum (Smith) (Wcislo 1997), L. lionotum was recently recognized as a senior synonym of Paralictus asteris Mitchell (Gibbs 2011). Material examined. Ingham Co.: East Lansing, White Hills Park, 4 Sep. 2016, J. Gibbs (1 ♀ JBWM); Van Buren Co.: Almena, 3 km N, N42.2843 W85.8537, 13 Aug. 2014, J. Gibbs (2 ♀ JBWM).

Lasioglossum (Dialictus) michiganense (Mitchell 1960) (platyparium group) Paralictus michiganensis Mitchell, 1960: 448. (Michigan holotype) Holotype. ♀ USA: Michigan: Wayne Co.: 2 Jun. 1940, Beebe (NMNH).

County records: Allegan, Ingham, Kalamazoo, Wayne. Notes. Lasioglossum michiganense is presumed to be a social parasite of other L. (Dialictus) (Michener 1978, Gibbs et al. 2012), but its host is unknown and range limits remain uncertain. The female examined was collected in a riparian area adjacent to a woodlot. Collection locations are frequently associated with nearby wooded habitat (e.g., Giles & Ascher 2006). However, it has also been reported from an urban park in the city of Chicago (Tonietto & Ascher 2009). Material examined. Allegan Co.: Trevor Nichols Res. Ctr. N42.6 W86.155, 26 May-5 Jun. 2015, Malaise trap, J. Gibbs & Y. Nozoe (1 ♀ JBWM); Ingham Co.: East Lansing. N42.748 W84.489, 23 Jul. 2016, J. Gibbs (1 ♀ JBWM; http://bugguide.net/node/view/1263039).

Lasioglossum (Dialictus) nigroviride (Graenicher 1911) County records: Allegan, Cheboygan, Chippewa, Clinton, Dickinson, Emmet, Gogebic, Ingham, Kalamazoo, Kalkaska, Keweenaw, Mackinac, Marquette, Midland, Osceola, Oscoda, Ottawa, Roscommon, Van Buren. Notes. Wood-nesting. This relatively large species was observed nesting in a rotten log at Six-mile Creek, Ithaca, New York (J. Gibbs, pers. obs.).

Lasioglossum (Dialictus) novascotiae (Mitchell 1960) (viridatum group) County records: Allegan, Mackinac.



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Lasioglossum (Dialictus) oblongum (Lovell 1905) (viridatum group) County records: Alger, Allegan, Clinton, Dickinson, Gogebic, Ingham, Jackson, Missaukee, Osceola, Ottawa, Saginaw, Van Buren. Notes. A wood-nesting species (Gibbs 2011).

Lasioglossum (Dialictus) obscurum (Robertson 1892) (viridatum group) County records: Allegan, Barry, Berrien, Cheboygan, Clinton, Ingham, Ionia, Jackson, Kalamazoo, Monroe, Saginaw, St. Clair, Van Buren, Wayne.

Lasioglossum (Dialictus) oceanicum (Cockerell 1916) County records: Allegan, Antrim, Barry, Berrien, Branch, Calhoun, Cass, Clare, Clinton, Eaton, Gladwin, Gratiot, Huron, Ingham, Ionia, Iosco, Jackson, Kalamazoo, Kalkaska, Kent, Lapeer, Leelanau, Livingston, Manistee, Mecosta, Midland, Montcalm, Muskegon, Newaygo, Oakland, Oceana, Osceola, Ottawa, Saginaw, Sanilac, St. Joseph, Tuscola, Van Buren, Wexford. Notes. There has been some confusion over the name of this bee. Halictus nymphaearum was proposed as a replacement for the preoccupied Halictus palustris Robertson (1890) (not Halictus palustris Morawitz 1876), and subsequently the species has most often been referred to as Dialictus nymphaearum (Robertson 1895) or as L. (D.) nymphaearum (Robertson 1895). Gibbs (2010b) resurrected the name L. oceanicum (Cockerell 1916) from synonymy as the valid name for this species after discovering that the apparent lectotype of H. palustris Robertson, labeled as such and deposited at ANSP, pertained to another well-known species L. albipenne (Robertson 1890) rather than to Dialictus nymphaearum sensu auct. (e.g., Mitchell, 1960). Gibbs (2011) reversed this decision based on an inability to find a valid lectotype designation, but this was in fact validly published by Cresson (1928) showing the decision by Gibbs (2010) to have been correct (contra Gibbs 2011). Thus, Lasioglossum oceanicum is the correct name for this species, i.e. the names Halictus palustris Robertson and H. nymphaearum return to synonymy under L. albipenne.

Lasioglossum (Dialictus) paradmirandum (Knerer & Atwood 1966) (viridatum group) County records: Barry, Berrien, Calhoun, Cass, Clinton, Eaton, Gratiot, Huron, Ingham, Ionia, Kalamazoo, Kent, Lapeer, Midland, Montcalm, Newaygo, Oakland, Ogemaw, Osceola, Ottawa, Saginaw, Shiawassee, Tuscola, Washtenaw, Wayne.

Lasioglossum (Dialictus) perpunctatum (Ellis 1913) County records: Allegan, Arenac, Barry, Benzie, Berrien, Calhoun, Cheboygan, Chippewa, Clare, Clinton, Dickinson, Eaton, Gratiot, Hillsdale, Huron, Ingham, Ionia, Jackson, Kalamazoo, Kalkaska, Lake, Lapeer, Leelanau, Livingston, Manistee, Mecosta, Midland, Montcalm, Montmorency, Oakland, Osceola, Ottawa, Roscommon, Shiawassee, St. Joseph, Tuscola, Van Buren, Wexford.

Lasioglossum (Dialictus) pictum (Crawford 1902) = Dialictus muskegonensis Mitchell, 1960: 439. (Michigan holotype) Holotype. ♂ USA: Michigan: Muskegon Co.: 2.viii.1944, R.R. Dreisbach (NMNH). Synonymy by Mitchell (1962).

County records: Allegan, Barry, Berrien, Cass, Clare, Clinton, Gladwin, Ingham, Jackson, Leelanau, Livingston,


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Macomb, Manistee, Mecosta, Midland, Monroe, Montcalm, Muskegon, Oceana, Ottawa, Shiawassee, St. Joseph, Van Buren. Notes. Michigan is at the eastern range limits of this species, as records of similar bees from the Atlantic states pertain to Lasioglossum (Dialictus) arantium Gibbs 2011.

Lasioglossum (Dialictus) pilosum (Smith 1853) (pilosum group) County records: Alcona, Allegan, Antrim, Arenac, Barry, Bay, Berrien, Calhoun, Cass, Cheboygan, Clare, Clinton, Dickinson, Eaton, Emmet, Genesee, Gladwin, Grand Traverse, Gratiot, Hillsdale, Huron, Ingham, Ionia, Iosco, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Lapeer, Leelanau, Livingston, Mackinac, Macomb, Manistee, Marquette, Mason, Mecosta, Midland, Montcalm, Muskegon, Newaygo, Oakland, Oceana, Ogemaw, Osceola, Oscoda, Otsego, Ottawa, Roscommon, Saginaw, Schoolcraft, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw, Wexford.

Lasioglossum (Dialictus) planatum (Lovell 1905) (viridatum group) County records: Charlevoix, Cheboygan, Dickinson, Grand Traverse, Huron, Keweenaw, Ottawa, Saginaw, Tuscola, Wexford.

Lasioglossum (Dialictus) platyparium (Robertson 1895) (platyparium group) County records: Clinton, Eaton, Van Buren, Washtenaw. Notes. Social parasite or cleptoparasite. Host unknown but presumably a L. (Dialictus).

Lasioglossum (Dialictus) pruinosum (Robertson 1892) (pilosum group) County records: Cass, Clinton, Ingham, Leelanau, Tuscola. Notes. Nest described by Melander & Brues (1903).

Lasioglossum (Dialictus) rufulipes (Cockerell 1938) County records: Dickinson, Marquette. Notes. Lasioglossum rufulipes has a boreal distribution. The UP of Michigan is at the southern edge of its range in the east (Gibbs 2011) and is otherwise unknown from the Eastern United States although it occurs as far east as Quebec in Canada. It was previously recorded from Michigan by Gibbs (2011). Material examined. Dickinson Co.: T44 R29W sec. 14, 5 Aug.1982, Solidago, D.K. Young (1 ♂ MSUC); Marquette Co.: Michigamme, 8 Aug. 1936, C. Sabrosky (1 ♂ MSUC).

Lasioglossum (Dialictus) sagax (Sandhouse 1924) (viridatum group) County records: Oceana. Notes. A taxonomically challenging and problematic species that would benefit from additional study.

Lasioglossum (Dialictus) sheffieldi Gibbs 2010 (perdifficile group) (New state record) County records: Alger.



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Notes. Lasioglossum sheffieldi is a sand dune specialist, originally described from coastal areas of the Canadian Maritime Provinces and the western bank of the St. Lawrence River in Quebec (Gibbs 2010b). Two additional specimens, one from a Manitoba dune system and the other from the west coast of Lake Michigan in Wisconsin, were examined but not included as paratypes due to their disjunct localities (Gibbs 2010b). A long series of males and females were recently examined from Pictured Rocks National Lakeshore, which fills an important gap in the known distribution of this species, and points to the likelihood of additional unknown localities in other dune and lakeshore habitats, which may connect the inland populations to the populations on the Atlantic Coast. Material examined. Alger Co.: Pictured Rocks National Lakeshore (PWRC).

Lasioglossum (Dialictus) smilacinae (Robertson 1899) County records: Barry, Berrien, Cheboygan, Clinton, Ingham, Jackson, Kalamazoo, Lake, Lapeer, Livingston, Manistee, Mecosta, Missaukee, Montcalm, Newaygo, Oakland, Osceola, Ottawa, Van Buren, Washtenaw. Notes. Nest descriptions by Brittain (1933) are referred to L. laevissimum (see above).

Lasioglossum (Dialictus) subversans (Mitchell 1960) Dialictus subversans Mitchell, 1960: 419. (Michigan holotype) Holotype. ♀ USA: Michigan: Otsego Co.: 26.iv.1944, R.R. Dreisbach (NMNH).

County records: Dickinson, Ontonagon, Otsego, Van Buren. Notes. Lasioglossum subversans was poorly documented in the literature until revisionary studies by Gibbs (2010b, 2011) who treated it as synonymous with Dialictus perpunctatulus Knerer & Atwood. Lasioglossum subversans appears to have a northern distribution, extending across Canada but only reaching south into the USA in northern Michigan and Maine. Material examined. Dickinson Co.: Ralph, 19 Jul. 1986, R.L. Fischer (1 ♀ MSUC); T43 R30W sec. 7, 12 Jun.1983, Malaise trap, M. Arduser (1 ♀ MSUC); T43 R29W sec. 14, 24 Jun.1983, Rubus allegheniensis, M. Arduser (1 ♀ MSUC); T44 R29W sec. 5, 7 Aug.1982, Solidago, D.K. Young (1 ♂ MSUC); T44 R29W sec. 22, 11 Aug.1982, D.K. Young (2 ♂ MSUC).

Lasioglossum (Dialictus) subviridatum (Cockerell 1938) (viridatum group) County records: Allegan, Charlevoix, Clinton, Ingham, Ottawa, Saginaw, Shiawassee, Van Buren, Wayne. Notes. Wood-nesting (Gibbs 2011). Nests of this species are common under the bark of fallen trees in wooded areas near East Lansing, Michigan (JG, pers. obs.). Ripiphorid beetles have been reared from L. subviridatum nests (J.G., unpublished data).

Lasioglossum (Dialictus) taylorae Gibbs 2010 (viridatum group) County records: Charlevoix, St. Clair. Notes. Lasioglossum taylorae was recently described from a small number of specimens. It was subsequently documented from Michigan without specific localities (Gibbs 2011). Material examined. Charlevoix Co.: Susan Creek Nature Preserve, N43.361 W85.1181, 11 Jun. 2017, J. Gibbs ex. Geum rivale (1 ♀ MSUC); St. Clair Co.: Pine River Nat. Ctr., N42.979 W82.654, 23 Jul. 2015, J. Gibbs (1 ♀ MSUC).


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Lasioglossum (Dialictus) tegulare (Robertson 1890) (tegulare group) County records: Allegan, Cass, Jackson, Ingham, Leelanau, Livingston, Saginaw, Tuscola, Van Buren. Notes. Many historical records, especially northward, pertain to L. ellisiae.

Lasioglossum (Dialictus) tenax (Sandhouse 1924) County records: Alger, Dickinson, Keweenaw, Marquette. Notes. This species has an alpine/boreal distribution. Michigan’s UP is near the southern edge of its range in the east. Nests in Alberta are solitary (Packer 1994).

Lasioglossum (Dialictus) timothyi Gibbs 2010 County records: Allegan, Crawford, Grand Traverse, Kalamazoo, Kalkaska, Lake, Manistee, Midland, Missaukee, Montmorency, Muskegon, Newaygo, Osceola, Ottawa.

Lasioglossum (Dialictus) trigeminum Gibbs 2011 County records: Gogebic, Ottawa. Notes. A primitively eusocial ground-nester (see Gibbs 2011; Michener 1966 as versatum).

Lasioglossum (Dialictus) versans (Lovell 1905) = Evylaeus divergenoides Mitchell, 1960: 351 (Michigan holotype) Holotype. ♂ USA: Michigan: Mackinac Co.: St. Ignace, 23 Jul. 1921, T.H. Hubbell (UMMZ). Synonymy by Gibbs (2011).

County records: Alcona, Alger, Allegan, Antrim, Arenac, Baraga, Barry, Bay, Benzie, Cass, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Crawford, Delta, Dickinson, Emmet, Gladwin, Gogebic, Grand Traverse, Houghton, Ingham, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Keweenaw, Lapeer, Leelanau, Luce, Mackinac, Manistee, Marquette, Mason, Midland, Missaukee, Montmorency, Muskegon, Oceana, Ogemaw, Ontonagon, Osceola, Oscoda, Otsego, Ottawa, Roscommon, Schoolcraft, Shiawassee, Van Buren, Wexford.

Lasioglossum (Dialictus) versatum (Robertson 1902) County records: Allegan, Barry, Calhoun, Cass, Clinton, Dickinson, Eaton, Gratiot, Huron, Ingham, Ionia, Jackson, Kalamazoo, Lapeer, Livingston, Mecosta, Montcalm, Oakland, Oceana, Osceola, Ottawa, Saginaw, St. Clair, St. Joseph, Tuscola, Van Buren, Washtenaw. Notes. Primitively eusocial ground-nesting species based on Kansas and Ontario populations (Breed 1975; Knerer & Atwood 1966). Note past taxonomic confusion with L. callidum (see above).

Lasioglossum (Dialictus) vierecki (Crawford 1904) County records: Allegan, Arenac, Barry, Berrien, Calhoun, Clare, Clinton, Delta, Gladwin, Gratiot, Ingham, Ionia, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Leelanau, Livingston, Manistee, Midland, Montcalm, Muskegon, Newaygo, Oakland, Oceana, Oscoda, Otsego, Ottawa, St. Joseph, Tuscola, Van Buren, Washtenaw, Wexford. Notes. A sandy soil loving species. Lasioglossum vierecki is a solitary species (Knerer 1969; Packer 1993).



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Lasioglossum (Dialictus) viridatum (Lovell 1905) (viridatum group) = Dialictus lepidus Mitchell, 1960: 438. (Michigan holotype) Holotype. ♂ USA: Michigan: Keweenaw Co.: 27 Aug. [year obscured], R.R. Dreisbach (NMNH). Synonymy by Mitchell (1962).

County records: Alger, Allegan, Antrim, Baraga, Benzie, Calhoun, Cass, Charlevoix, Chippewa, Clinton, Dickinson, Emmet, Gogebic, Huron, Kalamazoo, Keweenaw, Mackinac, Marquette, Midland, Monroe, Ontonagon, Roscommon, Saginaw, Schoolcraft, Shiawassee, St. Clair, St. Joseph, Tuscola, Van Buren, Wexford. Notes. Social nesting in Nova Scotia was reported by Atwood (1933), but the identification is unreliable (Zarrillo et al. 2016).

Lasioglossum (Dialictus) weemsi (Mitchell 1960) County records: Ottawa, Washtenaw.

Lasioglossum (Dialictus) zephyrum (Smith 1853) County records: Allegan, Bay, Berrien, Calhoun, Cheboygan, Clare, Clinton, Eaton, Huron, Ingham, Ionia, Kalamazoo, Kent, Lapeer, Leelanau, Manistee, Mason, Mecosta, Midland, Montcalm, Newaygo, Oakland, Oceana, Ogemaw, Sanilac, Shiawassee, Van Buren, Wayne. Notes. A primitively eusocial ground-nester, commonly found in southfacing banks or horizontal ground near streams and rivers (Batra 1964, 1966). A well-studied species of halictid bee; for a list of references see Gibbs (2010) and Moure & Hurd (1987).

Subgenus Evylaeus Robertson Taxonomy: Gibbs et al. (2013).

Lasioglossum (Evylaeus) cinctipes (Provancher 1888) County records: Antrim, Barry, Cass, Cheboygan, Clare, Clinton, Dickinson, Emmet, Genesee, Grand Traverse, Huron, Ingham, Iosco, Kalamazoo, Kalkaska, Keweenaw, Lapeer, Leelanau, Livingston, Marquette, Montcalm, Oceana, Ogemaw, Ontonagon, Ottawa, Van Buren, Wexford. Notes. A eusocial ground-nester (Atwood 1933; Knerer & Plateaux-Quénu 1966; Packer et al. 1989a; b).

Subgenus Hemihalictus Cockerell Revision: Gibbs et al. (2013). Biology. All species are expected to be solitary ground-nesters, but most have never been studied. Prior to Gibbs et al. (2013), the subgenus was monotypic, including only L. lustrans.

Lasioglossum (Hemihalictus) birkmanni (Crawford 1906) County records: Allegan, Berrien, Cheboygan, Ingham, Jackson, Kalamazoo, Muskegon, Oceana, Shiawassee, Van Buren.


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Notes. The application of this name has undergone a recent change (Gibbs et al. 2013). Recent examination of the lectotype of Halictus quadrimaculatus Robertson 1890 (described as 4-maculatus; Robertson’s name was preoccupied by Hylaeus quadrimaculatus Schenck, 1853 [=Lasioglossum interruptum (Panzer, 1798) of Europe] so was replaced by Halictus macoupinensis Robertson 1895), designated by Cresson (1928), revealed that that name macoupinensis has been used incorrectly by most authors. The type series was composite, and records of L. macoupinense in earlier literature, including some of the type series, do not match the lectotype and are instead referable to L. birkmanni. The name L. macoupinense is still in use, but study of the lectotype shows it to be a senior synonym of the bee widely known (e.g., Mitchell 1960; Hurd 1979; Moure & Hurd 1987) as Evylaeus divergens (Lovell) (see below). Observed nesting in the ground in Alabama (see Gibbs et al. 2013).

Lasioglossum (Hemihalictus) fedorense (Crawford 1906) County records: St. Joseph. Notes. Lasioglossum fedorense has been collected previously from the Indiana Dunes National Lakeshore (Grundel et al. 2011), which at its closest point is less than 10 km from the Michigan border. Lasioglossum fedorense prefers sandy habitats (Gibbs et al. 2013), which are abundant in the western part of the LP. The single specimen female examined is from Tamarack Lake only 6 km from the Indiana border. Material examined. St. Joseph Co.: Tamarack Lake, 14 Jun. 1986 (1 ♀ UMMZ).

Lasioglossum (Hemihalictus) foxii (Robertson 1895) County records: Allegan, Antrim, Barry, Benzie, Berrien, Cheboygan, Clinton, Crawford, Dickinson, Ingham, Kalamazoo, Kalkaska, Lake, Leelanau, Livingston, Manistee, Montcalm, Montmorency, Oceana, Oscoda, Otsego, Ottawa, Van Buren, Washtenaw. Notes. Lasioglossum foxii is known to nest in aggregations (Knerer & Atwood 1962).

Lasioglossum (Hemihalictus) inconditum (Cockerell 1916) County records: Alger, Chippewa, Dickinson, Iron, Keweenaw, Marquette, Schoolcraft. Notes. A ground-nesting species that is presumably solitary (Gibbs et al. 2013). Until recently, North American material of this species was identified as L. rufitarse (Zetterstedti 1838), a Palaearctic species.

Lasioglossum (Hemihalictus) lustrans (Cockerell 1897) County records: Allegan, Clare, Clinton, Delta, Dickinson, Kalamazoo, Kalkaska, Kent, Lake, Leelanau, Muskegon, Newaygo, Osceola, Ottawa, St. Joseph, Van Buren. Notes. This solitary species is best known from the southeastern and south-central United States, but it extends north locally to Michigan’s Upper Peninsula. It is relatively common along the east coast of Lake Michigan where it visits a variety of yellow composite flowers. Southern populations of L. lustrans have been reported to be very narrow oligoleges of Pyrrhopappus (Asteraceae), only occasionally visiting other members of the tribe Cichoriae (Daly 1961; Michener 1947b). Northern populations, however, occur outside the range of Pyrrhopappus (Estes & Thorp 1975) and regularly visit a broader array of species in the tribe. Arduser (2010) and Grundel et al. (2011) both report L. lustrans visiting Krigia species in the north. Lasioglossum lustrans has also been observed visiting other genera of Cichoriae in Michigan, including Crepis and Taraxacum (J.G. pers. obs.). Lasioglossum lustrans is recognizable by the absence of vein 1rs-m through most of its range, but a small proportion of northern specimens from Wisconsin and Michigan have this vein present (Gibbs 2010a; Gibbs et al. 2013).



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Lasioglossum (Hemihalictus) macoupinense (Robertson 1895) County records: Alger, Allegan, Antrim, Barry, Cheboygan, Clare, Clinton, Delta, Dickinson, Emmet, Gratiot, Huron, Ingham, Iron, Jackson, Kalamazoo, Mackinac, Manistee, Midland, Montcalm, Oakland, Oceana, Osceola, Ottawa, St. Clair, Van Buren, Wayne. Notes. The female lectotype of Halictus quadrimaculatus (see Cresson 1928) in the ANSP belongs to the species most often referred to as L. divergens in the literature (often in combination with either Halictus or Evylaeus) (Gibbs et al. 2013) although the name macoupinensis (a replacement name for H. quadrimaculatus) has been most often used to refer to L. birkmanni (see above).

Lasioglossum (Hemihalictus) nelumbonis (Robertson 1890) County records: Allegan, Barry, Calhoun, Dickinson, Ionia, Jackson, Kalamazoo, Lapeer, Livingston, Midland, Osceola, St. Joseph, Van Buren. Notes. Lasioglossum nelumbonis is associated with wetlands and has been recorded as an oligolege of Nymphaeaceae (Gibbs et al. 2013; Lippok et al. 2000).

Lasioglossum (Hemihalictus) pectorale (Smith 1853) County records: Allegan, Antrim, Barry, Berrien, Branch, Calhoun, Cass, Clinton, Eaton, Emmet, Grand Traverse, Gratiot, Hillsdale, Huron, Ingham, Ionia, Jackson, Kalamazoo, Kalkaska, Lake, Lapeer, Leelanau, Livingston, Manistee, Mason, Mecosta, Midland, Missaukee, Montcalm, Montmorency, Muskegon, Oakland, Oceana, Oscoda, Ottawa, Saginaw, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw, Wexford. Notes. A common and abundant species.

Lasioglossum (Hemihalictus) swenki (Crawford 1906) County records: Kalamazoo, Ottawa. Notes. This primarily North Central species apparently prefers sandy areas (e.g., Grundel et al. 2010). Gibbs et al. (2013) first reported the occurrence of this species from a single location near Lake Michigan. A closely related species, L. fedorense, is newly recorded from Michigan (see above). Material examined. Kalamazoo Co.: (no locality) (1 ♀ KCIC); Ottawa Co.: Grand Haven, 30 Aug. 1961, N. Aylesworth (1 ♀ MSUC).

Subgenus Lasioglossum Curtis s. s. Revision: McGinley (1986). Behavior: All members of Lasioglossum s. s. are expected to be solitary, ground-nesters (Gibbs et al. 2012b; Pesenko et al. 2000).

Lasioglossum (Lasioglossum) acuminatum McGinley 1986 (forbesii group) County records: Alger, Allegan, Antrim, Barry, Berrien, Cheboygan, Crawford, Delta, Dickinson, Emmet, Hillsdale, Huron, Ingham, Kalkaska, Lake, Leelanau, Mackinac, Manistee, Marquette, Mecosta, Menominee, Midland, Missaukee, Montmorency, Muskegon, Newaygo, Oakland, Oceana, Ogemaw, Oscoda, Otsego, Ottawa, Presque Isle, Roscommon, Saginaw, Schoolcraft, Tuscola, Van Buren, Wexford.


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Lasioglossum (Lasioglossum) athabascense (Sandhouse 1933) County records: Allegan, Antrim, Cheboygan, Chippewa, Dickinson, Emmet, Houghton, Keweenaw, Mackinac, Marquette, Ontonagon, Wayne.

Lasioglossum (Lasioglossum) coriaceum (Smith 1853) County records: Alger, Allegan, Antrim, Baraga, Barry, Bay, Benzie, Berrien, Branch, Calhoun, Cass, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Crawford, Dickinson, Eaton, Emmet, Genesee, Gladwin, Grand Traverse, Gratiot, Hillsdale, Houghton, Ingham, Ionia, Jackson, Kalamazoo, Kalkaska, Lake, Leelanau, Lenawee, Livingston, Mackinac, Manistee, Marquette, Mecosta, Midland, Missaukee, Monroe, Montcalm, Oakland, Oceana, Oscoda, Otsego, Ottawa, Presque Isle, Roscommon, Saginaw, Schoolcraft, St. Clair, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne.

Lasioglossum (Lasioglossum) forbesii (Robertson 1890) (forbesii group) County records: Allegan, Berrien, Clinton, Ingham, Ottawa, Tuscola. Notes. The rarest member of the subgenus in the state. However, before McGinley’s (1986) revision, this name was applied broadly to multiple species, e.g., L. paraforbesii McGinley, now recognized as distinct.

Lasioglossum (Lasioglossum) fuscipenne (Smith 1853) County records: Allegan, Barry, Cheboygan, Kalamazoo, Muskegon, Oakland, Ottawa, Van Buren.

Lasioglossum (Lasioglossum) paraforbesii McGinley 1986 (forbesii group) County records: Allegan, Barry, Berrien, Cass, Cheboygan, Chippewa, Clinton, Eaton, Grand Traverse, Huron, Ingham, Jackson, Kalamazoo, Kalkaska, Lake, Leelanau, Lenawee, Mackinac, Mecosta, Midland, Montcalm, Newaygo, Oakland, Oceana, Oscoda, Tuscola, Van Buren.

Subgenus Leuchalictus Warncke Revision: McGinley (1986). Biology. Both representatives of the subgenus in North America are believed to be accidental introductions (Ebmer 2011; Giles & Ascher 2006; Zayed et al. 2007). The record of a third member of the subgenus, L. dynastes (Bingham) in Ohio (Ebmer 2011) is incorrect and due to a poorly labeled specimens. The collector, an Ohio resident, actually found the specimens in the Himalayan Region of India: Ranichauri, District of Tehri Garhwal, State of Uttarakhand (R. Williams, pers. comm.). Both L. (Leuchalictus) below are solitary, ground-nesters (Pesenko et al. 2000).

Lasioglossum (Leuchalictus) leucozonium (Schrank 1781) County records: Alger, Allegan, Antrim, Barry, Berrien, Calhoun, Cass, Chippewa, Clare, Clinton, Dickinson, Eaton, Genesee, Grand Traverse, Gratiot, Houghton, Huron, Ingham, Iosco, Iron, Jackson, Kalamazoo, Kalkaska, Keweenaw, Lake, Lapeer, Leelanau, Livingston, Luce, Mackinac, Marquette, Montcalm, Muskegon, Newaygo,



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Oakland, Oceana, Ontonagon, Oscoda, Ottawa, Presque Isle, Saginaw, Schoolcraft, St. Joseph, Tuscola, Van Buren, Wexford. Notes. A very common exotic species (Zayed et al. 2007). It is a solitary ground-nester (Atwood 1933; Knerer 1969; Knerer & Atwood 1966; Pesenko et al. 2000).

Lasioglossum (Leuchalictus) zonulum (Smith 1848) County records: Alger, Baraga, Charlevoix, Cheboygan, Chippewa, Clinton, Delta, Dickinson, Eaton, Emmet, Genesee, Gogebic, Gratiot, Houghton, Huron, Iron, Keweenaw, Lapeer, Leelanau, Livingston, Mackinac, Marquette, Mason, Mecosta, Menominee, Midland, Missaukee, Montcalm, Muskegon, Oceana, Ontonagon, Otsego, Roscommon, Saginaw, Schoolcraft, St. Clair, St. Joseph, Tuscola. Notes. An exotic species (Giles & Ascher 2006). Closely related to L. leucozonium, with similar nesting habits (Atwood 1933; Pesenko et al. 2000).

Subgenus Sphecodogastra Ashmead Revision: Gibbs et al. (2013). Biology. A biologically diverse subgenus including social and solitary nesters. Most members of the subgenus sensu Gibbs et al. (2013) are polylectic, but the type species and its relatives, i.e. Sphecodogastra sensu stricto, are Onagraceae specialists (McGinley 2003).

Lasioglossum (Sphecodogastra) comagenense (Knerer & Atwood 1964) (fulvicorne group) County records: Antrim, Dickinson, Gogebic, Keweenaw, Marquette. Notes. A socially plastic ground-nester (Packer et al. 1989a, Packer et al. 1989b, Batra 1990b).

Lasioglossum (Sphecodogastra) oenotherae (Stevens 1920) (lusorium group) County records: Ingham, Kent, Livingston, Washtenaw. Notes. The nesting biology of Ontario populations have been studied in detail (Knerer & Mackay 1969). Records of this species are largely absent from the Midwest, although the species occurs widely from the Central Plains to Maritime Canada (Gibbs et al. 2013; McGinley 2003). Multiple attempts by JG were made to capture female L. oenotherae in Ingham County from Oenothera spp. (Onagraceae) in the early and late hours of the day without success, but two males were obtained from other flowers during the middle of the day when females would not typically be active (McGinley 2003). Material examined. Ingham Co.: Lake Lansing, Haslett, N42.749 W84.4, 20 Aug. 2013 (1 ♂ JBWM); Harrison Park, East Lansing, N42.754 W84.486, 25 Aug. 2013 (1 ♂ JBWM); Livingston Co.: E. S. George Reserve, 3 Jul. 19(5/9)5, Oenothera fruticosa (as tetragona) (UMMZ). Kent Co.: Grand Rapids, 19 Jun. 2014, Oenothera biennis L. (♀ http://bugguide.net/node/view/940949).

Lasioglossum (Sphecodogastra) quebecense (Crawford 1907) (fulvicorne group) County records: Allegan, Cass, Cheboygan, Clare, Crawford, Dickinson, Ingham, Keweenaw, Leelanau, Mackinac, Marquette, Montmorency, Osceola, Oscoda, Ottawa, Schoolcraft, Van Buren, Wexford. Notes. Lasioglossum quebecense has been recorded as frequent visitor of Vaccinium (Boulanger et al. 1967) and has potential value as pollinator of spring crops.


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Lasioglossum (Sphecodogastra) seillean Gibbs & Packer 2013 (fulvicorne group) County records: Alger, Keweenaw. Notes. A boreal species first described in Gibbs et al. (2013). Michigan specimens are all females, but identification is most reliable by examination of male genitalia due to close similarity to L. quebecense.

Lasioglossum (Sphecodogastra) texanum (Cresson 1872) (texanum group) County records: Clinton, Ingham, Kalamazoo, Kent, Livingston. Notes. A solitary Oenothera (Onagraceae) specialist that flies at moonlight (Kerfoot 1967a; b; McGinley 2003). Lasioglossum texanum is rarely encountered in the state.

Lasioglossum (Sphecodogastra) truncatum (Robertson 1901) (calceatum group) County records: Grand Traverse, Lake, Leelanau.

Genus Sphecodes Latreille Taxonomy: Mitchell (1960). Sphecodes is in particular need of revision. Biology. All members of the genus are cleptoparasites or social parasites, usually of other Halictinae but with exceptions, see below. Definitive host records are few and information on host specificity is generally lacking.

Sphecodes antennariae Robertson 1891 (mandibularis group) County records: Kalamazoo, Washtenaw.

Sphecodes aroniae Mitchell 1960 (ranunculi group) (New state record) County records: Midland, Muskegon. Notes. Sphecodes aroniae is very similar to S. ranunculi, which was previously known from the state, and the former has been under-recorded in published studies (Goldstein & Ascher 2016). Females recorded below were distinguished from S. ranunculi (Robertson) by the flat distinct genal carina and the male by the greatly expanded angular mesotibia. Both sexes are also recognizable by the flat procoxa. Material examined. Midland Co.: (no locality), 5–11 Jun. 1961, R.R. Dreisbach (1 ♂ MSUC); Muskegon Co.: Tyler Rd., W of Dalton, 17 km N of Muskegon, N43.319 W86.264, 8 Jun. 2013 (2 ♀ JBWM, MSUC: RI), 5 Jun. 2014 (1 ♀ JBWM).

Sphecodes atlantis Mitchell 1956 (mandibularis group) County records: Allegan, Berrien, Clare, Ingham, Leelanau, Mecosta, Midland, Muskegon, Otsego, Washtenaw. Notes. A known cleptoparasite of Lasioglossum pilosum (see Michener 1978).

Sphecodes autumnalis Mitchell 1956 (mandibularis group) County records: Alger, Gladwin, Midland, Shiawassee. THE BEES OF MICHIGAN


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Notes. A cleptoparasite of Perdita octomaculata (see Michener 1978).

Sphecodes banksii Lovell 1909 (mandibularis group) County records: Allegan, Ottawa. Notes. Local in occurrence and perhaps sand associated, possibly a cleptoparasite of L. vierecki.

Sphecodes clematidis Robertson 1897 (dichrous group) County records: Ingham, Ionia, Otsego, Shiawassee.

Sphecodes confertus Say 1837 (confertus group) County records: Allegan, Cheboygan, Clinton, Ingham, Livingston, Muskegon, Oceana, Ottawa, Shiawassee, Van Buren, Wayne.

Sphecodes coronus Mitchell 1956 (mandibularis group) Holotype. ♂ USA, North Carolina, Wake Co.: 16 Jun. 1955, on Daucus carota, T.B. Mitchell (NMNH: 75238). = Sphecodes carolinus Mitchell, 1956 Holotype. ♀ USA, North Carolina, Grandfather Mt., 24 Jun. 1954, on Rubus, T.B. Mitchell (NMNH: 76237).

County records: Alger, Arenac, Charlevoix, Dickinson, Iosco, Keweenaw, Muskegon. Notes. The synonymy of these two names has not been well established in the literature, although S. coronus has been used to include both sexes in faunal studies (Goldstein & Ascher 2016) and DNA barcode data support the synonymy (Sheffield et al. 2009; Sheffield & Perron 2014).

Sphecodes cressonii (Robertson 1903) (mandibularis group) County records: Allegan, Barry, Berrien, Calhoun, Clinton, Keweenaw, Livingston, Montcalm, Ottawa, Washtenaw.

Sphecodes davisii Robertson 1897 (mandibularis group) Holotype. ♂ USA, Michigan (?) (Michigan Agricultural College), 17 Apr. 1886 (INHS 179717) = Sphecodes persimilis Lovell & Cockerell 1907

County records: Alger, Antrim, Calhoun, Ingham, Kent, Leelanau, Livingston, Oakland, Ottawa, Tuscola, Van Buren. Notes. This is a common and widespread species, originally described from a single male specimen. It is considered a senior synonym of S. persimilis, which was described from the female. Mitchell (1960) pointed out the possible synonymy and Abrams & Eickwort (1980) considered this probable. Mike Arduser and JSA have each applied this synonymy, but its establishment was relatively obscure since it was not done as part of a published revision but rather has appeared online and implicitly in faunal checklists (Arduser 2010; Ascher et al. 2014; Goldstein & Ascher 2016). Sphecodes davisii is a cleptoparasite of Agapostemon virescens (Fabricius) and A. sericeus (Förster) (Abrams & Eickwort 1980).


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Sphecodes dichrous Smith 1853 (dichrous group) Syntype. ♀ ♂ USA, E. Doubleday (NHM). = Sphecodes knetschi Cockerell 1898 (New synonymy) Syntype. ♀ ♂ USA, Illinois, Terra Cotta, 25 Aug. 1897, 5 Sep. 1897, goldenrod, R. Knetsch (NMNH).

County records: Clinton, Ingham, Jackson, Livingston, Muskegon, Washtenaw. Notes. Mitchell (1960) records this species under both names. Although the synonymy, first recognized by Mike Arduser, has been applied implicitly, it has not been published formally.

Sphecodes fattigi Mitchell 1956 (mandibularis group) Holotype. ♂ USA, Florida, Gainesville, 25 Apr. 1921, P. W. Fattig (CUIC). = Sphecodes prostygius Mitchell, 1960 (New synonymy) Holotype. ♀ USA, New Hampshire, Meredith Center, 3 Aug. 1957, Solidago, R. A Morse (CUIC).

County records: Alger, Clinton, Lake, Missaukee, Van Buren. Notes. This unpublished synonymy by M. Arduser has been applied in databases and implicitly in other studies but has not been formally stated.

Sphecodes heraclei heraclei Robertson 1897 (dichrous group) County records: Ionia, Livingston. Notes. Recorded by Evans (1986) from the E.S. George Reserve, Livingston County, and recently recollected by T. Wood (pers. comm.) at the Portland State Game Area, Ionia County.

Sphecodes illinoensis (Robertson 1903) (mandibularis group) County records: Allegan, Leelanau, Livingston, Oceana, Ottawa, Roscommon, Van Buren, Washtenaw.

Sphecodes johnsonii Lovell 1909 (mandibularis group) County records: Cheboygan, Clinton, Ingham, Van Buren.

Sphecodes levis Lovell & Cockerell 1907 (mandibularis group) County records: Van Buren. Notes. Mitchell (1960) record. Two female specimens from Van Buren Co. at JBWM. This species has been recorded a a cleptoparasite of Lasioglossum quebecense (see Michener 1978).

Sphecodes mandibularis Cresson 1872 (mandibularis group) Lectotype. ♀ USA, Texas, Bosque Co.: G.W. Belfrage (ANSP: 2133). Designated by Cresson (1916). = Sphecodes stygius Robertson 1893 Lectotype. ♀ USA, Illinois, Carlinville, 12 Aug. 1891, C.A. Robertson (INHS: 11944). Designated by W.E. LaBerge in Webb (1980).



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County records: Allegan, Antrim, Barry, Benzie, Charlevoix, Clinton, Kalamazoo, Leelanau, Livingston, Montcalm, Van Buren. Notes. The synonymy of these two names, proposed by M. Arduser, has not been well established in the literature, although S. mandibularis has been used in faunal studies for the species (Wolf & Ascher 2009, Ascher et al. 2014, Goldstein & Ascher 2016). Mitchell (1960) treated the species as S. stygius rather than S. mandibularis because the female lectotype of the latter from Bosque County, Texas was extralimital.

Sphecodes minor Robertson 1898 (dichrous group) County records: Barry, Ingham, Shiawassee. Notes. Recorded as a cleptoparasite of Lasioglossum cinctipes, with two separate broods that attack nests in spring and summer during the solitary and social phases of the host (Knerer & Atwood 1967).

Sphecodes pimpinellae Robertson 1900 (mandibularis group) Holotype. ♀ USA, Illinois, Carlinville, C. Robertson (INHS) = Sphecodes wheeleri Mitchell, 1956 (New synonymy) Holotype. ♀ USA, North Carolina, Marion, 18 Jun. 1941, Leucanthemum, T. B. Mitchell (NMNH).

County records: Berrien, Jackson. Notes. Sphecodes pimpinellae, a cleptoparasite of Augochlorella (Ordway 1964), is newly recorded from southern Michigan (Lettow et al. in review). The synonymy, first suggested by Ordway (1964) and endorsed in an unpublished study by Mike Arduser, is provisionally accepted here but cannot be considered well established due to incomplete understanding of observed morphological variation. The names S. pimpinellae and S. wheeleri have been applied to females with and without a subapical tooth on the mandible, respectively. Sphecodes pimpinellae has been infrequently recorded across the east and Midwest, while S. wheeleri was described from a few areas in New York, Connecticut and North Carolina (Mitchell 1956). The Michigan female specimen lacks the subapical tooth, so should the synonymy be rejected it would be S. wheeleri. Material examined. Berrien Co.: SWMREC, 19 Jul. 2016 (1 ♂ MSUC:RI); Jackson Co.: Clark Lake, MacCready Res., N42°7ʹ W84°24ʹ, 4 Jun. 2012, M.C. Lettow (1 ♀ MSUC [at JBWM]).

Sphecodes prosphorus Lovell & Cockerell 1907 (dichrous group) County records: Crawford, Ionia, Presque Isle.

Sphecodes pycnanthemi Robertson 1897 (mandibularis group) (New state record) County records: Hillsdale. Notes. Sphecodes pycnanthemi is a poorly known species, described from Carlinville, Illinois. It has also been recorded from Indiana (Jean 2010). A New York specimen was compared directly to the lectotype. A recently collected specimen from Michigan matches the New York material. Material examined. Hillsdale Co.: Pittsford State Game Area, N41.866 W84.522, 3 Jun. 2017, T.J. Wood (TJWC). Additional material examined. NEW YORK: Tompkins Co.: Ithaca, Six-mile Creek, N42.432 W76.484, 19 May 2012, J. Gibbs (JBWM—New state record).

Sphecodes ranunculi Robertson 1897 (ranunculi group) County records: Alger, Ingham, Livingston, Muskegon, Shiawassee, Van Buren.


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Sphecodes townesi Mitchell 1956 (mandibularis group) County records: Ingham, Iosco.

Nomiinae Dieunomiini Genus Dieunomia Cockerell Revision: Blair (1935).

Subgenus Dieunomia Cockerell s. s. Dieunomia (Dieunomia) heteropoda heteropoda (Say 1824) County records: Allegan, Berrien, Kalamazoo, Ottawa, Van Buren. Notes. Dieunomia heteropoda was recently documented from the southwestern portion of the state based on specimens collected since 2003 (Gibbs et al. 2014). These records are near the northeastern edge of its range. Although the large size and dark color of this bee makes it easy to recognize, the earliest known record in the state is only from 2003, suggesting that there has been a genuine range expansion. However, Montgomery (1957) recorded it from Starke County in northwest Indiana. Dieunomia heteropoda can be collected on large Asteraceae, such as Helianthus, Coreopsis, Rudbeckia and Centaurea. Females have also been observed collecting nectar from Polygonum (Polygonaceae). Nests and mating behavior have been described from western states (Cane 1991; Cross & Bohart 1960; Wcislo 1993; Wcislo & Buchmann 1995). Material examined. Allegan Co.: 4.5 km south of Pullman, 17 Jul. 2013 (MSUC: RI), 15 Aug. 2013, J. Gibbs (1 ♀ JBWM), 11 Jul. 2014 (2 ♀ JBWM); Allegan State Game Area, T2N R14W Sec. 18, 2 Aug. 2003 (1 ♀ MSUC); Berrien Co.: 4 km west of Berrien Springs (1 ♀ MSUC: RI); Kalamazoo Co.: Oshtemo Twp., N41.172, W85.345, 13 Jul. 2011, D.A. Landis (1 ♂ MSUC: RI); Ottawa Co.: Grand River Park, 27 Jul. 2008 (1 sex not recorded MVWC); Van Buren Co.: 4 km NNW of Almena, 18 Jul. 2011 Coreopsis (1 ♂), 20 Jul. 2011 (1 ♀), 8 Aug. 2011 Helianthus (1 ♀ 1 ♂), 10 Jul. 2012 Rudbeckia (5 ♀ 1 ♂) (MSUC: RI), 4 km NNW of Almena, 14 Aug. 2014, J. Gibbs (1 ♀ JBWM).

Rophitinae Genus Dufourea Lepeletier Taxonomy: Mitchell (1960); Dumesh & Sheffield (2012); Gibbs et al. (2014). Behavior. Dufourea are solitary ground-nesters that primarily specialize on particular floral hosts.

Dufourea harveyi (Cresson 1878) County records: Kalamazoo. Notes. Dufourea harveyi, a specialist on Potentilla, was recently resurrected from synonymy with D. fimbriata fimbriata (Cresson) by Gibbs et al. (2014). Three specimens deposited at MSUC collected between 1960 and 2009 were recently documented from Barry, Jackson and Kalamazoo Counties in a review of Michigan Dufourea (Gibbs et al. 2014). The species had not previously been recorded from the eastern United States, although a specimen



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from northern Ontario had been reported (Dumesh & Sheffield 2012; as D. fimbriata). Dufourea fimbriata is fully confirmed only from its type locality of Colorado and from California where subspecies sierrae (Michener) occurs. The four species of Dufourea known from Michigan were reviewed by Gibbs et al. (2014).

Dufourea maura (Cresson 1878) County records: Keweenaw. Notes. Specialist on Campanula (Campanulaceae). Only known in the Eastern United States from Isle Royale (Arduser 1986).

Dufourea monardae (Viereck 1924) County records: Calhoun, Dickinson, Huron, Ingham, Ionia, Kalamazoo, Livingston, Oakland, Oceana. Notes. Specialist on Lamiaceae, in particular Monarda fistulosa L.

Dufourea novaeangliae (Robertson 1897) County records: Allegan, Ingham, Wexford. Notes. Solitary, ground-nester and a specialist on Pontederia cordata L. (Pontederiaceae) studied in detail in New York (Eickwort et al. 1986).

MEGACHILIDAE Megachilinae Anthidiini Genus Anthidiellum Cockerell Subgenus Loyolanthidium Urban Taxonomy: Mitchell (1962); Urban (2001). Biology. Nests of resin are made on the outer surface of plants or rocks (Baker et al. 1985; Grigarick & Stange 1968).

Anthidiellum (Loyolanthidium) notatum notatum (Latreille 1809) (New state record) County records: Berrien, Cass, Ingham, Livingston. Notes. This species has not been recorded previously from Michigan although it was expected to be found here based on its wide distribution including southern Ontario (J.G. pers. obs.) and northern Indiana (Grundel et al. 2011). Several specimens were collected recently in the southern LP and a single historical specimen has been identified. Recent collections from two sites in different counties include males of two subspecies: A. notatum notatum and A. notatum boreale (Robertson) (see below). Anthidiellum boreale is distinguished from the nominal subspecies (widespread in the Eastern United States) by the truncated paraocular maculations, absence of axillar maculations and presence of a medial longitudinal carina on T6 and T7 of males (Mitchell 1962). Nest from North Carolina described by Baker et al. (1985). Material examined. Berrien Co.: 3.5 km ENE Niles, 9 Aug. 2011, Trifolium hybridum (1 ♂ MSUC: RI); SWMREC, 21 Aug. 2015, Lespedeza hirta (1 ♀ MSUC: RI); Cass Co.: Edward Lowe Foundation, N41.945


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W86.01, 26 Aug. 2013 (1 ♂ JBWM); Edward Lowe Foundation, N41.968 W85.993, 27 Aug. 2013 (1 ♂ JBWM), (2 ♀ Cirsium arvense (L.) Scop., 1 ♀ Cichorium intybus L., 1 ♂ Securigera varia L. (Lassen) MSUC: RI); Ingham Co.: Michigan State University, south campus/horticultural garden, N42.721 W84.473, Lotus corniculatus L. 21 Jul. 2014 (1 ♂), 4 Aug. 2013 (1 ♂), 23 Aug. 2013 (1 ♂), (JBWM); Livingston Co.: E. S. George Reserve, 11 Aug. 1975 (F.C. Evans), Monarda fistulosa (1 ♂ UMMZ). Additional material. CANADA: Ontario: North York Dist.: York University, N43.775 W79.505 Lotus corniculatus, 19 Jul. 2008 (1♂), 11 Aug. 2008 (1♂), (JBWM).

Anthidiellum (Loyolanthidium) notatum boreale (Robertson 1902) (New state record) County records: Cass, Ingham. Notes. The holotype male is from Carlinville, Illinois (INHS). Mitchell (1962) lists Illinois in the distribution of both subspecies known from Michigan. The collection of males together at the same site suggests the subspecies are syntopic. Additional taxonomic study is needed to determine if A. n. boreale warrants subspecific or specific status. Urban (2001) described and reviewed the subgenus (as a genus), but did not examine any specimens of A. n. boreale, nor did she indicate its current taxonomic status, whereas she elevated the western A. gilense (Cockerell 1897) and A. robertsoni (Cockerell 1904) to specific rank. Material examined. Cass Co.: Edward Lowe Foundation, N41.968 W85.993, 27 Aug. 2013 (1 ♂ JBWM; MSUC: RI); Ingham Co.: Michigan State University, arboretum, N42.721 W84.475, Lotus corniculatus L., 11 Aug. 2013 (1 ♂ JBWM).

Genus Anthidium Fabricius Revision: Gonzalez & Griswold (2013). Behavior. Wool-carder bees use plant hairs to make nests in pre-existing cavities or the ground.

Subgenus Anthidium Fabricius s. s. Anthidium (Anthidium) manicatum manicatum (Linnaeus 1758) County records: Allegan, Bay, Ingham, Jackson, Kent, Leelanau, Lenawee, Mecosta, Newaygo, Oakland, Ottawa, Saginaw, Van Buren, Washtenaw, Wayne. Notes. Exotic. First recorded in North America from New York (Jaycox 1967), this cavity-nesting species has spread rapidly across North America (Gibbs & Sheffield 2009). See also O'Brien et al. (2013).

Anthidium (Anthidium) psoraleae Robertson 1902 County records: Barry, Berrien, Lake, Livingston, Washtenaw. Notes. This species is found in the central USA between the Rockies and Mississippi River, from southern Ontario south to Oklahoma. It has been collected in southern Michigan, including Barry, Berrien, Livingston and Washtenaw Counties (Gonzalez & Griswold 2013; O’Brien et al. 2013). New site records are provided here that extend its range northward to Lake County. Material examined. Barry Co.: 24 Jun. 1956, Trifolium, R.W. Hodges (1 ♀ MSUC); Lake Co.: Hillsdale College Bio. Sta., Luther, 27–28 Jun. 2015, J. Gibbs (1 ♀ 2 ♂ JBWM).

Anthidium (Anthidium) tenuiflorae Cockerell 1907 (New state record and new for Eastern USA) (Fig. 17A) County records: Keweenaw (Isle Royale). THE BEES OF MICHIGAN


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Notes. Gonzalez & Griswold (2013) report this primarily western species from as far east as Minnesota in the US and Saskatchewan in Canada. The Minnesota records are from near Basswood Lake, which is less than 200 km west of Isle Royale, Michigan where the bee was collected in 1957 and also recently by S. Wilson (pers. comm.). A single bee from Manitoba has been examined from Garson Quarry Station, 1 July 1996 (JBWM) suggesting that it should also occur in western Ontario, Canada. Notes on the nest were made by Hicks (1926) in Colorado. Material examined. Keweenaw Co.: Isle Royale, 3 Jul. 1957, R.W. Hodges (1 ♀ MSUC); Isle Royale, 13 Jul. 1957, R.W. Hodges (1 ♀ MSUC); Isle Royale, 27 Jul. 1957, R.W. Hodges (1 ♀ MSUC).

Subgenus Proanthidium Friese Anthidium (Proanthidium) oblongatum oblongatum (Illiger 1806) County records: Allegan, Barry, Berrien, Clinton, Ingham, Ionia, Kent, Ottawa, Van Buren, Washtenaw, Wayne. Notes. Exotic. First recorded in the northeast by Hoebeke & Wheeler (1999), it had reached Illinois by 2008 (Tonietto & Ascher 2009). See also O'Brien et al. (2013).

FIGURE 17. Anthidiine bees from Michigan (not to scale). A) Anthidium (Anthidium) tenuiflorae Cockerell female, metasomal terga. B) Stelis (Stelis) subemarginata Cresson female from Michigan, lateral habitus. Courtesy of Sam Droege, Patuxent Wildlife Research Center.

Genus Dianthidium Cockerell Subgenus Dianthidium Cockerell s. s. Taxonomy: Schwarz 1926; Mitchell (1962). Biology: O’Brien (2007).

Dianthidium (Dianthidium) simile (Cresson 1864) County records: Berrien, Cass, Charlevoix, Cheboygan, Grand Traverse, Huron, Iosco, Kalkaska, Lake, Leelanau, Mackinac, Manistee, Mason, Mecosta, Midland, Montcalm, Montmorency, Oceana, Oscoda, Otsego, Ottawa, Presque Isle, Shiawassee. Notes. A resin bee with underground nests, studied in detail in Michigan by O’Brien (2007).


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Genus Stelis Panzer Taxonomy: Parker & Bohart (1979); Mitchell (1962). Biology: Stelis are all cleptoparasites of megachiline bees.

Subgenus Dolichostelis Parker & Bohart Revision: Parker & Bohart (1979).

Stelis (Dolichostelis) louisae Cockerell 1911 County records: Cass, Clinton, Ingham, Kalamazoo, Washtenaw, Wayne. Notes. Stelis louisae is a cleptoparasite of Megachile (Chelostomoides) including M. campanulae (Robertson) (Baker et al. 1985; Parker et al. 1987).

Subgenus Stelis Panzer s. s. Stelis (Stelis) coarctatus Crawford 1916 (coarctatus group) = Stelis (Microstelis) vernalis Mitchell, 1962: 43. New synonymy (by F. Parker and T. Griswold).

County records: Cheboygan, Kalamazoo, Livingston, Mason, Missaukee. Notes. Mitchell (1962) did not include S. coarctatus in his treatment of the eastern species. The holotype female, deposited at NMNH, is from Kansas. The holotype female of S. vernalis, also deposited at NMNH, is from North Carolina. This species has been recorded as a cleptoparasite of Heriades carinata Cresson (see Sheffield et al. 2008). It was reared from Heriades carinata nests in Michigan by Matthews (1965 as vernalis), but also from Hoplitis (Proteriades) shoshone (Parker) in Nevada (Parker 1976), suggesting a broad host range.

Stelis (Stelis) foederalis Smith 1854 (foederalis group) = Stelis michiganensis Mitchell, 1962: 45. (Michigan holotype; Fig. 18). New synonymy (by F. Parker and T. Griswold). Holotype. ♀ USA: Michigan: Luce Co.: 8 Jul. 1946, R.R. Dreisbach (NMNH 537077 on indefinite loan from NCSU).

County records: Baraga, Delta, Dickinson, Kalkaska, Keweenaw, Luce, Mackinac, Marquette, Montmorency, Schoolcraft. Notes. Recorded as a cleptoparasite of Osmia atriventris Cresson and Hoplitis spoliata (Provancher) (see Fye 1965; Medler 1967b as Chelynia).

Stelis (Stelis) labiata (Provancher 1888) (subemarginata group) County records: Dickinson, Iron. Notes. Recorded as acleptoparasite of Hoplitis species, including H. producta (Cresson) and H. spoliata (Medler 1961, 1967b; Medler & Lussenhop 1968).



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Stelis (Stelis) lateralis Cresson 1864 (lateralis group) County records: Antrim, Cass, Cheboygan, Dickinson, Eaton, Emmet, Gratiot, Isabella, Jackson, Saginaw, St. Joseph, Washtenaw. Notes. Recorded as a cleptoparasite of Hoplitis, including H. pilosifrons (Cresson), H. producta and H. spoliata (Graenicher 1905; Hicks 1926; Medler 1961, 1967b; Michener 1955; Rau 1928), recorded in earlier studies as sexmaculata Ashmead.

Stelis (Stelis) nitida Cresson 1878 (foederalis group) County records: Baraga, Keweenaw, Schoolcraft.

Stelis (Stelis) subemarginata Cresson 1878 (subemarginata group) (New state record) (Fig. 17B) County records: Keweenaw. Notes. A single specimen record confirmed by S. Droege and examination of photographs. Fye (1965) recorded this species from Black Sturgeon Lake, Ontario; approximately 125 km north of Isle Royale as a cleptoparasite of Hoplitis spoliata (see also Sheffield et al. 2008). Graenicher (1935) recorded it from a nest of Osmia simillima Smith from Milwaukee, Wisconsin on the shore of Lake Michigan, but his identification might apply to Stelis labiata (Medler & Lussenhop 1968). Recorded as a cleptoparasite of H. spoliata and O. simillima. Material examined (photograph). Keweenaw Co.: Isle Royale National Park, 48.1252 -88.5111, 14 Jul. 2013, S. Wilson (1 ♀ PWRC).

FIGURE 18. Stelis michiganensis Mitchell holotype female (= S. foederalis Smith). A) Lateral habitus. B) Face. Photographs provided with the permission of the National Museum of Natural History, Smithsonian Institution, 10th and Constitution Ave. N.W., Washington, DC 20560-0193. (http://www.nmnh.si.edu/).

Genus Trachusa Panzer Subgenus Heteranthidium Cockerell Taxonomy: Mitchell (1962); Snelling (1966a); Brooks & Griswold (1988).


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Trachusa (Heteranthidium) zebrata (Cresson 1872) (zebrata group) County records: Kalamazoo, Lake, Livingston. Notes. A prairie associated species, uncommonly observed in Michigan.

Megachilini Genus Coelioxys Latreille Taxonomy: Mitchell (1962, 1980); Baker (1975); Rocha Filho & Packer (2016). Biology. Coelioxys are primarily cleptoparasites of Megachile (Baker 1975; Baker et al. 1985; Graenicher 1905; Hurd 1979; Ivanochko 1979).

Subgenus Boreocoelioxys Mitchell Revision: Baker (1975).

Coelioxys (Boreocoelioxys) moestus Cresson 1864 County records: Arenac, Calhoun, Cheboygan, Dickinson, Huron, Iron, Kalamazoo, Keweenaw, Marquette, Mecosta, Midland, Montcalm, Oscoda, Washtenaw. Notes. Hosts include Megachile frigida Smith and M. texana Cresson (Pengelly 1955), M. centuncularis (L.) (Medler 1959; Sheffield et al. 2008), M. lapponica Thomson (Peck & Bolton 1946), M. relativa Cresson (Medler & Koerber 1958; Medler & Lussenhop 1968; Sheffield et al. 2008; Strickler & Scriber 1994), and possibly M. rotundata (Fab.) (Hobbs 1968) (see C. funerarius Smith entry below), M. melanophaea Smith (Pengelly 1955) and M. pusilla (Baker 1975).

Coelioxys (Boreocoelioxys) octodentatus Say 1824 County records: Berrien, Cheboygan, Clinton, Huron, Ingham, Kalamazoo, Kalkaska, Lenawee, Livingston, Midland, Oakland, Shiawassee, Van Buren, Washtenaw, Wayne. Notes. Hosts include Megachile brevis Say (Baker 1971; Hicks 1926; Michener 1953; Pengelly 1955), M. mendica Cresson (Medler 1965), M. centuncularis (Medler & Lussenhop 1968; Pengelly 1955), M. perihirta Cockerell and M. rotundata (Baker 1975), and M. texana and possibly M. frigida and M. melanophaea (Pengelly 1955).

Coelioxys (Boreocoelioxys) porterae Cockerell 1900 County records: Alger, Alpena, Cheboygan, Clinton, Delta, Emmet, Hillsdale, Huron, Iron, Kalamazoo, Keweenaw, Mackinac, Marquette, Mecosta, Midland, Muskegon, St. Joseph. Notes. Hosts include Megachile frigida (Baker 1975) and M. relativa (Mitchell 1962; Sheffield et al. 2008).

Coelioxys (Boreocoelioxys) rufitarsis Smith 1854 County records: Alger, Bay, Cheboygan, Clinton, Crawford, Dickinson, Gladwin, Huron, Ingham, Isabella, Jackson, Kent, Keweenaw, Livingston, Mackinac, Menominee, Midland, Oakland, Ogemaw, Ontonagon, Oscoda, Otsego, Saginaw, Sanilac, Shiawassee, Washtenaw, Wayne.



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Notes. Host records include Megachile latimanus Say and M. melanophaea (Graenicher 1905, 1935; Pengelly 1955), M. montivaga Cresson (Hicks 1926), M. perihirta (Baker 1975), and M. texana (Pengelly 1955). Two spellings were used by Smith (1854) in the original publication describing this species, Coelioxys rufitarsus in the heading of the species description (pg. 271) and C. rufitarsis in the appendix (pg. 460). No subsequent uses of either spelling by Smith are known to us. Dalla Torre (1896) cited only the spelling C. rufitarsis, incorrectly attributing this to page 271 in Smith (1854) and incorrectly attributing use of this spelling to Cresson (1864) and Provancher (1882), whereas those authors had used rufitarsus. Robertson (1897), Lovell & Cockerell (1907), and Hurd (1979) all referenced the original spelling, but adopted C. rufitarsis, although none noted the source of this spelling, i.e. they did not cite the page number in question from Smith’s appendix. Crawford (1914) and Hobbs (1956) used both spellings apparently in error. Thus, none of these authors satisfies the requirements of first reviser (see ICZN article 24.2.1). Moure et al. (2007), however, cited both original spellings correctly and adopted rufitarsis, establishing this spelling as valid. The spelling rufitarsis is in prevailing usage (see ICZN article 33.2.3.1), i.e. has been used by a substantial majority of authors (Baker 1975; Beaulne 1942; Bizecki Robson 2013; Bried & Dillon 2012; Cockerell 1900, 1925; Colla et al. 2009; Crawford 1914; Dalla Torre 1896; Donovall & VanEngelsdorp 2010; Gardner & Spivak 2014; Genaro 2001; Graenicher 1927a; Grixti & Packer 2006; Hobbs 1956; Hurd 1979; Jean 2010; Kuhlman & Burrows 2017; Leavengood & Serrano 2005; Linsley 1951; MacKay & Knerer 1979; Mitchell 1962; Neff & Simpson 1991; Onuferko et al. 2015; Pearson 1933; Peck & Bolton 1946; Richards et al. 2011; Robertson 1929; Rocha Filho & Packer 2016; Rozen & Kamel 2006; Scott et al. 2011; Shapiro et al. 2014; Sheffield et al. 2009; Stephen & Rao 2007; Wagner et al. 2014; Wojcik et al. 2008; Wolf & Ascher 2009). While only a limited number of authors have used rufitarsus (Blake et al. 2010; Cockerell 1903b; Crawford 1914; Cresson 1864; Hobbs 1956; Ivanochko 1979; Michener 2007; Mitchell 1973; Montgomery 1957; Pengelly 1955; Provancher 1882; Sladen 1915; Thomson et al. 1982; Woodcock et al. 2014).

Coelioxys (Boreocoelioxys) sayi Robertson 1897 County records: Allegan, Barry, Benzie, Berrien, Branch, Calhoun, Cass, Gratiot, Hillsdale, Huron, Ingham, Ionia, Iosco, Isabella, Jackson, Kalamazoo, Lenawee, Midland, Missaukee, Monroe, Montcalm, Muskegon, Newaygo, Osceola, Ottawa, Shiawassee, St. Joseph, Van Buren, Washtenaw, Wayne. Notes. Recorded as a cleptoparasite of Megachile brevis (Baker 1975) and M. mendica (Medler 1965).

Subgenus Coelioxys Latreille Revision: Baker (1975).

Coelioxys (Coelioxys) sodalis Cresson 1878 County records: Alger, Cheboygan, Delta, Dickinson, Ingham, Kalamazoo, Keweenaw, Manistee, Marquette, Midland, Wayne. Notes. Hosts include Megachile melanophaea (Graenicher 1927a, 1935; Pengelly 1955), M. mucida (Gibbs 2017), M. frigida and M. texana (Pengelly 1955), and possibly M. rotundata (Hobbs 1968). Coelioxys sodalis is now considered the only Nearctic member of the subgenus Coelioxys (Rocha Filho & Packer 2016).

Subgenus Cyrtocoelioxys Mitchell Key: Baker (1975).


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Coelioxys (Cyrtocoelioxys) modestus Smith 1854 County records: Berrien, Ingham, Kalamazoo, Livingston, Menominee, Montmorency, St. Joseph, Washtenaw, Wayne. Notes. Hosts include resin collecting species in the subgenus Megachile (Chelostomoides), including Megachile campanulae (Krombein 1967; O’Neill & O’Neill 2016). Records from leafcutting species, such as M. centuncularis (Graenicher 1927b), and M. relativa (Fye 1965), may be due to confusion with C. moestus (Krombein 1967).

Subgenus Paracoelioxys Gribodo Revision: Baker (1975) as subgenus Schizocoelioxys Mitchell.

Coelioxys (Paracoelioxys) funerarius Smith 1854 County records: Charlevoix, Cheboygan, Dickinson, Iron, Keweenaw, Lake, Marquette, Sanilac. Notes. Hosts include Megachile frigida (Mitchell 1962), M. inermis Provancher (Medler 1958; Sheffield et al. 2008), M. latimanus (Graenicher 1927a), M. relativa (Fye 1965; Medler & Koerber 1958; Medler & Lussenhop 1968; Sheffield et al. 2008), and possibly M. rotundata (Hobbs 1968), although the known host M. relativa was also present in that study. In Michigan, both M. relativa and M. inermis are known hosts based on trap nest data (Packer et al. 1995; Strickler & Scriber 1994).

Subgenus Synocoelioxys Mitchell Revision: Baker (1975).

Coelioxys (Synocoelioxys) alternatus Say 1837 County records: Alpena, Cass, Gladwin, Ingham, Jackson, Kalamazoo, Kent, Lapeer, Livingston, Saginaw, Shiawassee, St. Joseph, Van Buren, Wayne. Notes. Recorded as a cleptoparasite of Megachile pugnata say (Medler 1964a), an association frequently seen in Michigan (JG, pers. obs).

Subgenus Xerocoelioxys Latreille s. s. Revision: Baker (1975); Rocha-Filho & Packer (2016).

Coelioxys (Xerocoelioxys) immaculatus Cockerell 1912 (New state record) County records: Livingston, Oceana. Notes. This cleptoparasite of Megachile, including M. addenda Cresson in New Jersey (Cane et al. 1996), has been collected once in Michigan and in adjacent areas of Indiana (Grundel et al. 2011; Jean 2010). Material examined. Oceana Co.: Hart, 19 Jun. 1939, C.W. Sabrosky (1 ♀ MSUC).



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Genus Megachile Latreille Taxonomy: Mitchell (1934, 1935a; b, 1936a; b, 1937a; b; c, 1962); Parker (1978); Ivanochko (1979); Sheffield et al. (2011b) Biology. Solitary leaf-cutting bees, including both ground and cavity-nesters. Summarized in Sheffield et al. (2011b), Ivanochko (1979) and Hobbs & Lilly (1954).

Subgenus Acentron Mitchell Taxonomy: Mitchell (1934).

Megachile (Acentron) albitarsis Cresson 1872 County records: Livingston.

Subgenus Callomegachile Michener Megachile (Callomegachile) sculpturalis Smith 1853 County records: Cheboygan, Ingham, Jackson, Kent, Monroe, Oakland, Osceola, Washtenaw, Wayne. Notes. Megachile scultupuralis, a distinctive exotic species of East Asian origin, has recently spread across the Northeast and Midwest regions of the country (Hinojosa-Díaz 2008; Magnum & Brooks 1997). It was first recorded from Michigan in 2008 from Wayne and Cheboygan Counties in the southeast and north of the LP, respectively (O’Brien & Craves 2008). The species, known to have been present in the state from at least 2006 (see below; the first Illinois specimen was also collected that year, see Tonietto & Ascher 2009), is active in midsummer. It is known to nest in active Xylocopa virginica burrows in dead wood (Laport & Minckley 2012; Roulston & Malfi 2012) and has been observed in Michigan removing nesting materials of other Megachile species from trap nests (JG pers. obs.). Material examined. Ingham Co.: East Lansing, Sep. 2006, P. Giordano (1 ♀ MSUC); East Lansing, MSU, N42.72 W84.473, 18 Jul. 2014 (2 ♂), 21 Aug. 2013 (1 ♀) (JBWM); East Lansing, MSU, Natural Science Bldg., 13 Jul. 2011, Stphynolobium japonicum, G. Parsons (3 ♀ 11 ♂ MSUC); East Lansing, MSU, Natural Science Bldg., 20 Aug. 2009, Sophora japonica, G. Parsons (16 ♀ MSUC); East Lansing, MSU, Radiology Bldg. N42717 W84.467, 20 Jul. 2014, J. Gibbs (1 ♀ JBWM); Lansing area, Nielsen residence, T4N R2W S7, 2 Jul.2006, M.C. Nielsen (1 ♀ MSUC); Lansing, Saginaw/MLK blvd., stem nest side of house, 18 Aug. 2008, R. Isaacs (2 ♀ MSUC); Kent Co.: (unknown locality) 3 Jul. 2012, M. Vanderwerp (1 ? MVWC); Oakland Co.: 26 Jul. 2014, M. Vanderwerp (MVWC); Osceola Co.: Leroy, 1.5 km SW, N44.0311 W85.4729, 26 Jun. 2015, J. Gibbs (1 ♀ JBWM); Shiawassee Co.: North of Perry (C. Sheffield and J. Gibbs pers. obs. 2006).

Subgenus Chelostomoides Robertson Revision: Mitchell (1937c).

Megachile (Chelostomoides) campanulae (Robertson 1903) (exilis group) County records: Allegan, Barry, Berrien, Cheboygan, Clinton, Ingham, Jackson, Kalamazoo, Livingston, Marquette, Midland, Missaukee, Montmorency, Oakland, Oscoda, Presque Isle, St. Joseph, Van Buren, Washtenaw, Wayne. Notes. A resin-collecting species that nests in stems. Nests described from Kansas, New York (Krombein 1967), and Wisconsin (Medler 1966; Medler & Lussenhop 1968).


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Megachile (Chelostomoides) rugifrons (Smith 1854) (rugifrons group) County records: Clinton, Livingston, Midland.

Subgenus Eutricharaea Thomson Taxonomy: Parker (1978); Mitchell (1980).

Megachile (Eutricharaea) apicalis Spinola 1808 (New state record) (Fig. 19A) County records: Ingham. Notes. A single female specimen was collected adjacent to a railway line in 2013. After repeated attempts to recollect the species at the same locality, an additional female and male were found in 2014. Megachile apicalis prefers Centaurea (Asteraceae) (Müller & Bansac 2004), which was abundant at the collection site. The female of the species is similar to the alfalfa leafcutter bee, M. rotundata, but can be distinguished by the apical projection of the clypeus and the darker scopal hairs on S4 in addition to S5 (Parker 1978; Sheffield et al. 2011b). Megachile rotundata is commonly observed in the same area. A continuing expansion of its range is likely, similar to that seen in other cavity nesting megachilid bees (Gibbs & Sheffield 2009; Hinojosa-Díaz 2008). In other US localities, this species can be very abundant and aggressively competes for nesting sites (Barthell & Thorp 1995; Stephen 2003; Thorp 1996). Further range extensions of this species might be facilitated by the spread of the invasive plant Centaurea stoebe L. (Spotted Knapweed) (Richardson et al. 2000). Material examined. Ingham Co.: Michigan State University, south campus, N42.721 W84.473, 9 Jun. 2013 (1 ♀ MSUC); Michigan State University, south campus, N42.721 W84.473, 23 Aug. 2014 (1 ♀ 1 ♂ JBWM).

FIGURE 19. Megachilid bees from Michigan (not to scale). A) Megachile (Eutricharaea) apicalis Spinola female, face. B) Ashmeadiella (Ashmeadiella) bucconis bucconis (Say) male, oblique habitus.

Megachile (Eutricharaea) pusilla Pérez 1884 (New state record) County records: Clinton, Ingham. Notes. Megachile pusilla is an exotic species that has evidently been present in central Michigan since at least



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1956, but it has been infrequently collected. Its close similarity to the abundant M. rotundata, differing from the latter by the narrower shape of the vertex and gena dorsally and fainter T2 fovea, may partially explain why it is not commonly recognized. The identity of North American populations were only recently clarified (Soltani et al. 2017). Previously, these bees were identified as M. concinna Smith, but this species sensu stricto is now unknown from the continental United States, although there are North American records from the Greater and Lesser Antilles. Cavity nesting was described from Arizona and Tennessee as M. concinna (Butler & Ritchie 1965; Butler & Wargo 1963; Krombein 1967). Material examined. Ingham Co.: Bath, 22 Jun. 1986, R.L. Fischer (1 ♀ MSUC); Lansing, 31 Aug. 1956, R.W. Hodges (1 ♀ MSUC); Lansing, 5 Sep. 1956, R.W. Hodges (1 ♀ MSUC).

Megachile (Eutricharaea) rotundata (Fabricius 1787) County records: Alger, Barry, Cass, Houghton, Ingham, Kalamazoo, Kalkaska, Kent, Lake, Leelanau, Mecosta, Oakland, Washtenaw, Wayne, Wexford. Notes. Exotic. The nesting biology has been well studied as M. rotundata is actively managed for alfalfa pollination (Hurd 1979; Pitts-Singer & Cane 2011). It was found in abundance in trap-nests placed on MSU campus, East Lansing, Michigan (JG, pers. obs.).

Subgenus Leptorachis Mitchell Taxonomy: Mitchell (1934).

Megachile (Leptorachis) petulans Cresson 1878 (New state record) County records: Kalamazoo, Newaygo, Van Buren. Notes. Megachile petulans is known from a few locations in the southern tier of counties. It has previously been recorded from the neighboring states of Indiana and Ohio (Mitchell 1962). Material examined. Kalamazoo Co.: Gull Lake Bio. Sta. (KBS), 18 Jul. 1956 (1 ♂, Melilotus alba; 1 ♂, Monarda fistulosa), 1 Aug. 1956 (2 ♂) 11 Aug. 1969 (1 ♂), 14 Aug. 1964 (1 ♂), R.L. Fischer (MSUC); Newaygo Co.: (no locality) 15 Jul. 1950, R.R. Dreisbach (1 ♀ MSUC); Van Buren Co.: (no locality) 25 Jul. 1942, R.R. Dreisbach (1 ♀ MSUC).

Subgenus Litomegachile Mitchell Taxonomy: Mitchell (1935a); Bzdyk (2012).

Megachile (Litomegachile) brevis Say 1837 County records: Allegan, Alpena, Barry, Branch, Calhoun, Cass, Cheboygan, Gratiot, Hillsdale, Ingham, Isabella, Kalamazoo, Leelanau, Livingston, Mason, Newaygo, Oakland, Oceana, Otsego, Ottawa, Saginaw, Shiawassee, St. Joseph, Van Buren. Notes. The nesting biology of M. brevis in Kansas has been studied in detail (Michener 1953). Additional studies of this cavity and stem-nester are available from Kansas (Baker et al. 1985), Missouri (Rau 1934), Wisconsin (Medler & Lussenhop 1968), and Oregon (Rockwood 1951). The latter may pertain to M. onobrychidis Cockerell, which was formerly considered a subspecies of M. brevis (see Sheffield et al. 2011; Bzdyk 2012).


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Megachile (Litomegachile) mendica Cresson 1878 County records: Alger, Allegan, Barry, Berrien, Cass, Charlevoix, Cheboygan, Clinton, Delta, Eaton, Emmet, Gladwin, Gratiot, HIllsdale, Huron, Ingham, Ionia, Iosco, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Leelanau, Lenawee, Livingston, Manistee, Mecosta, Midland, Missaukee, Monroe, Montcalm, Muskegon, Newaygo, Oakland, Oceana, Osceola, Ottawa, Roscommon, Saginaw, Shiawassee, St. Joseph, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Trap-nests described from Wisconsin (Medler 1965; Medler & Lussenhop 1968), Kansas and North Carolina (Baker et al. 1985), and New York, Maryland, North Carolina and Florida (Krombein 1967). Nesting in an old log in Missouri reported by Rau (1922). There is a published record of M. mendica nesting in the ground in Texas (Williams et al. 1986), but this cannot be confirmed. The presumed depository at Texas A&M does not have any material matching the paper (K. Wright, in litt.).

Megachile (Litomegachile) texana Cresson 1878 County records: Barry, Cass, Charlevoix, Cheboygan, Clinton, Dickinson, Gladwin, Grand Traverse, Ingham, Iosco, Iron, Isabella, Kalkaska, Kent, Leelanau, Livingston, Mackinac, Manistee, Marquette, Midland, Missaukee, Oakland, Oceana, Otsego, Ottawa, Shiawassee, Van Buren, Washtenaw, Wayne. Notes. Several biological notes on this ground-nesting species are available (Hurd 1979), including from Missouri (Rau 1922), Colorado (Hicks 1926), New York (Eickwort et al. 1981), North Carolina (Krombein 1953) and Florida (Krombein 1970). Shallow nests, a few inches long, were also observed in flat sandy soil in Michigan (Gibbs 2017). One was adjacent to a highbush blueberry field in Van Buren County and a second was in a clearing at Rose Lake, Shiawassee County.

Subgenus Megachile Latreille s. s. Revision: Mitchell (1935b as Delomegachile).

Megachile (Megachile) centuncularis (Linnaeus 1758) County records: Allegan, Barry, Berrien, Cass, Cheboygan, Genesee, Huron, Ingham, Kalamazoo, Kalkaska, Kent, Livingston, Mackinac, Marquette, Mason, Midland, Newaygo, Oakland, Ottawa, St. Joseph, Van Buren, Washtenaw, Wayne, Wexford. Notes. Possibly exotic. Holarctic distribution that apparently does not extend into northwestern Canada and Alaska. Cavity-nests in North America have been described repeatedly (Hurd 1979), from California (Michelbacher & Hurd 1954), New York (Krombein 1967), Massachusetts (Packard 1869), Pennsylvania (Gentry 1874), Colorado (Hicks 1926), and Wisconsin (Medler 1959; Medler & Lussenhop 1968).

Megachile (Megachile) inermis Provancher 1888 County records: Alger, Alpena, Antrim, Arenac, Baraga, Bay, Benzie, Cheboygan, Chippewa, Clare, Crawford, Delta, Dickinson, Emmet, Gladwin, Gogebic, Grand Traverse, Iosco, Iron, Jackson, Kalkaska, Keweenaw, Lake, Leelanau, Livingston, Luce, Mackinac, Manistee, Marquette, Menominee, Midland, Missaukee, Montmorency, Oakland, Oceana, Ontonagon, Oscoda, Otsego, Presque Isle, Saginaw, Schoolcraft, Shiawassee, Van Buren. Notes. Cavity-nests described from Manitoba and Sastkatchewan (Peck & Bolton 1946; Stephen 1955, 1956), Wisconsin (Medler 1958; Medler & Lussenhop 1968), and Ontario (Fye 1965).



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Megachile (Megachile) lapponica Thomson 1872 (New state record) County records: Keweenaw. Notes. This record comes from a specimen identified by S. Droege. Due to a paucity of records of this boreal species from the Eastern United States and its similarity to M. relativa, additional collections and molecular confirmation would be desirable. North American populations of this bee were called M. nivalis Friese until Sheffield et al. (2011) placed this in synonymy with M. lapponica, described from Sweden and now considered a naturally Holarctic species. Nests in logs in Saskatchewan (Peck & Bolton 1946). Specimen record: Keweenaw Co.: Isle Royale, Davidson Island, N48.1252, W88.5111, 4 Aug. 2013, S. Wilson (1 ♀ PWRC).

Megachile (Megachile) montivaga Cresson 1878 County records: Allegan, Baraga, Barry, Berrien, Cass, Dickinson, Jackson, Kalamazoo, Kent, Keweenaw, Livingston, Marquette, Mecosta, Menominee, Ontonagon, Schoolcraft, Van Buren. Notes. Megachile montivaga nests in plant stems, including Cirsium (Orr et al. 2015), Helianthus (Hicks 1926), Rhus (Rau 1934), and Verbascum (Hicks 1926), and uses flower petals for nest construction, a trait correlated with reduced cutting edges of its mandibles. Trap-nesting described from Kansas (Baker et al. 1985) and Missouri (1934).

Megachile (Megachile) relativa Cresson 1878 County records: Alcona, Alger, Alpena, Antrim, Baraga, Barry, Benzie, Cheboygan, Crawford, Delta, Dickinson, Emmet, Gogebic, Ingham, Iosco, Iron, Jackson, Kalamazoo, Kent, Keweenaw, Leelanau, Luce, Mackinac, Manistee, Marquette, Mecosta, Menominee, Midland, Montmorency, Oakland, Ogemaw, Ontonagon, Osceola, Otsego, Ottawa, Presque Isle, Roscommon, Saginaw, Schoolcraft, Wexford. Notes. Stem-nesting described from Ontario (Fye 1965) and Wisconsin (Medler & Koerber 1958; Medler & Lussenhop 1968).

Subgenus Megachiloides Mitchell Revision: Mitchell (1936b).

Megachile (Megachiloides) dakotensis Mitchell 1926 (New state record) County records: Kalkaska, Wexford. Notes. This relatively large prairie-associated species is primarily known in Michigan from a relatively long series from a single location. Given these locations are in the north-central part of the Lower Peninsula, it seems likely that this species is far more widespread in the state. It has been recorded without specific locality data from both Illinois and Wisconsin (Mitchell 1962; Wolf & Ascher 2009). Material examined. Kalkaska Co.: (no locality), ex. window pane trap, 10 Jul. 1966 (5 ♀), 16 Jul. 1966 (2 ♀), 21 Jul. 1966 (16 ♀, 1 ♂), L.F. Wilson (MSUC); Wexford Co.: T24N R9W Sec. 22 (approximately 5 km N of Manton), 7–14 Jul. 1965, ex. Pit trap in Scotch pine plantation, J.H. Shaddy (1 ♂ MSUC).

Subgenus Sayapis Titus Revision: Mitchell (1937b).


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Megachile (Sayapis) frugalis frugalis Cresson 1872 County records: Allegan, Berrien, Ionia, Kalamazoo, Kent, Livingston, Ottawa, Washtenaw.

Megachile (Sayapis) inimica Cresson 1872 sayi Cresson 1878 County records: Kalamazoo. Notes. Trap-nests described from Arizona (Krombein 1967) and Wisconsin (Medler & Lussenhop 1968).

Megachile (Sayapis) pugnata pugnata Say 1837 County records: Allegan, Barry, Berrien, Cass, Cheboygan, Clinton, Dickinson, Eaton, Emmet, Gladwin, Gogebic, Gratiot, Ingham, Iron, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lake, Lapeer, Livingston, Macomb, Mecosta, Midland, Monroe, Montmorency, Oakland, Otsego, Ottawa, Roscommon, Saginaw, St. Joseph, Van Buren, Washtenaw, Wayne. Notes. Specialist on Asteraceae. This cavity-nester has been studied in Wisconsin (Medler 1964a; Medler & Lussenhop 1968). In Utah, it has been explored for its potential as a managed pollinator of commercial sunflower (Tepedino & Frohlich 1982). Megachile pugnata is commonly found in trap-nests placed in both MSU campus, East Lansing, Michigan and prairie restorations in the LP (JG, pers. obs.).

Subgenus Xanthosarus Robertson Revision: Mitchell (1936a).

Megachile (Xanthosarus) addenda Cresson 1878 County records: Allegan, Barry, Benzie, Berrien, Eaton, Grand Traverse, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Livingston, Mason, Newaygo, Ottawa, St. Joseph, Van Buren, Wexford. Notes. Nesting biology and potential as a wild pollinator of cranberry in New Jersey discussed by Cane et al. (1996).

Megachile (Xanthosarus) frigida frigida Smith 1853 County records: Alger, Baraga, Charlevoix, Cheboygan, Delta, Dickinson, Emmet, Ingham, Iosco, Iron, Keweenaw, Lake, Mackinac, Manistee, Marquette, Mason, Midland, Washtenaw. Notes. A nest in a poplar log was described from Manitoba (Stephen 1956). Graenicher (1905) records this species nesting in logs as M. addenda (Medler & Lussenhop 1968).

Megachile (Xanthosarus) gemula gemula Cresson 1878 County records: Alcona, Alger, Allegan, Alpena, Baraga, Barry, Berrien, Cheboygan, Clare, Clinton, Delta, Dickinson, Emmet, Gladwin, Gogebic, Grand Traverse, Iron, Kalamazoo, Kalkaska, Kent, Keweenaw, Leelanau, Mackinac, Manistee, Marquette, Midland, Missaukee, Montmorency, Ontonagon, Osceola, Oscoda, Otsego, Ottawa, Presque Isle, Roscommon, Schoolcraft, Shiawassee, St. Joseph, Van Buren.



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Notes. Trap nesting in bored stems described from Ontario (Fye 1965) and Wisconsin (Medler & Lussenhop 1968).

Megachile (Xanthosarus) latimanus Say 1823 County records: Alger, Allegan, Alpena, Arenac, Baraga, Barry, Bay, Berrien, Branch, Cheboygan, Clinton, Crawford, Delta, Dickinson, Eaton, Emmet, Genesee, Gladwin, Gratiot, Hillsdale, Houghton, Huron, Ingham, Iosco, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Lapeer, Leelanau, Livingston, Mackinac, Macomb, Manistee, Marquette, Mason, Mecosta, Menominee, Midland, Missaukee, Montcalm, Montmorency, Newaygo, Oakland, Oceana, Ogemaw, Osceola, Oscoda, Otsego, Ottawa, Presque Isle, Roscommon, Saginaw, Shiawassee, St. Clair, St. Joseph, Van Buren, Washtenaw, Wayne, Wexford. Notes. Ground-nests described from Wisconsin (Graenicher 1905).

Megachile (Xanthosarus) melanophaea melanophaea Smith 1853 County records: Alger, Baraga, Cheboygan, Clinton, Delta, Dickinson, Emmet, Gogebic, Huron, Iron, Kalkaska, Kent, Keweenaw, Luce, Mackinac, Mason, Midland, Montmorency, Oceana, Ontonagon, Osceola, Oscoda, Otsego, Ottawa, Roscommon, Washtenaw, Wexford. Notes. Ground-nester (Graenicher 1905).

Megachile (Xanthosarus) mucida Cresson 1878 County records: Berrien, Ingham, Ionia, Muskegon, Van Buren. Notes. Megachile mucida is poorly represented in historical collections, especially from the northern portion of its range, but it is now quite common in late spring in the Lansing area of Michigan. Its adult emergence was among the earliest of Megachile species in the area and its coloration is distinctive, except with respect to separation from its close relatives M. gemula and M. melanophaea. From those, M. mucida can be distinguished by characters of the female mandible and of the male tarsi (Mitchell 1962) and subtle differences in the distribution of pale hairs. Gibbs (2017) provided details on its distribution in Michigan and described its nesting habits.

Megachile (Xanthosarus) perihirta Cockerell 1898 County records: Mackinac, Schoolcraft. Notes. A widespread western species but poorly documented in the Eastern United States. The occurrence in Michigan was confirmed from a male specimen deposited at the UMMZ. The biology of this ground-nesting species has been studied in Alberta (Hobbs 1956; Hobbs & Lilly 1954; Sladen 1918).

Osmiini Genus Ashmeadiella Cockerell Revision: Michener (1939).

Subgenus Ashmeadiella Cockerell s. s. Ashmeadiella (Ashmeadiella) bucconis bucconis (Say 1837) (New state record) (Fig. 19B)


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County records: Otsego. Notes. Most species of Ashmeadiella are western, but bucconis was described from “Indiana” and it has also been recorded from Illinois and Wisconsin (Jean 2010; Michener 1939; Mitchell 1962). Nests of the subspecies A. b. denticulata (Cresson) were described from Arizona by Krombein (1967). Leaf pulp is used in cell partitions. Material examined. Otsego Co.: (no locality) 7 Jul. 1959 R. & K. Dreisbach (1 ♂ MSUC).

Genus Chelostoma Latreille Subgenus Prochelostoma Robertson Taxonomy: Eickwort (1980); Buck et al. (2005).

Chelostoma (Prochelostoma) philadelphi (Robertson 1891) County records: Allegan, Livingston, St. Joseph, Washtenaw, Wayne. Notes. A native species oligolectic on Philadelphus (Saxifragaceae). Two non-native Chelostoma are likely to occur in the state (see Appendix 1). Nests in Virginia described by Krombein (Krombein 1959, 1967). Mud is used in cell partitions.

Genus Heriades Spinola Subgenus Neotrypetes Robertson Taxonomy: Michener (1938); Mitchell (1962). Biology: Solitary stem-nesters that use resin in nest construction (Krombein 1967; Matthews 1965).

Heriades (Neotrypetes) carinata Cresson 1864 County records: Alcona, Allegan, Antrim, Arenac, Barry, Bay, Berrien, Calhoun, Cheboygan, Clare, Clinton, Delta, Dickinson, Eaton, Emmet, Gladwin, Gogebic, Grand Traverse, Gratiot, Huron, Ingham, Iosco, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Lake, Lapeer, Leelanau, Lenawee, Livingston, Mackinac, Mecosta, Menominee, Midland, Missaukee, Montcalm, Montmorency, Muskegon, Newaygo, Oakland, Oceana, Osceola, Oscoda, Otsego, Presque Isle, Roscommon, Shiawassee, St. Clair, Tuscola, Van Buren, Washtenaw, Wayne, Wexford. Notes. Nesting biology described from Missouri (Rau 1922), New York and North Carolina (Krombein 1967), Wisconsin (Medler & Lussenhop 1968), and Oregon and Michigan (Matthews 1965).

Heriades (Neotrypetes) leavitti Crawford 1913 County records: Cheboygan, Huron, Ingham, Kalamazoo, Lake, Livingston, Missaukee, Saginaw, Shiawassee, Van Buren. Notes. Nest from Florida described by Krombein (1967).

Heriades (Neotrypetes) variolosa variolosa (Cresson 1872) County records: Cheboygan, Montmorency, Oscoda, Otsego, Ottawa, Saginaw. Notes. Nest from Minnesota described by Fischer (1955).



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Genus Hoplitis Klug Taxonomy: Michener (1947a); Mitchell (1962); Sedivy et al. (2013). Biology. Solitary stem-nesters. Cell partitions are made of leaf pulp and sometimes pebbles are incorporated.

Subgenus Alcidamea Cresspm Hoplitis (Alcidamea) albifrons albifrons (Kirby 1837) (tuberculata group) County records: Alger, Baraga, Barry, Cheboygan, Dickinson, Iron, Keweenaw, Livingston, Marquette, Schoolcraft. Notes. Nest in Ontario described by Fye (1965).

Hoplitis (Alcidamea) pilosifrons (Cresson 1864) (producta group) County records: Allegan, Arenac, Barry, Benzie, Berrien, Calhoun, Cass, Cheboygan, Clinton, Dickinson, Gratiot, Huron, Ingham, Ionia, Isabella, Jackson, Kalamazoo, Lake, Leelanau, Livingston, Manistee, Mason, Midland, Montcalm, Montmorency, Oakland, Oceana, Osceola, Ottawa, Saginaw, Sanilac, Shiawassee, St. Clair, St. Joseph, Tuscola, Van Buren, Washtenaw, Wexford. Notes. A stem-nesting leaf-cutter bee, based on nests from Kansas (Michener 1955)

Hoplitis (Alcidamea) producta producta (Cresson 1864) (producta group) County records: Allegan, Antrim, Barry, Benzie, Berrien, Branch, Cass, Charlevoix, Cheboygan, Chippewa, Clare, Clinton, Crawford, Delta, Dickinson, Eaton, Genesee, Gladwin, Gratiot, Hillsdale, Houghton, Ingham, Iron, Isabella, Jackson, Kalamazoo, Kalkaska, Kent, Keweenaw, Lapeer, Leelanau, Lenawee, Livingston, Luce, Macomb, Manistee, Marquette, Mason, Mecosta, Menominee, Midland, Missaukee, Monroe, Montmorency, Newaygo, Oakland, Oceana, Osceola, Oscoda, Ottawa, Roscommon, Saginaw, Schoolcraft, Shiawassee, St. Clair, St. Joseph, Van Buren, Washtenaw, Wayne, Wexford. Notes. Nests decribed from Colorado (Hicks 1926), Missouri (Rau 1928), and Wisconsin (Graenicher 1905; Medler 1961; Medler & Lussenhop 1968).

Hoplitis (Alcidamea) spoliata (Provancher 1888) (tuberculata group) County records: Alcona, Allegan, Alpena, Antrim, Barry, Bay, Berrien, Cass, Cheboygan, Dickinson, Eaton, Grand Traverse, Gratiot, Houghton, Huron, Iosco, Jackson, Kalamazoo, Kent, Keweenaw, Lake, Lapeer, Livingston, Marquette, Mecosta, Menominee, Osceola, Otsego, Roscommon, Saginaw, Van Buren, Washtenaw, Wexford. Notes. Nests described from Colorado (Hicks 1926), Ontario (Fye 1965), Wisconsin (Medler 1967b; Medler & Lussenhop 1968), and Kansas (Baker et al. 1985).

Hoplitis (Alcidamea) truncata truncata (Cresson 1878) (truncata group) County records: Calhoun, Cass, Barry, Berrien, Cheboygan, Dickinson, Huron, Ingham, Jackson, Kalamazoo, Montmorency, Oscoda, Schoolcraft, Shiawassee, Washtenaw.


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Genus Osmia Panzer Taxonomy: Sandhouse (1939); Mitchell (1962); Rust (1974). Biology. Typically solitary cavity-nesters or ground-nesting. Reviewed by Cane et al. (2007) and Rust (1974). Cell partitions are typically made of leaf pulp, but mud may also be used.

Subgenus Cephalosmia Sladen Revision: Rust (1974).

Osmia (Cephalosmia) subaustralis Cockerell 1900 County records: Alger, Alpena, Cheboygan, Kalkaska, Montmorency, Newaygo, Oscoda, Otsego, Wexford. Notes. A nest in a beetle burrow in an old log from Utah is described by Rust (1974).

Subgenus Diceratosmia Robertson Revision: Michener (1949).

Osmia (Diceratosmia) conjuncta Cresson 1864 County records: Allegan, Antrim, Barry, Berrien, Cheboygan, Clare, Clinton, Hillsdale, Huron, Ingham, Kalamazoo, Leelanau, Mackinac, Macomb, Mason, Midland, Missaukee, Oakland, Otsego, Presque Isle, Shiawassee, Van Buren. Notes. Nests from Missouri were found in abandoned snail shells (Rau 1937).

Subgenus Helicosmia Thomson Revision: Rust (1974 as Chalcosmia).

Osmia (Helicosmia) caerulescens (Linnaeus 1758) County records: Calhoun, Cheboygan, Clinton, Ingham, Iosco, Jackson, Kalamazoo, Kent, Leelanau, Livingston, Manistee, Midland, Monroe, Montcalm, Oceana, Presque Isle, Sanilac, Schoolcraft, Shiawassee, Van Buren, Washtenaw, Wayne. Notes. Probably exotic, based on lack of records from Alaska and, until recently, from western Canada and the Pacific Northwest (now well established in, e.g., the Seattle vicinity, as shown by records on http://bugguide.net) but historically was restricted to Eastern North America. A relatively early Idaho record was considered “probably adventive” by Hurd (1979). Cavity-nesting described from New York (Krombein 1967) and Wisconsin (Medler 1967a). Its potential as an alfalfa pollinator has been studied in France (Tasei & Picart 1972).

Osmia (Helicosmia) georgica Cresson 1878 County records: Allegan, Jackson, Kent, Livingston, Midland, Ottawa, Sanilac, St. Joseph, Washtenaw. Notes. Specialist on Asteraceae. Cavity-nesting described from Connecticut (Hartman et al. 1944) and Tennessee (Hawkins 1975; Krombein 1967). THE BEES OF MICHIGAN


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Osmia (Helicosmia) texana Cresson 1872 County records: Gratiot, Ingham, Kalamazoo, Livingston, Macomb, Shiawassee. Notes. Nests in cavities including multiple reports of this species in old Anthophora nests from Colorado, Alberta, and Utah (Hicks 1926; Hobbs et al. 1961; Rust 1974).

Subgenus Melanosmia Schmiedeknecht Taxonomy: Rightmyer et al. (2010).

Osmia (Melanosmia) albiventris Cresson 1864 County records: Cheboygan, Kalamazoo, Keweenaw, Livingston, Marquette, Montmorency, Shiawassee, St. Clair, St. Joseph. Notes. Trap-nests described from Wisconsin (Medler 1967a; Medler & Lussenhop 1968), Georgia and South Carolina (Jenkins & Matthews 2004).

Osmia (Melanosmia) atriventris Cresson 1864 County records: Alger, Alpena, Allegan, Antrim, Barry, Berrien, Charlevoix, Cheboygan, Emmet, Grand Traverse, Ingham, Jackson, Keweenaw, Lake, Lapeer, Leelanau, Livingston, Marquette, Midland, Muskegon, Oceana, Ottawa, Saginaw, Shiawassee, Van Buren, Washtenaw. Notes. Cavity-nesting described in Ontario (Fye 1965) and South Carolina (Horn & Hanula 2004).

Osmia (Melanosmia) bucephala Cresson 1864 County records: Alger, Allegan, Barry, Berrien, Cheboygan, Dickinson, Ingham, Kalamazoo, Kent, Keweenaw, Luce, Marquette, Midland, Montmorency, Oscoda, Schoolcraft, Van Buren. Notes. Nests from Maryland and Washington, D.C. described by Krombein (1967) and Packard (1867), respectively. Unique among Osmia for its use of wood fibers in cell construction; partitions are made of leaf pulp.

Osmia (Melanosmia) collinsiae Robertson 1905 County records: Alpena, Lapeer, Montmorency, Oscoda, Wexford.

Osmia (Melanosmia) distincta Cresson 1864 County records: Allegan, Barry, Berrien, Clinton, Eaton, Hillsdale, Ingham, Isabella, Kent, Livingston, Muskegon, Oakland, Ottawa, Shiawassee, Van Buren. Notes. Specialist on Penstemon (Scrophulariaceae).

Osmia (Melanosmia) felti Cockerell 1911 (Newly confirmed state record) County records: Keweenaw, Marquette.


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Notes. Osmia felti was recorded from southwest Michigan based on a single female identified by JSA (Tuell et al. 2009). The specimen, deposited at MSUC: RI, was reexamined by JG and determined to be O. virga Sandhouse. However, O. felti is confirmed to occur in the UP based on specimens below. Material examined. Keweenaw Co.: Isle Royale, N46.6616 W86.0252, 17 May 2011 (1 ♂ PWRC), 19 Aug. 2010 (1 ♀ PWRC); Isle Royale, N47.8763 W89.2326, 19 Aug. 2013, S. Wilson (1 ♀ MSUC: RI); Marquette Co.: Salmon Trout River Br. Huron Mt. Club, 8 Aug. 1981, D. Gosling (1 ♀ MSUC).

Osmia (Melanosmia) inermis (Zetterstedt 1838) (inermis group) County records: Alger, Marquette, Montmorency, Oscoda. Notes. Holarctic (Müller 2016; Rightmyer 2010). Nests under stones (Cane et al. 2007). Sheffield et al. (2015) demonstrated the use of artificial terracotta nests to encourage this species for pollination of lowbush blueberry.

Osmia (Melanosmia) inspergens Lovell & Cockerell 1907 County records: Kalkaska, Keweenaw, Montmorency, Oscoda, Saginaw, Schoolcraft. Notes. Nests under stones (Krombein 1967).

Osmia (Melanosmia) laticeps Thomson 1872 (inermis group) County records: Alger, Marquette. Notes. A specialist on Vaccinium (Rightmyer et al. 2010).

Osmia (Melanosmia) nigriventris (Zetterstedt 1838) (nigriventris group) County records: Marquette. Notes. Holarctic (Müller 2016; Rightmyer et al. 2010). Nests in wood (Cane et al. 2007).

Osmia (Melanosmia) proxima Cresson 1864 County records: Alcona, Alger, Allegan, Bay, Charlevoix, Cheboygan, Dickinson, Emmet, Iron, Keweenaw, Lapeer, Mackinac, Marquette, Ontonagon, Oscoda, Presque Isle, Roscommon, Wexford. Notes. Nests described by Fye (1965) and Medler (1967a) from Ontario and Wisconsin, respectively.

Osmia (Melanosmia) pumila Cresson 1864 County records: Allegan, Barry, Berrien, Cass, Ingham, Ionia, Jackson, Kent, Lapeer, Livingston, Ottawa, Presque Isle, Saginaw, St. Clair, Tuscola, Van Buren, Washtenaw. Notes. Nest described Maryland (Krombein 1967), New York (Goodell 2003), and Wisconsin (Medler 1967a; Medler & Lussenhop 1968).

Osmia (Melanosmia) simillima Smith 1853 County records: Alger, Allegan, Barry, Bay, Berrien, Branch, Calhoun, Cheboygan, Clinton, Dickinson, Emmet, Gladwin, Huron, Ingham, Ionia, Kalamazoo, Keweenaw, Lake, Lapeer, Leelanau, Livingston, Manistee, Mecosta, Midland, Missaukee, Montcalm, Ottawa, Roscommon, Saginaw, Shiawassee, St. Clair, Van Buren. Notes. Recorded nesting in oak galls, possibly from Massachusetts (Packard 1867). Scott (1993) describes this



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species nesting communally in buried wood at a sandy site at Pictured Lakes National Lakeshore, Michigan. A nest in driftwood on the west shore of Lake Michigan was reported by Graenicher (1905).

Osmia (Melanosmia) subarctica Cockerell 1912 (New status) Osmia (Nothosomia) michiganensis Mitchell, 1962: 92 (New synonymy; Michigan holotype; Fig. 20) Holotype. ♂ USA: Michigan: Grand Traverse Co.: 27 May 1950, R.R. Dreisbach (USNM ENT 00536966) (on indefinite loan from NCSU).

County records: Dickinson, Grand Traverse. Notes. Osmia subarctica is poorly known due to its northern range and past taxonomic confusion. Osmia subarctica was treated as a synonym of O. tersula Cockerell by Sandhouse (Sandhouse 1939) based on the identical locality information (“Hudson’s Bay, 44-17”, types deposited at the Natural History Museum, London) and apparent large impunctate margins of the metasomal terga shared by both species and this synonymy was followed by Hurd (1979). However, upon closer inspection of the holotype of O. subarctica by MGR, there are actually fine punctures that narrow the true impunctate margin of the terga. More importantly, the punctures of the clypeus and the curvature of the hind tibial spur are both different from what is found in females of O. tersula. In O. subarctica, the interspaces between the punctures of the clypeus are raised and the punctures themselves are elongated, giving the entire clypeus a wrinkled look, while in O. tersula the interspaces are flat and the punctures are round. In O. subarctica, the hind tibial spur is moderately thick, but strongly curved on its apical fifth, while in O. tersula the hind tibial spur is slightly thinner and the much less strongly curved apically. Mitchell (1962) described O. michiganensis based on a single male specimen and there are few subsequent records. Earlier specimens recognized by Mike Arduser as O. michiganensis are not O. subarctica (M. Arduser, in litt.), but rather belong to O. aff. trevoris, which follows below. A published record of O. michiganensis from Jackson Co., Michigan (Fiedler et al. 2012) was a misidentified O. atriventris. The first report of this species outside of Michigan is from the Apostle Islands in far northern Wisconsin, by S. Droege (in litt. 2010).

Osmia (Melanosmia) tarsata Provancher 1888 Osmia tarsata Provancher 1888: 328. Osmia kenoyeri Cockerell 1915: 483. Synonymy by Sheffield & Perron (2014) Osmia hendersoni Cockerell 1907: 365. New synonymy. Holotype. ♀ USA: Colorado: Arapahoe Peak, 1 Sep. 1906, S. A. Rohwer (NMNH 27891).

County records: Alger, Chippewa. Notes. This distinctive species occurs in the UP. Its taxonomy has been confused in the past, having been treated as a synonym of O. bucephala by Sandhouse (1939), but resurrected by Mitchell (1962) based on examination of the type material. In the same work, Mitchell recorded the female from Michigan as Osmia integra Cresson, a misidentification, because integra as now understood has a more western distribution. Sheffield & Perron (2014) recently synonymized O. kenoyeri with O. tarsata, which expands its known range of O. tarsata considerably. A series of males and females at PWRC was identified by S. Droege (as O. kenoyeri). Mike Arduser (in litt.) reports this species nesting in stable sand dunes and outwash plains at Grand Sable Dunes, Pictured Rocks National Lakeshore. Nests excavated in 1987 had seven to nine globular cells composed of masticated Fragaria leaves attached to roots at the base of little bluestem grasses. Material examined. Alger Co.: Pictured Rocks, N46.6635 W86.0111, 9 Jun. 2011, (2 ♀ PWRC); Pictured Rocks, N46.6616 W86.0252, 20 May 2011, (4 ♀ 13 ♂ PWRC), 14 Jul. 2011, (1 ♀ PWRC); Chippewa Co.: (no locality) 7 Jun. 1959, (1 ♀ MSUC); Paradise Bog T49N R7W, Sec. 15, on bearberry, 22 May 1993, (1 ♀ MSUC).


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FIGURE 20. Osmia michiganensis Mitchell holotype male (= O. subarctica Cockerell). A) Lateral habitus. B) Face. Photographs provided with the permission of the National Museum of Natural History, Smithsonian Institution, 10th and Constitution Ave. N.W., Washington, DC 20560-0193. (http://www.nmnh.si.edu/).

Osmia (Melanosmia) tersula Cockerell 1912 County records: Chippewa, Delta, Dickinson, Keweenaw, Marquette, Montmorency, Ontonagon, Oscoda, Otsego, Roscommon, Wexford. Notes. Trap-nests from Wisconsin described by Medler (1967a).

Osmia (Melanosmia) aff. trevoris Cockerell 1897 County records: Alger, Allegan, Chippewa, Kalkaska, Keweenaw, Marquette, Missaukee, Montcalm, Montmorency, Oscoda, Schoolcraft. Notes. Females of O. aff. trevoris can be recognized by the following combination of characters: clypeal margin with small triangular notch medially, clypeal setae brown, mandibular carinae parallel and scopa dark brown (M. Arduser, unpublished data). Males would run to couplet 27 in Mitchell (1962), being similar to both O. collinsiae Robertson and especially O. subarctica (see above). The species was compared to O. inurbana Cresson and O. trevoris by MGR, likely synonymous names based on male and female types, and widely collected in western North America. Material from Michigan, Indiana, and Ohio need further study to resolve the taxonomic status of these specimens. Material examined. Kalkaska Co.: (no locality), 1 Jun. 1966, ex. window pane trap, L.F. Wilson (2 ♀ MSUC); Montcalm Co.: Flat River Game Area, 14 May 1955, R.L. Fischer (1 ♀ MSUC); Montmorency Co.: (no locality), 5 Jul. 1966 ex. window pane trap, P.C. Kennedy (2 ♀ MSUC); (no locality), 18 Jul. 1966 ex. window pane trap (1 ♀ MSUC); Oscoda Co.: (no locality), 14 Jun. 1966 ex. window pane trap, L.F. Wilson (3 ♀ MSUC).

Osmia (Melanosmia) virga Sandhouse 1939 County records: Allegan, Berrien, Clinton, Crawford, Muskegon, Osceola, Ottawa, Van Buren. Notes. A potentially valuable native pollinator of blueberry due to its preference for Vaccinium. Mitchell (1962) recognized only the male, contributing to this species being overlooked or confused with O. felti (see Giles & Ascher 2006; Goldstein & Ascher 2016).



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Subgenus Osmia Panzer s. s. Revision: Rust (1974).

Osmia (Osmia) cornifrons (Radoszkowski 1887) County records: Allegan, Berrien, Grand Traverse, Ingham, Ionia, Kalamazoo, Kent, Leelanau, Oakland, Oceana, Ottawa, Van Buren, Washtenaw. Notes. This non-native species was intentionally introduced to North America from Japan for pollination studies in both Utah and Maryland, in 1965 and 1977, respectively (Batra 1979; Rust 1974), and became established in the Mid-Atlantic states following the latter. It was shipped to Leelanau County, Michigan from Indiana in 2005 for pollination trials on cherry with subsequent imports from Pennsylvania (Rothwell 2006; JG, pers. obs.). It is a potentially valuable economic pollinator of cherry and apple (Batra 1979, 2007). The species is now well established in Michigan in both urban settings and semi-natural areas near urban centers. It can be collected in large numbers in the spring.

Osmia (Osmia) lignaria lignaria Say 1837 County records: Alger, Allegan, Berrien, Clinton, Delta, Dickinson, Genesee, Gogebic, Grand Traverse, Ingham, Ionia, Iron, Kalamazoo, Kent, Keweenaw, Livingston, Marquette, Montcalm, Oakland, Ontonagon, Ottawa, Roscommon, Shiawassee, Van Buren, Washtenaw, Wayne. Note. Nests have been described by Rau (1937), Krombein (1967), Medler (1967a) and others. The western subspecies O. l. propinqua Cresson in particular, has been used as a managed crop pollinator (Bosch et al. 2000; Bosch & Kemp 2001, 2002).

Osmia (Osmia) taurus Smith 1873 (New state record) (Fig. 21) County records: Kalamazoo. Notes. This exotic species, native to East Asia and widespread in Japan, is closely related to O. cornifrons (above). Although it was first detected many years after the introduction of O. cornifrons (Buchmann & Ascher 2005; Ebmer 2011), it seems plausible that it was introduced accidentally at the same time. The species has spread along the eastern states from New York to Georgia largely along the Appalachian Mountains. A single male was collected from a semi-natural area near a low-density population center and agricultural lands. A series of O. cornifrons were collected at the same site. The species were distinguished using a published key (Yasumatsu & Hirashima 1950). The spread of O. taurus into the Midwest has not been well-documented, except from photographs identified online from Indiana (http://bugguide.net/node/view/913803l) and Ohio (http:// bugguide.net/node/view/1220268), and a recent dissertation on bees from Cleveland (Prajzner 2016). The first Michigan record of O. taurus could easily have been overlooked due to the abundance of O. cornifrons at this site, which look very similar. Osmia cornifrons, being readily distinguishable with the naked eye from species other than O. taurus, were of little interest to the collector at the time. This example illustrates one benefit of collecting insects in series even when the identity is presumed to be known. Material examined. Kalamazoo Co.: Kellogg Bird Sanctuary, N42.394 W85.385, 16 Apr. 2016, J. Gibbs & Y. Nozoe (1 ♂ MSUC).


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FIGURE 21. Osmia (Osmia) taurus Smith male from Michigan. A) Lateral habitus. B) Genital capsule.

MELITTIDAE Mellitinae Genus Macropis Panzer Taxonomy: Michez & Patiny (2005); Mitchell (1960) Biology. Solitary oil-collecting specialists on Lysimachia (see Cane et al. 1983).

Subgenus Macropis Panzer s. s. Macropis (Macropis) nuda (Provancher 1882) County records: Allegan, Alpena, Arenac, Berrien, Cheboygan, Dickinson, Gladwin, Hillsdale, Huron, Ingham, Iosco, Isabella, Lapeer, Mackinac, Midland, Newaygo, Osceola, Otsego, Saginaw, Van Buren. Notes. To our knowledge, only a single male of this oil-collecting bee has been collected in Michigan since 1959, although recent collections have been made locally in other states (Wagner & Ascher 2008) and in eastern Canadian provinces (Sheffield et al. 2004; Wagner & Ascher 2008). The bee has been collected historically across the entire LP, and its host plant is present widely in the state (Voss & Reznicek 2012). Macropis nuda was among the species in the northeast modeled to be in decline based on historical records (Bartomeus et al. 2013a). The possibility of a decline is supported by lack or extreme scarcity of this and other Macropis species in recent bee surveys made using bowl traps in portions of the mid-Atlantic region where it was once common such as the vicitiny of Washington, DC (S. Droege, pers. comm.). The lack of recent Michigan records of this species and of an associated cleptoparasite (Epeoloides pilosulus, last collected in 1944; see entry above) may be due in part to insufficient collection effort focused on this species. Macropis specializes on Lysimachia (Cane et al. 1983; Michez & Patiny 2005), a plant that grows in moist, often shaded ground (Voss & Reznicek 2012). Localities with L. ciliata L. present, sometimes in abundance near extensive riparian zones, were visited by JG in Ingham, Clinton, and Benzie Counties, in some cases on numerous occasions, specifically to collect Macropis, but without success. Material examined. Allegan Co.: Allegan, 2 Jul. 1936 (1 ♀ MSUC); Hillsdale Co.: Pittsford SGA, 41.866, 84.522, 8 Jul. 2017 (1 ♂ TJWC); Ingham Co.: East Lansing, 20 Jul. 1937 (1 ♀ MSUC); Mackinac Co.: (no locality) 11 Aug. 1959 (1 ♂ MSUC); Midland Co.: (no locality) 5 Jul. 1938 (1 ♀ 1 ♂ MSUC); Osceola Co.: 3 Aug. 1940 (1 ♂ MSUC); Otsego Co.: 5 Jul. 1940 (1 ♂ MSUC); Saginaw Co.: (no locality) 14 Jul. 1940 (1 ♀ MSUC).



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Acknowledgements We thank Julianna Wilson for providing information on collection localities, Sam Droege for providing access to Michigan specimen information in his database and sharing opinions on Nomada (any mistakes in our treatment of this genus are our own), Mike Arduser (Missouri Department of Conservation) and Thomas Wood (MSU) for valuable comments on the manuscript, and Barry O’Connor and Mark O’Brien for allowing visits to UMMZ. Thomas Wood provided a number of valuable records collected in 2017. Thanks also to Ann Fraser who allowed examination of KCIC material, and Anthony Cognato and Gary Parsons for providing uninhibited access to MSUC. Hadel Go (AMNH) provided images of Protandrena cockerelli from other localities. Loans of material were generously made by V. Scott (UCMC), J. Leather (OSAC), T.L. Griswold, H. Ikerd (BBSL), V. Lee, N. Penny, W. Pulawski, R. Zuparko (CAS), Rob Jean (Environmental Solutions & Innovations, Inc.), D. Dmitriev, C. Favret, J. Zahniser (INHS) and S. Brady, D. Furth and B. Harris (NMNH). Post-doctoral support for JG was made available by funding from the United States Department of Agriculture-National Institute for Food and Agriculture, Specialty Crop Research Initiative: Developing Sustainable Pollination Strategies for U.S. Specialty Crops (201251181-20105) and by Project GREEEN.

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(Hymenoptera: Megachilidae) in Canada and Alaska. Canadian Journal of Arthropod Identification, 18, 1–107. https://doi.org/10.3752/cjai.2011.18 Sheffield, C.S., Rigby, S.M., Smith, R.F. & Kevan, P.G. (2004) The rare cleptoparasitic bee Epeoloides pilosula (Hymenoptera: Apoidea: Apidae) discovered in Nova Scotia, Canada, with distributional notes. Journal of the Kansas Entomological Society, 77, 161–164. https://doi.org/10.2317/0310.23.1 Sheffield, C.S., Wilkes, M.A., Cutler, C.G. & Hermanutz, L. (2015) An artificial nesting substrate for Osmia species that nest under stones, with focus on Osmia inermis (Hymenoptera: Megachilidae). Insect Conservation and Diversity, 8, 189–192. https://doi.org/10.1111/icad.12095 Shinn, A.F. (1967) A revision of the bee genus Calliopsis and the biology and ecology of C. andreniformis (Hymenoptera, Andrenidae). The University of Kansas Science Bulletin, 46, 753–936. https://doi.org/10.5962/bhl.part.20081 Sladen, F.W.L. (1915) Characters separating the species of the bee genus Cœlioxys occurring in Ontario. The Canadian Entomologist, 47, 205–208. https://doi.org/10.4039/ent47205-7 Sladen, F.W.L. (1918) Pollination of alfalfa by bees of the genus Megachile. Table of Canadian species of the latimanus group. The Canadian Entomologist, 50, 301–304. https://doi.org/10.4039/ent50301-9 Smith, A.A., Bentley, M. & Reynolds, H.L. (2013) Wild Bees Visiting Cucumber on Midwestern U.S. Organic Farms Benefit From Near-Farm Semi-Natural Areas. Journal of Economic Entomology, 106, 97–106. https://doi.org/10.1603/ec12007 Smith, F. (1854) Catalogue of hymenopterous insects in the collection of the British Museum. Part II. Apidae. British Museum, London, 465 pp. https://doi.org/10.5962/bhl.title.20999 Smith, J.B. (1901) Notes on some digger bees.―I. Journal of the New York Entomological Society, 9, 29–40. Snelling, R.R. (1966a) A new species of Heteranthidium from California (Hymenoptera: Megachilidae). Los Angeles County Museum Contributions in Science, 97, 1–8. Snelling, R.R. (1966b) Studies on North American bees of the genus Hylaeus. 3. The Nearctic subgenera (Hymenoptera: Colletidae). Bulletin of the Southern California Academy of Sciences, 65(3), 164–175. Snelling, R.R. (1968) Studies on North American bees of the genus Hylaeus. 4. The subgenera Cephalylaeus, Metziella and Hylaeana (Hymenoptera: Colletidae). Los Angeles County Museum Contributions in Science, 144, 1–6. Snelling, R.R. (1970) Studies on North American bees of the genus Hylaeus. 5. The subgenera Hylaeus, s. str. and Paraprosopis (Hymenoptera: Colletidae). Los Angeles County Museum Contributions in Science, 180, 1–59. Soltani, G.G., Bénon, D., Alvarez, N. & Praz, C.J. (2017) When different contact zones tell different stories: Putative ring species in the Megachile concinna species complex (Hymenoptera: Megachilidae). Biological Journal of the Linnean Society, 121, 815–832. https://doi.org/10.1093/biolinnean/blx023 Soucy, S.L. (2002) Nesting biology and socially polymorphic behavior of the sweat bee Halictus rubicundus (Hymenoptera: Halictidae). Annals of the Entomological Society of America, 95, 57–65. https://doi.org/10.1603/0013-8746(2002)095[0057:nbaspb]2.0.co;2 Stehr, F.W. & Nielsen, M.C. (1994) Roland L. Fischer. American Entomologist, 40, 62–63. https://doi.org/10.1093/ae/40.1.62b Stephen, W.P. (1954) A revision of the bee genus Colletes in America North of Mexico. The University of Kansas Science Bulletin, 36, 149–527. Stephen, W.P. (1955) Alfalfa pollination in Manitoba. Journal of Economic Entomology, 48, 543–548. https://doi.org/10.1093/jee/48.5.543 Stephen, W.P. (1956) Notes on the biologies of Megachile frigida Smith and M. inermis Provancher (Hymenoptera: Megachilidae). The Pan-Pacific Entomologist, 32, 95–101. Stephen, W.P. (1966a) Andrena (Cryptandrena) viburnella. I. Bionomics. Journal of the Kansas Entomological Society, 39, 42– 51. Stephen, W.P. (1966b) Andrena (Cryptandrena) viburnella. II. External morphology of the larva and pupa. Journal of the Kansas Entomological Society, 39, 51–53. Stephen, W.P. (2003) Solitary bees in North American agriculture: A perspective. In: Strickler, K. & Cane, J.H. (Eds.), For Nonnative Crops, Whence Pollinators of the Future? Entomological Society of America, Lanham, Maryland, pp. 41–66. Stephen, W.P. & Rao, S. (2007) Sampling native bees in proximity to a highly competitive food resource (Hymenoptera: Apiformes). Journal of the Kansas Entomological Society, 80, 369–376. https://doi.org/10.2317/0022-8567(2007)80[369:snbipt]2.0.co;2 Stevens, O.A. (1921) Further notes on evening flowers, panurgine and halictine bees. The Canadian Entomologist, 53, 65–69. https://doi.org/10.4039/ent5365-3 Stockhammer, K.A. (1966) Nesting habits and life cycle of a sweat bee, Augochlora pura (Hymenoptera: Halictidae). Journal of the Kansas Entomological Society, 39, 157–192.



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(1972) Observations préliminaires sur la biologie d’Osmia (Chalcosmia) coerulescens L., (Hymenoptera Megachilidae), pollinisatrice de la luzerne (Medicago sativa L.). Apidologie, 3, 149–165. https://doi.org/10.1051/apido:19720203 Tepedino, V.J. & Frohlich, D.R. (1982) Mortality factors, pollen utilization, and sex ratio in Megachile pugnata Say (Hymenoptera: Megachilidae), a candidate for commercial sunflower pollination. Journal of the New York Entomological Society, 90, 269–274. Terrell, E.E. & Batra, S.W.T. (1984) Insects collect pollen of eastern wildrice, Zizania aquatica (Poaceae). Castanea, 49, 32– 34. Thomson, J.D., Maddison, W.P. & Plowright, R.C. (1982) Behavior of bumble bee pollinators of Aralia hispida Vent. (Araliaceae). Oecologia, 54, 326–336. https://doi.org/10.1007/bf00380001 Thorp, R.W. (1996) Resource overlap among native and introduced bees in California. In: Matheson, A., Buchmann, S.L., O’Toole, C., Westrich, P. & Williams, I.H. (Eds.), The Conservation of Bees. 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(1967) New species of Pseudopanurgus from Arizona (Hymenoptera, Apoidea). American Museum Novitates, 2298, 1–23. Timberlake, P.H. (1968) A revisional study of the bees of the genus Perdita F. Smith, with special reference to the fauna of the Pacific coast (Hymenoptera, Apoidea). Part VII. University of California Publications in Entomology, 49, 1–196. Timberlake, P.H. (1969) A contribution to the systematics of North American species of Synhalonia (Hymenoptera, Apoidea). University of California Publications in Entomology, 57, 1–76. Timberlake, P.H. (1973) Revision of the genus Pseudopanurgus of North America. University of California Publications in Entomology, 72, i–vi + 1–58. Timberlake, P.H. (1976) Revision of the North American bees of the genus Protandrena Cockerell (Hymenoptera: Apoidea). Transactions of the American Entomological Society, 102, 133–227. Tonietto, R.K. & Ascher, J.S. 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Molecular phylogenetics and evolution, 103, 245–259. https://doi.org/10.1016/j.ympev.2016.07.004 Tucker, E.M. & Rehan, S.M. (2016) Wild bee pollination networks in northern New England. Journal of Insect Conservation, 20, 325–337. https://doi.org/10.1007/s10841-016-9870-1 Tuell, J.K., Ascher, J.S. & Isaacs, R. (2009) Wild bees (Hymenoptera: Apoidea: Anthophila) of the Michigan highbush blueberry agroecosystem. Annals of the Entomological Society of America, 102, 275–287. https://doi.org/10.1603/008.102.0209 Tuell, J.K., Fiedler, A.K., Landis, D. & Isaacs, R. (2008) Visitation by wild and managed bees to eastern US native plants for use in conservation programs. Environmental Entomology, 37, 707–718. https://doi.org/10.1603/0046-225X(2008)37[707:VBWAMB]2.0.CO;2 Tuell, J.K. & Isaacs, R. (2009) Elevated pan traps to monitor bees in flowering crop canopies. Entomologia Experimentalis et


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APPENDIX 1. Species expected to occur in Michigan based on known ranges The following species are expected to occur in Michigan based on their occurrence in multiple neighboring states or provinces, but have not yet been reported in the state. ANDRENIDAE Andrena (Trachandrena) quintilis Robertson 1898 Andrena quintilis is widespread in the Northcentral Region, and occurs in Illinois and Wisconsin. It could possibly occur in the southwest LP or western UP. Pseudopanurgus parvus (Robertson 1892) Widely distributed in eastern North America the Great Plains, making Michigan a probable location of occurrence. Pseudopanurgus pauper (Cresson 1878) A specialist on Ceanothus, with a range including New England and west to Minnesota. APIDAE Melecta pacifica Cresson 1878 atlantica Linsley 1948 A widespread species that occurs in Illinois and Ohio. This species could potentially reach southern Michigan. Nomada affabilis affabilis Cresson 1878 (superba group) The range of N. affabilis includes adjacent states to the south and west of Michigan. It is likely a cleptoparasite of Eucera species, so occurrence in the southeast portion of the state is possible. Nomada ceanothi Cockerell 1907 (ruficornis group) This species has been recorded in Wisconsin and Ontario, making Michigan a likely locality. Due to taxonomic challenges in the genus, this species may have been overlooked. Nomada xanthura Cockerell 1908 Recorded from the northeastern United States, including Wisconsin (Alexander & Schwarz 1994; Wolf & Ascher 2009). JSA considers this a possible senior synonym of N. ochlerata. A number of other Nomada from the eastern United States, including N. gracilis Cresson, N. inepta Mitchell, N. composita, N. detrita Mitchell, N. mendica Mitchell, and the Palaearctic N. flava Panzer are closely related and require comparative study. Nomada inepta and N. gracilis are potential synonyms (Sheffield et al. 2009; Wolf & Ascher 2009). Xeromelecta interrupta (Cresson 1872) A cleptoparasite of Anthophora abrupta, with a range including Illinois, Indiana, and Wisconsin. The range of X. interrupta could include southwestern Michigan. COLLETIDAE Hylaeus (Hylaeus) communis Nylander This European species was recently reported from North America based on specimens from southern Québec, Canada (Martins et al. 2017). As reported by the discoverers of the North American population this species has generalist foraging and habitat preferences. Due to its adaptability it is likely to spread to Michigan and elsewhere in eastern North America. Hylaeus (Paraprosopis) pictipes Nylander This species is newly recorded from North America (Gibbs & Dathe 2017). It was first identified in 2015 from Ontario, but has since been recorded in Pennsylvania and Ohio. It has likely been in Ohio since at least 2011 (S. Droege, pers. comm.) Hylaeus (Spatulariella) punctatus (Brullé) This introduced species of European origin occurs in both Chicago, Illinois near the southwestern corner of Michigan (Ksiazek et al. 2014) and approximately 700 km east by northeast in Toronto, Ontario (Sheffield et al. 2011a). Furthermore, it is now known to occur widely in North America including Colorado (Scott et al., 2011; see also http://bugguide.net/node/view/ 337636) and California (where first detected in North America). Thus, it is probable that the species also occurs in urban areas of Michigan, but no specimens are yet known to us. A series of Hylaeus recently collected from Detroit by M.C. Simao (University of Michigan) were examined, but no H. punctatus were found among them. HALICTIDAE Augochloropsis (Paraugochloropsis) sumptuosa (Smith 1853)



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The range of A. sumptuosa includes much of the eastern United States and extends north into Wisconsin (Wolf & Ascher 2009). The southeastern corner of Michigan is a possible area of occurrence. Lasioglossum (Dialictus) katherineae Gibbs 2011 The range of this species includes Iowa, Wisconsin and Maine and extends south to Georgia (Zarrillo et al. 2016). This species is rare in collections with the exception of a sandplain site in the Montague Plains Wildlife Management Area, Franklin County, Massachusetts (Zarrillo et al. 2016). Lasioglossum (Hemihalictus) pectinatum (Robertson 1890) Lasioglossum pectinatum is uncommonly collected, likely due to its specialization on Physalis, a plant with rather obscure flowers (Gibbs et. al. 2013; Voss & Reznicek 2012). It occurs in the northeastern United States and west to southeastern Minnesota (JG, unpublished). Additional effort to collect from the host plant would help clarify the range of this uncommon bee. Sphecodes solonis Graenicher 1911 This northern species occurs in New England, New York, Minnesota, Wisconsin and much of Canada, suggesting that it likely occurs in the UP. MEGACHILIDAE Coelioxys (Boreocoelioxys) banksi Crawford 1914 There are no known specimens for Michigan, however it has been recorded in Wisconsin, southern Ohio and southern Ontario (Baker 1975), and more recently from Illinois (Tonietto and Ascher, 2009), which strongly suggests it occurs in the state. Coelioxys banksi has been reared from M. relativa nests using trap-nests (Medler & Koerber 1958). Coelioxys (Glyptocoelioxys) germanus Cresson 1878 Ascher et al. (2014) speculated that C. germanus was a cleptoparasite of M. petulans, based on the subgeneric range limits and matching flight seasons. The new record of M. petulans for Michigan raises the possibility that C. germanus is also present; it is currently known from adjacent states to the south. Paranthidium (Paranthidium) jugatorium jugatorium (Say) The genus Paranthidium has not been recorded from Michigan. This distinctive resin bee is oligolectic on Asteraceae, with a preference for Helianthus. It may occur in southern Michigan, based on its widespread distribution that includes Minnesota, Wisconsin, Indiana, Ohio and New York State.


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APPENDIX 2. Species excluded from the Michigan list of bees Several validly described bee species have been previously recorded from Michigan, but reported occurrence in the state is based on doubtful or erroneous identifications or otherwise cannot be confirmed. We enumerate these excluded species below. ANDRENIDAE Andrena (Callandrena sensu lato) helianthiformis Viereck and Cockerell, 1914 Two specimens collected from Helianthus strumosis L. were recorded from Ingham County (Tuell et al. 2008). These specimens, deposited at MSUC: RI, were re-examined by JG and determined to be A. helianthi. Andrena (Melandrena) sayi Robertson 1891 Mitchell (160) record. JG found a short series (MSUC) with a Mitchell determination label to be A. commoda. The closest record from Bouseman and LaBerge (1979) is from Carroll County, Indiana. Andrena (Parandrena) nida Mitchell 1960 This species, associated with sandbar willow, Salix exigua Nutt. (JSA, unpublished), is recorded from Michigan by a single male paratype from Newaygo County deposited at MSUC. The specimen, cited as nida in the revision (LaBerge and Ribble, 1972), was reexamined by JG and compared to another male A. nida paratype from Vicksburg, Mississippi. The Michigan paratype is actually A. andrenoides based on the punctation of the clypeus and sculpturing of the metapostnotum. The mandible is entirely dark, but the Michigan paratype is also stylopized, which might explain why this character does not match a typical A. andrenoides. Material examined. Newaygo Co.: (no locality), 13 May 1956, R. & K. Dreisbach (1 paratype ♂ MSUC). Additional material examined. MISSISSIPPI: Warren Co.: Vicksburg, 11 Mar. 1955, R.R. Dreisbach (1 paratype ♂ MSUC). Andrena (Scrapteropsis) fenningeri Viereck 1922 This species was recorded from Michigan by Mitchell (1960). Several males identified by him are deposited at MSUC, but Mitchell incorrectly associated the males of this species and described the males of A. fenningeri separately as A. verna Mitchell (LaBerge 1971). As a result, Mitchell’s records seem unreliable. A specimen from Crawford Co., incorrectly labeled as a paratype of A. verna is at MSUC, but it is not conspecific with A. verna paratypes from other states. Mitchell (1960) did not include any Michigan material in his description of A. verna. Andrena fenningeri reaches northern Indiana, so it may eventually be found in southwest Michigan. Perdita (Perdita) drymariae Timberlake 1960 Two female paratypes from Baraga County are deposited at MSUC. These specimens are so disjunct from the remaining members of the species in New Mexico and Mexico, that these undoubtedly represent a case of mislabeled specimens (Timberlake 1968). The paratypes do not match any of the other Perdita species from Michigan, and key easily to P. drymariae in Mitchell (1960). Perdita drymariae belongs to the sphaeralceae species group. No members of the sphaeralceae species group occur east of the Mississippi River (Hurd 1979). APIDAE Anthophora (Lophanthophora) ursina Cresson 1869 Fiedler et al. (2012) recorded this species from the MSU MacCready Reserve in Jackson County. Voucher material from this study (MSUC: DL) was re-examined and the specimen in question was determined to be a male Melissodes desponsus. Eucera (Synhalonia) belfragei (Cresson 1872) Recorded by Mitchell (1962). This species is poorly documented outside of Texas. The nearest fully confirmed record with details available is from Parke County in western Indiana (Timberlake 1969). Melissodes (Melissodes) comptoides Robertson 1898 County records: none. Recorded by Mitchell (1962). This species occurs in the Great Plains and the southeastern states. The type locality is Carlinville in Southern Illinois, but an INHS record determined by LaBerge is north of Chicago suggesting that occurrence as far north as Michigan is plausible. The species is included based on Mitchell’s record and its probable occurrence in southwest Michigan. Nomada florilega Lovell and Cockerell 1905 (ruficornis group) County records: Livingston, Cheboygan. Notes. Recorded by Mitchell (1962), and a specimen (MSUC) with his determination label does match his key, but this species has not been recognized routinely by currently active workers and its status requires clarification. An OSUC specimen identified by P. H. Timberlake has not been examined.



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Nomada subrutila Lovell and Cockerell 1905 (ruficornis group) County records: Livingston. Notes. A record by Evans (1986) is unverified. A Mitchell determined specimen from Washtenaw County appears to be N. luteoloides. Nomada texana Cresson 1872 (vegana group) This species has been reported to occur in Michigan (Mitchell 1962), but no specific locality or specimens were indicated. A recent study of N. texana and related species suggests this species has a western and southern distribution, although it does reach as close to Michigan as northern Indiana (Droege et al. 2010). We know of no material from Michigan. Specimens identified as N. texana at MSUC were found to be either N. fervida Smith or N. tiftonensis Cockerell. HALICTIDAE Augochlorella neglectula Cockerell 1897 This species was recorded by Dreisbach (1945), but the species reaches its northern limits in southwestern USA (Coelho 2004; Ordway 1966a). Lasioglossum (Dialictus) nymphale (Smith 1853) Recorded from Michigan by Mitchell (1960) and Tuell et al. (2009) based on misidentified individuals of L. pictum. In the literature, L. nymphale has been recorded as far as Minnesota (Moure & Hurd 1987), but it is a southeastern species, which extends north only along the east coast (Gibbs 2011). Lasioglossum (Dialictus) simplex (Robertson 1901) Apparently a social parasite of L. versatum (Michener 1966, 1978), but see Gibbs (2011). No Michigan specimens have been examined, but it is was recorded from Michigan by Mitchell (1960). Due to identification issues this record is deemed questionable, but the type locality in Carlinville, Illinois and the widespread distribution of its putative host, make its occurrence in the state plausible. MEGACHILIDAE Osmia (Diceratosmia) subfasciata Cresson, 1872 Osmia subfasciata is a primarily southern species, ranging north to the southern tier of Indiana counties (Jean 2010). Eight males of O. subfasciata were recorded from Michigan by Tuell et al. (2009). Only one of these specimens could be found at MSUC and was determined by JG to be a male O. conjuncta. The same study reported 6 females and 3 males of O. conjuncta and a number of undetermined Osmia. Thirty-nine additional O. conjuncta males were found among this material. Both O. conjuncta and O. subfasciata have distinctive lateral emarginations of metasomal tergum 6, so it seems most likely that all records of O. subfasciata from Michigan were misidentified O. conjuncta.


GIBBS ET AL.



147

NMNH (HT) NMNH (HT)

Mitchell, 1960: 233 Mitchell, 1960: 230 Mitchell, 1960: 212 Mitchell, 1960: 181 Mitchell, 1960: 177 Mitchell, 1960: 111 Cockerell, 1916: 9

Andrena (Gonandrena) monroensis

Andrena (Leucandrena) bifurcata

Andrena (?Leucandrena) chippewaensis Andrena (Trachandrena) compacta

Andrena (Trachandrena) arenakensis

Andrena (Bythandrena) acra

Bombus ternarius var. expallidus

Depository

Andrena (Thysandrena) crenata

NMNH (HT)

Mitchell, 1962: 414 Mitchell, 1962: 386 Mitchell, 1962: 396. Mitchell, 1962: 431 Mitchell, 1962: 473 Cockerell, 1916: 2 Mitchell, 1960: 391. Mitchell, 1960: 443 Mitchell, 1960: 440 Crawford, 1906: 5 Mitchell, 1960: 448. Mitchell, 1960: 439 Mitchell, 1960: 419 Mitchell, 1960: 351 Mitchell, 1960: 438 Robertson, 1897: 319 Mitchell, 1962: 92 Mitchell, 1962: 45

Nomada dreisbachi

Nomada (Heminomada) colorata

Nomada (?Heminomada) ochlerata

Nomada sobrina

Triepeolus michiganensis

Prosopis gaigei

Dialictus dreisbachi

Dialictus solidaginis

Dialictus otsegoensis

Halictus lineatulus*

Paralictus michiganensis

Dialictus muskegonensis

Dialictus subversans

Evylaeus divergenoides

Dialictus lepidus

Sphecodes davisii

Osmia (Nothosomia) michiganensis

Stelis michiganensis

* Type locality is uncertain, but Michigan seems probable.

NMNH (HT)

Mitchell, 1962: 382.

Nomada (?Heminomada) autumnalis

NMNH (HT)

NMNH (HT)

INHS (HT)

NMNH (HT)

NMNH (HT)

NMNH (HT)

NMNH (HT)

NMNH? (HT)

NMNH (ST)

NMNH (LT)

NCSU? (HT)

NMNH (HT)

NMNH (HT)

NMNH (HT)

NMNH (HT)

UMMZ (HT)

NMNH? (HT)

?UMMZ (HT)

SEMC (HT)

LaBerge, 1961: 568 Mitchell, 1962: 45

Melissodes (Eumelissodes) subillata

Epeolus lanhami

NMNH (HT)

NMNH (HT)

NMNH (HT)

NMNH (HT)

NMNH (HT)

NMNH (HT)

Authority Mitchell, 1960: 221

Name

R.R. Dreisbach

R.R. Dreisbach

Robertson, 1897

R.R. Dreisbach

R.R. Dreisbach

R.R. Dreisbach

R.R. Dreisbach

Beebe

R.H. Wolcott

R.R. Dreisbach

R.R. Dreisbach

R. & K. Dreisbach

F.M. Gaige

R.R. Dreisbach

R.R. Dreisbach

I.J. Cantrall

R. & K. Dreisbach

A.W. Andrews

U.N. Lanham

L.H. Shinners

F.M. Gaige

R.L. Fischer

R.R. Dreisbach

R. & K. Dreisbach

R. & K. Dreisbach

R.R. Dreisbach

R.R. Dreisbach

R.R. Dreisbach

Collector

Stelis (Stelis) foederalis Smith, 1854; new synonymy

Osmia (Melanosmia) subarctica; new synonmy

Sphecodes davisii

Lasioglossum (Dialictus) viridatum (Lovell 1905); syn. by Mitchell (1962)

Lasioglossum (Dialictus) versans (Lovell 1905); syn. by Gibbs (2011).

Lasioglossum (Dialictus) subversans

Lasioglossum (Dialictus) pictum (Crawford, 1902); syn. by Mitchell (1962).

Lasioglossum (Dialictus) michiganense

Lasioglossum (Dialictus) lineatulum (Crawford, 1906)

Lasioglossum (Dialictus) leucocomum (Lovell 1908); syn. by Gibbs (2011).

Lasioglossum (Dialictus) laevissimum (Smith, 1853); syn. by Gibbs (2010).

Lasioglossum (Dialictus) dreisbachi

Hylaeus (Prosopis) gaigei

Triepeolus michiganensis

Nomada sobrina

Nomada luteoloides Robertson, 1895; syn. by Schwarz and Gusenleitner (2004). Nomada ochlerata

Nomada dreisbachi

Nomada autumnalis Mitchell, 1962

Epeolus americanus (Cresson, 1878); syn. by Onuferko (2017).

Melissodes (Eumelissodes) subillatus

Andrena (Trachandrena) hippotes Robertson, 1895; syn. by LaBerge (1973). Andrena (Tylandrena) wilmattae Cockerell 1906; syn. by LaBerge and Bouseman (1970). Bombus (Pyrobombus) ternarius Say 1837; syn. by Mitchell (1962).

Andrena (Trachandrena) ceanothi Viereck 1917; syn. by LaBerge (1973).

Andrena (Simandrena) wheeleri Graenicher, 1904; new synonymy

Andrena (Euandrena) nigrihirta (Ashmead, 1890); syn. by LaBerge and Ribble (1975). Andrena (Gonandrena) platyparia Robertson, 1895; syn. by LaBerge and Ribble (1972). Andrena (Ptilandrena) erigeniae Robertson, 1891; syn. by LaBerge (1987).

Current status

SUPPLEMENTARY TABLE 1. Michigan name-bearing type specimens including holotypes (HT), lectotypes (LT) and syntypes (ST).

SUPPLEMENTARY TABLE 2. Checklist of the bees of Michigan Family

Genus

Subgenus

Species

Authority

ANDRENIDAE ANDRENINAE Andrenini Andrena

(Andrena)

carolina

Viereck 1909

Andrena

(Andrena)

clarkella

(Kirby 1802)

Andrena

(Andrena)

frigida

Smith 1853

Andrena

(Andrena)

mandibularis

Robertson 1892

Andrena

(Andrena)

milwaukeensis

Graenicher 1903

Andrena

(Andrena)

rufosignata

Cockerell 1902

Andrena

(Andrena)

thaspii

Graenicher 1903

Andrena

(Andrena)

tridens

Robertson 1902

Andrena

(Callandrena s.l.)

aliciae

Robertson 1891

Andrena

(Callandrena s.l.)

asteris

Robertson 1891

Andrena

(Callandrena s.l.)

gardineri

Cockerell 1906

Andrena

(Callandrena s.l.)

helianthi

Robertson 1891

Andrena

(Callandrena s.l.)

krigiana

Robertson 1901

Andrena

(Callandrena s.l.)

placata

Mitchell 1960

Andrena

(Callandrena s.l.)

rudbeckiae

Robertson 1891

Andrena

(Callandrena s.l.)

simplex

Smith 1853

Andrena

(Cnemidandrena)

canadensis

Dalla Torre 1896

Andrena

(Cnemidandrena)

chromotricha

Cockerell 1899

Andrena

(Cnemidandrena)

hirticincta

Provancher 1888

Andrena

(Cnemidandrena)

nubecula

Smith 1853

Andrena

(Cnemidandrena)

parnassiae

Cockerell 1902

Andrena

(Cnemidandrena)

peckhami

Cockerell 1902

Andrena

(Cnemidandrena)

robervalensis

Mitchell 1960

Andrena

(Conandrena)

bradleyi

Viereck 1907

Andrena

(Euandrena)

algida

Smith 1853

Andrena

(Euandrena)

geranii

Robertson 1891

Andrena

(Euandrena)

nigrihirta

(Ashmead 1890)

Andrena

(Euandrena)

polemonii

Robertson 1891

Andrena

(Gonandrena)

fragilis

Smith 1853

Andrena

(Gonandrena)

integra

Smith 1853

Andrena

(Gonandrena)

persimulata

Viereck 1917

Andrena

(Gonandrena)

platyparia

Robertson 1895

Andrena

(Holandrena)

cressonii cressonii

Robertson 1891

Andrena

(Iomelissa)

violae

Robertson 1891

Andrena

(Larandrena)

miserabilis

Cresson 1872

Andrena

(Leucandrena)

barbilabris

(Kirby 1802)

Andrena

(Leucandrena)

erythronii

Robertson 1891

Andrena

(Melandrena)

barbara

Bouseman & LaBerge 1979

Andrena

(Melandrena)

carlini

Cockerell 1901 ......continued on the next page


GIBBS ET AL.

SUPPLEMENTARY TABLE 2. (Continued) Family

Genus

Subgenus

Species

Authority

Andrena

(Melandrena)

commoda

Smith 1879

Andrena

(Melandrena)

confederata

Viereck 1917

Andrena

(Melandrena)

dunningi

Cockerell 1898

Andrena

(Melandrena)

hilaris

Smith 1853

Andrena

(Melandrena)

nivalis

Smith 1853

Andrena

(Melandrena)

pruni

Robertson 1891

Andrena

(Melandrena)

regularis

Malloch 1917

Andrena

(Melandrena)

vicina

Smith 1853

Andrena

(Micrandrena)

illinoiensis

Robertson 1891

Andrena

(Micrandrena)

melanochroa

Cockerell 1898

Andrena

(Micrandrena)

neonana

Viereck 1917

Andrena

(Micrandrena)

nigrae

Robertson 1905

Andrena

(Micrandrena)

personata

Robertson 1897

Andrena

(Micrandrena)

salictaria

Robertson 1905

Andrena

(Micrandrena)

ziziae

Robertson 1891

Andrena

(Parandrena)

andrenoides

(Cresson 1878)

Andrena

(Parandrena)

wellesleyana

Robertson 1897

Andrena

(Plastandrena)

crataegi

Robertson 1893

Andrena

(Ptilandrena)

distans

Provancher 1888

Andrena

(Ptilandrena)

erigeniae

Robertson 1891

Andrena

(Rhacandrena)

brevipalpis

Cockerell 1930

Andrena

(Rhacandrena)

robertsonii

Dalla Torre 1896

Andrena

(Scaphandrena)

arabis

Robertson 1897

Andrena

(Scrapteropsis)

alleghaniensis

Viereck 1907

Andrena

(Scrapteropsis)

imitatrix

Cresson 1872

Andrena

(Scrapteropsis)

morrisonella

Viereck 1917

Andrena

(Simandrena)

nasonii

Robertson 1895

Andrena

(Simandrena)

wheeleri

Graenicher 1904

Andrena

(Taeniandrena)

wilkella

(Kirby 1802)

Andrena

(Thysandrena)

bisalicis

Viereck 1908

Andrena

(Thysandrena)

w-scripta

Viereck 1904

Andrena

(Trachandrena)

ceanothi

Viereck 1917

Andrena

(Trachandrena)

forbesii

Robertson 1891

Andrena

(Trachandrena)

heraclei

Robertson 1897

Andrena

(Trachandrena)

hippotes

Robertson 1895

Andrena

(Trachandrena)

mariae

Robertson 1891

Andrena

(Trachandrena)

miranda

Smith 1879

Andrena

(Trachandrena)

nuda

Robertson 1891

Andrena

(Trachandrena)

rehni

Viereck 1907

Andrena

(Trachandrena)

rugosa

Cockerell 1906

Andrena

(Trachandrena)

sigmundi

Cockerell 1902

Andrena

(Trachandrena)

spiraeana

Robertson 1895 ......continued on the next page



149


Genus

Subgenus

Species

Authority

Andrena

(Trachandrena)

virginiana

Mitchell 1960

Andrena

(Tylandrena)

erythrogaster

(Ashmead 1890)

Andrena

(Tylandrena)

perplexa

Smith 1853

Andrena

(Tylandrena)

wilmattae

Cockerell 1906

Callopsis

(Calliopsis)

andreniformis

Smith 1853

Callopsis

(Verbenapis)

nebraskensis

Crawford 1902

Perdita

(Cockerellia)

albipennis pallidipennis

Cresson 1868 / Graenicher 1910

Perdita

(Cockerellia)

bequaerti indianensis

Viereck 1917

Perdita

(Perdita)

gerhardi gerhardi

Viereck 1904

Perdita

(Perdita)

halictoides

Smith 1853

Perdita

(Perdita)

maculigera maculipennis

Cockerell 1896 / Graenicher 1914

Perdita

(Perdita)

octomaculata octomaculata

(Say 1824)

Perdita

(Perdita)

swenki

Crawford 1915

Protandrena

bancrofti

Dunning 1897

Protandrena

cockerelli

Dunning 1897

Pseudopanurgus

aestivalis aestivalis

(Provancher 1882)

Pseudopanurgus

albitarsis

(Cresson 1872)

Pseudopanurgus

andrenoides

(Smith 1853)

Pseudopanurgus

compositarum

(Robertson 1893)

Pseudopanurgus

labrosiformis

(Robertson 1898)

Pseudopanurgus

rugosus

(Robertson 1895)

PANURGINAE Callopsini

Perditini

Protandrenini

APIDAE APINAE Anthophorini Anthophora

(Clisodon)

terminalis

Cresson 1869

Anthophora

(Melea)

abrupta

Say 1837

Anthophora

(Melea)

bomboides

Kirby 1837

Anthophora

(Mystacanthophora)

walshii

Cresson 1869

laboriosa

(Fabricius 1804)

Habropoda Apini Apis

(Apis)

mellifera

Linnaeus 1758

Bombus

(Bombias)

auricomus

(Robertson 1903)

Bombus

(Bombus)

affinis

Cresson 1863

Bombus

(Bombus)

terricola

Kirby 1837

Bombus

(Cullumanobombus)

fraternus

(Smith 1854)

Bombini

......continued on the next page


GIBBS ET AL.


Genus

Subgenus

Species

Authority

Bombus

(Cullumanobombus)

griseocollis

(DeGeer 1773)

Bombus

(Cullumanobombus)

rufocinctus

Cresson 1863

Bombus

(Psithyrus)

ashtoni

(Cresson 1864)

Bombus

(Psithyrus)

citrinus

(Smith 1854)

Bombus

(Psithyrus)

fernaldae

(Franklin 1911)

Bombus

(Psithyrus)

insularis

(Smith 1861)

Bombus

(Pyrobombus)

bimaculatus

Cresson 1863

Bombus

(Pyrobombus)

frigidus

Smith 1854

Bombus

(Pyrobombus)

impatiens

Cresson 1863

Bombus

(Pyrobombus)

perplexus

Cresson 1863

Bombus

(Pyrobombus)

sandersoni

Franklin 1913

Bombus

(Pyrobombus)

ternarius

Say 1837

Bombus

(Pyrobombus)

vagans vagans

Smith 1854

Bombus

(Subterraneobombus)

borealis

Kirby 1837

Bombus

(Thoracobombus)

fervidus

(Fabricius 1798)

Bombus

(Thoracobombus)

pensylvanicus

(DeGeer 1773)

Eucera

(Synhalonia)

atriventris

(Smith 1854)

Eucera

(Synhalonia)

dubitata

(Cresson 1878)

Eucera

(Synhalonia)

hamata

(Bradley 1942)

Florilegus

(Florilegus)

condignus

(Cresson 1878)

Melissodes

(Apomelissodes)

apicatus

Lovell & Cockerell 1906

Melissodes

(Eumelissodes)

agilis

Cresson 1878

Melissodes

(Eumelissodes)

denticulatus

Smith 1854

Melissodes

(Eumelissodes)

dentiventris

Smith 1854

Melissodes

(Eumelissodes)

druriellus

(Kirby 1802)

Melissodes

(Eumelissodes)

illatus


Melissodes

(Eumelissodes)

niveus

Robertson 1895

Melissodes

(Eumelissodes)

subillatus

LaBerge 1961

Melissodes

(Eumelissodes)

tinctus

LaBerge 1961

Melissodes

(Eumelissodes)

trinodis

Robertson 1901

Melissodes

(Eumelissodes)

wheeleri

Cockerell 1906

Melissodes

(Heliomelissodes)

desponsus

Smith 1854

Melissodes

(Melissodes)

bimaculatus bimaculatus

(Lepeletier 1825)

Melissodes

(Melissodes)

communis commnuis

Cresson 1878

Peponapis

(Peponapis)

pruinosa

(Say 1837)

Svastra

(Epimelissodes)

obliqua obliqua

(Say 1837)

Holcopasites

calliopsidis calliopsidis

(Linsley 1943)

Epeolus

ainslei

Crawford 1932

Eucerini

NOMADINAE Ammobatoidini Epeolini ......continued on the next page



151


Genus

Subgenus

Species

Authority

Epeolus

americanus

(Cresson 1878)

Epeolus

autumnalis

Robertson 1902

Epeolus

bifasicatus

Cresson 1864

Epeolus

canadensis

Mitchell 1962

Epeolus

interruptus

Robertson 1900

Epeolus

lectoides

Robertson 1901

Epeolus

minimus

(Robertson 1902)

Epeolus

pusillus

Cresson 1864

Epeolus

scutellaris

Say 1824

Triepeolus

concavus

(Cresson 1878)

Triepeolus

cressonii

(Robertson 1897)

Triepeolus

donatus

(Smith 1854)

Triepeolus

eliseae

new species

Triepeolus

helianthi

(Robertson 1897)

Triepeolus

lunatus

(Say 1824)

Triepeolus

michiganensis

Mitchell 1962

Triepeolus

pectoralis

(Robertson 1897)

Triepeolus

remigatus

(Fabricius 1804)

Triepeolus

simplex

Robertson 1903

Nomada

alpha alpha

Cockerell 1905a

Nomada

armatella

Cockerell 1903

Nomada

articulata

Smith 1854

Nomada

australis

Mitchell 1962

Nomada

autumnalis

Mitchell 1962

Nomada

banksi

Cockerell 1907

Nomada

bella

Cresson 1863

Nomada

besseyi

Swenk 1913

Nomada

bethunei

Cockerell 1903

Nomada

binotata

(Robertson 1903)

Nomada

composita

Mitchell 1962

Nomada

cressonii

Robertson 1893

Nomada

cuneata

(Robertson 1903)

Nomada

decepta

Mitchell 1962

Nomada

dentariae

(Robertson 1903)

Nomada

denticulata

Robertson 1902

Nomada

depressa

Cresson 1863

Nomada

dreisbachi

Mitchell 1962

Nomada

erigeronis

Robertson 1897

Nomada

fervida

Smith 1854

Nomada

gracilis

Cresson 1863

Nomada

illinoensis

Robertson 1900

Nomadini



GIBBS ET AL.


Genus

Subgenus

Species

Authority

Nomada

imbricata

Smith 1854

Nomada

lehighensis

Cockerell 1903

Nomada

lepida

Cresson 1863

Nomada

luteoloides

Robertson 1895

Nomada

maculata

Cresson 1863

Nomada

obliterata

Cresson 1863

Nomada

ochlerata

Mitchell 1962

Nomada

ovata

(Robertson 1903)

Nomada

parva

Robertson 1900

Nomada

perplexa

Cresson 1863

Nomada

placida

Cresson 1863

Nomada

pygmaea

Cresson 1863

Nomada

quadrimaculata

(Robertson 1903)

Nomada

rubicunda

Olivier 1811

Nomada

sayi

Robertson 1893

Nomada

sobrina

Mitchell 1962

Nomada

sphaerogaster

Cockerell 1903

Nomada

superba

Cresson 1863

Nomada

tiftonensis

Cockerell 1903

Nomada

vegana

Cockerell 1903

Nomada

vicina

Cresson 1863

Nomada

vincta

Say 1837

Epeoloides

pilosulus

(Cresson 1878)

Osirini XYLOCOPINAE Ceratinini Ceratina

(Zadontomerus)

calcarata

Robertson 1900

Ceratina

(Zadontomerus)

dupla

Say 1837

Ceratina

(Zadontomerus)

mikmaqi

Rehan & Sheffield 2011

Ceratina

(Zadontomerus)

strenua

Smith 1879

Xylocopa

(Xylocopoides)

virginica virginica

(Linnaeus 1771)

Colletes

aberrans

Cockerell 1897

Colletes

aestivalis

Patton 1879

Colletes

americanus

Cresson 1868

Colletes

banksi

Swenk 1908

Colletes

brevicornis

Robertson 1897

Colletes

compactus compactus

Cresson 1868

Colletes

consors mesocopus

Cresson 1868 / Swenk 1907

Colletes

hyalinus

Provancher 1888

Xylocopini COLLETIDAE COLLETINAE




153


Genus

Subgenus

Species

Authority

Colletes

impunctatus lacustris

Nylander 1852/Swenk 1906

Colletes

inaequalis

Say 1837

Colletes

kincaidii

Cockerell 1898

Colletes

latitarsis

Robertson 1891

Colletes

mandibularis

Smith 1853

Colletes

nudus

Robertson 1898

Colletes

simulans armatus

Cresson 1868 / Patton 1879

Colletes

solidaginis

Swenk 1906

Colletes

thoracicus

Smith 1853

Colletes

validus

Cresson 1868

Colletes

willistoni

Robertson 1891

HYLAEINAE Hylaeus

(Cephalylaeus)

basalis

(Smith 1853)

Hylaeus

(Hylaeus)

annulatus

(Linnaeus 1758)

Hylaeus

(Hylaeus)

fedorica

(Cockerell 1909)

Hylaeus

(Hylaeus)

leptocephalus

(Morawitz 1871["1870"])

Hylaeus

(Hylaeus)

mesillae cressonii

(Cockerell 1896) / (Cockerell 1907)

Hylaeus

(Hylaeus)

rudbeckiae

(Cockerell & Casad 1895)

Hylaeus

(Hylaeus)

saniculae

(Robertson 1896)

Hylaeus

(Hylaeus)

verticalis

(Cresson 1869)

Hylaeus

(Metziella)

sparsus

(Cresson 1869)

Hylaeus

(Paraprosopis)

floridanus

(Robertson 1893)

Hylaeus

(Prosopis)

affinis

(Smith 1853)

Hylaeus

(Prosopis)

gaigei

(Cockerell 1916)

Hylaeus

(Prosopis)

illinoisensis

(Robertson 1896)

Hylaeus

(Prosopis)

modestus modestus

Say 1837

Hylaeus

(Prosopis)

nelumbonis

(Robertson 1890)

Hylaeus

(Spatulariella)

hyalinatus

Smith 1842

Augochlora

(Augochlora)

pura pura

(Say 1837)

Augochlorella

aurata

(Smith 1853)

Augochlorella

persimilis

(Viereck 1910)

HALICTIDAE HALICTINAE Augochlorini

Augochloropsis

(Paraugochloropsis)

metallica fulgida

(Fabricius 1793) / (Smith 1853)

Agapostemon

(Agapostemon)

sericeus

(Förster 1771)

Agapostemon

(Agapostemon)

splendens

(Lepeletier 1841)

Agapostemon

(Agapostemon)

texanus

Cresson 1872

Agapostemon

(Agapostemon)

virescens

(Fabricius 1775)

Halictus

(Nealictus)

parallelus

Say 1837

Halictini



GIBBS ET AL.


Genus

Subgenus

Species

Authority

Halictus

(Odontalictus)

ligatus

Say 1837

Halictus

(Protohalictus)

rubicundus

(Christ 1791)

Halictus

(Seladonia)

confusus confusus

Smith 1853

Lasioglossum

(Dialictus)

abanci

(Crawford 1932)

Lasioglossum

(Dialictus)

achilleae

(Mitchell 1960)

Lasioglossum

(Dialictus)

admirandum

(Sandhouse 1924)

Lasioglossum

(Dialictus)

albipenne

(Robertson 1890)

Lasioglossum

(Dialictus)

anomalum

(Robertson 1892)

Lasioglossum

(Dialictus)

atwoodi

Gibbs 2010

Lasioglossum

(Dialictus)

bruneri

(Crawford 1902)

Lasioglossum

(Dialictus)

callidum

(Sandhouse 1924)

Lasioglossum

(Dialictus)

cattellae

(Ellis 1913)

Lasioglossum

(Dialictus)

ceanothi

(Mitchell 1960)

Lasioglossum

(Dialictus)

coeruleum

(Robertson 1893)

Lasioglossum

(Dialictus)

coreopsis

(Robertson 1902)

Lasioglossum

(Dialictus)

cressonii

(Robertson 1890)

Lasioglossum

(Dialictus)

dreisbachi

(Mitchell 1960)

Lasioglossum

(Dialictus)

ellisiae

(Sandhouse 1924)

Lasioglossum

(Dialictus)

ephialtum

Gibbs 2010

Lasioglossum

(Dialictus)

fattigi

(Mitchell 1960)

Lasioglossum

(Dialictus)

floridanum

(Robertson 1892)

Lasioglossum

(Dialictus)

foveolatum

(Robertson 1902)

Lasioglossum

(Dialictus)

gotham

Gibbs 2011

Lasioglossum

(Dialictus)

hartii

(Robertson 1892)

Lasioglossum

(Dialictus)

heterognathum

(Mitchell 1960)

Lasioglossum

(Dialictus)

hitchensi

Gibbs 2012

Lasioglossum

(Dialictus)

illinoense

(Robertson 1892)

Lasioglossum

(Dialictus)

imitatum

(Smith 1853)

Lasioglossum

(Dialictus)

laevissimum

(Smith 1853)

Lasioglossum

(Dialictus)

leucocomum

(Lovell 1908)

Lasioglossum

(Dialictus)

lineatulum

(Crawford 1906)

Lasioglossum

(Dialictus)

lionotum

(Sandhouse 1923)

Lasioglossum

(Dialictus)

michiganense

(Mitchell 1960)

Lasioglossum

(Dialictus)

nigroviride

(Graenicher 1911)

Lasioglossum

(Dialictus)

novascotiae

(Mitchell 1960)

Lasioglossum

(Dialictus)

oblongum

(Lovell 1905)

Lasioglossum

(Dialictus)

obscurum

(Robertson 1892)

Lasioglossum

(Dialictus)

oceanicum

(Cockerell 1916)

Lasioglossum

(Dialictus)

paradmirandum

(Knerer & Atwood 1966)

Lasioglossum

(Dialictus)

perpunctatum

(Ellis 1913)

Lasioglossum

(Dialictus)

pictum

(Crawford 1902)

Lasioglossum

(Dialictus)

pilosum

(Smith 1853) ......continued on the next page



155


Genus

Subgenus

Species

Authority

Lasioglossum

(Dialictus)

planatum

(Lovell 1905)

Lasioglossum

(Dialictus)

platyparium

(Robertson 1895)

Lasioglossum

(Dialictus)

pruinosum

(Robertson 1892)

Lasioglossum

(Dialictus)

rufulipes

(Cockerell 1938)

Lasioglossum

(Dialictus)

sagax

(Sandhouse 1924)

Lasioglossum

(Dialictus)

sheffieldi

Gibbs 2010

Lasioglossum

(Dialictus)

smilacinae

(Robertson 1899)

Lasioglossum

(Dialictus)

subversans

(Mitchell 1960)

Lasioglossum

(Dialictus)

subviridatum

(Cockerell 1938)

Lasioglossum

(Dialictus)

taylorae

Gibbs 2010

Lasioglossum

(Dialictus)

tegulare

(Robertson 1890)

Lasioglossum

(Dialictus)

tenax

(Sandhouse 1924)

Lasioglossum

(Dialictus)

timothyi

Gibbs 2010

Lasioglossum

(Dialictus)

trigeminum

Gibbs 2011

Lasioglossum

(Dialictus)

versans

(Lovell 1905)

Lasioglossum

(Dialictus)

versatum

(Robertson 1902)

Lasioglossum

(Dialictus)

vierecki

(Crawford 1904)

Lasioglossum

(Dialictus)

viridatum

(Lovell 1905)

Lasioglossum

(Dialictus)

weemsi

(Mitchell 1960)

Lasioglossum

(Dialictus)

zephyrum

(Smith 1853)

Lasioglossum

(Evylaeus)

cinctipes

(Provancher 1888)

Lasioglossum

(Hemihalictus)

birkmanni

(Crawford 1906)

Lasioglossum

(Hemihalictus)

fedorense

(Crawford 1906)

Lasioglossum

(Hemihalictus)

foxii

(Robertson 1895)

Lasioglossum

(Hemihalictus)

inconditum

(Cockerell 1916)

Lasioglossum

(Hemihalictus)

lustrans

(Cockerell 1897)

Lasioglossum

(Hemihalictus)

macoupinense

(Robertson 1895)

Lasioglossum

(Hemihalictus)

nelumbonis

(Robertson 1890)

Lasioglossum

(Hemihalictus)

pectorale

(Smith 1853)

Lasioglossum

(Hemihalictus)

swenki

(Crawford 1906)

Lasioglossum

(Lasioglossum)

acuminatum

McGinley 1986

Lasioglossum

(Lasioglossum)

athabascense

(Sandhouse 1933)

Lasioglossum

(Lasioglossum)

coriaceum

(Smith 1853)

Lasioglossum

(Lasioglossum)

forbesii

(Robertson 1890)

Lasioglossum

(Lasioglossum)

fuscipenne

(Smith 1853)

Lasioglossum

(Lasioglossum)

paraforbesii

McGinley 1986

Lasioglossum

(Leuchalictus)

leucozonium

(Schrank 1781)

Lasioglossum

(Leuchalictus)

zonulum

(Smith 1848)

Lasioglossum

(Sphecodogastra)

comagenense

(Knerer & Atwood 1964)

Lasioglossum

(Sphecodogastra)

oenotherae

(Stevens 1920)

Lasioglossum

(Sphecodogastra)

quebecense

(Crawford 1907)

Lasioglossum

(Sphecodogastra)

seillean

Gibbs & Packer 2013 ......continued on the next page


GIBBS ET AL.


Genus

Subgenus

Species

Authority

Lasioglossum

(Sphecodogastra)

texanum

(Cresson 1872)

Lasioglossum

(Sphecodogastra)

truncatum

(Robertson 1901)

Sphecodes

antennariae

Robertson 1891

Sphecodes

aroniae

Mitchell 1960

Sphecodes

atlantis

Mitchell 1956

Sphecodes

autumnalis

Mitchell 1956

Sphecodes

banksii

Lovell 1909

Sphecodes

clematidis

Robertson 1897

Sphecodes

confertus

Say 1837

Sphecodes

coronus

Mitchell 1956

Sphecodes

cressonii

(Robertson 1903)

Sphecodes

davisii

Robertson 1897

Sphecodes

dichrous

Smith 1853

Sphecodes

fattigi

Mitchell 1956

Sphecodes

heraclei heraclei

Robertson 1897

Sphecodes

illinoensis

(Robertson 1903)

Sphecodes

johnsonii

Lovell 1909

Sphecodes

levis


Sphecodes

mandibularis

Cresson 1872

Sphecodes

minor

Robertson 1898

Sphecodes

pimpinellae

Sphecodes

prosphorus


Sphecodes

pycnanthemi

Robertson 1897

Sphecodes

ranunculi

Robertson 1897

Sphecodes

townesi

Mitchell 1956

heteropoda heteropoda

(Say 1824)

Dufourea

harveyi

(Cresson 1878)

Dufourea

maura

(Cresson 1878)

Dufourea

monardae

(Viereck 1924)

Dufourea

novaeangeliae

(Robertson 1897)

NOMIINAE Dieunomia

(Dieunomia)

ROPHITINAE

MEGACHILIDAE MEGACHILINAE Anthidiini Anthidiellum

(Loyolanthidium)

notatum boreale

(Robertson 1902)

Anthidiellum

(Loyolanthidium)

notatum notatum

(Latreille 1809)

Anthidium

(Anthidium)

manicatum manicatum

(Linnaeus 1758)

Anthidium

(Anthidium)

psoraleae

Robertson 1902

Anthidium

(Anthidium)

tenuiflorae

Cockerell 1907

Anthidium

(Proanthidium)

oblongatum oblongatum

(Illiger 1806)

Dianthidium

(Dianthidium)

simile

(Cresson 1864) ......continued on the next page



157


Genus

Subgenus

Species

Authority

Stelis

(Dolichostelis)

louisae

Cockerell 1911

Stelis

(Stelis)

coarctatus

Crawford 1916

Stelis

(Stelis)

foederalis

Smith 1854

Stelis

(Stelis)

labiata

(Provancher 1888)

Stelis

(Stelis)

lateralis

Cresson 1864

Stelis

(Stelis)

nitida

Cresson 1878

Stelis

(Stelis)

subemarginata

Cresson 1878

Trachusa

(Heteranthidium)

zebrata

(Cresson 1872)

Coelioxys

(Boreocoelioxys)

moestus

Cresson 1864

Coelioxys

(Boreocoelioxys)

octodentatus

Say 1824

Coelioxys

(Boreocoelioxys)

porterae

Cockerell 1900

Coelioxys

(Boreocoelioxys)

rufitarsis

Smith 1854

Coelioxys

(Boreocoelioxys)

sayi

Robertson 1897

Coelioxys

(Coelioxys)

sodalis

Cresson 1878

Coelioxys

(Cyrtocoelioxys)

modestus

Smith 1854

Coelioxys

(Paracoelioxys)

funerarius

Smith 1854

Coelioxys

(Synocoelioxys)

alternatus

Say 1837

Coelioxys

(Xerocoelioxys)

immaculatus

Cockerell 1912

Megachile

(Acentron)

albitarsis

Cresson 1872

Megachile

(Callomegachile)

sculpturalis

Smith 1853

Megachile

(Chelostomoides)

campanulae

(Robertson 1903)

Megachile

(Chelostomoides)

rugifrons

(Smith 1854)

Megachile

(Eutricharaea)

apicalis

Spinola 1808

Megachile

(Eutricharaea)

pusilla

Pérez 1884

Megachile

(Eutricharaea)

rotundata

(Fabricius 1787)

Megachile

(Leptorachis)

petulans

Cresson 1878

Megachile

(Litomegachile)

brevis

Say 1837

Megachile

(Litomegachile)

mendica

Cresson 1878

Megachile

(Litomegachile)

texana

Cresson 1878

Megachile

(Megachile)

centuncularis

(Linnaeus 1758)

Megachile

(Megachile)

inermis

Provancher 1888

Megachile

(Megachile)

lapponica

Thomson 1872

Megachile

(Megachile)

montivaga

Cresson 1878

Megachile

(Megachile)

relativa

Cresson 1878

Megachile

(Megachiloides)

dakotensis

Mitchell 1926

Megachile

(Sayapis)

frugalis frugalis

Cresson 1872

Megachile

(Sayapis)

inimical sayi

Cresson 1872 / Cresson 1878

Megachile

(Sayapis)

pugnata pugnata

Say 1837

Megachile

(Xanthosarus)

addenda

Cresson 1878

Megachile

(Xanthosarus)

frigida frigida

Smith 1853

Megachile

(Xanthosarus)

gemula gemula

Cresson 1878

Megachilini



GIBBS ET AL.


Genus

Subgenus

Species

Authority

Megachile

(Xanthosarus)

latimanus

Say 1823

Megachile

(Xanthosarus)

melanophaea melanophaea

Smith 1853

Megachile

(Xanthosarus)

mucida

Cresson 1878

Megachile

(Xanthosarus)

perihirta

Cockerell 1898

Ashmeadiella

(Ashmeadiella)

bucconis bucconis

(Say 1837)

Chelostoma

(Prochelostoma)

philadelphi

(Robertson 1891)

Heriades

(Neotrypetes)

carinata

Cresson 1864

Heriades

(Neotrypetes)

leavitti

Crawford 1913

Heriades

(Neotrypetes)

variolosa variolosa

(Cresson 1872)

Hoplitis

(Alcidamea)

albifrons albifrons

(Kirby 1837)

Hoplitis

(Alcidamea)

pilosifrons

(Cresson 1864)

Hoplitis

(Alcidamea)

product producta

(Cresson 1864)

Hoplitis

(Alcidamea)

spoliata

(Provancher 1888)

Hoplitis

(Alcidamea)

truncata truncata

(Cresson 1878)

Osmia

(Cephalosmia)

subaustralis

Cockerell 1900

Osmia

(Diceratosmia)

conjuncta

Cresson 1864

Osmia

(Helicosmia)

caerulescens

(Linnaeus 1758)

Osmia

(Helicosmia)

georgica

Cresson 1878

Osmia

(Helicosmia)

texana

Cresson 1872

Osmia

(Melanosmia)

albiventris

Cresson 1864

Osmia

(Melanosmia)

atriventris

Cresson 1864

Osmia

(Melanosmia)

bucephala

Cresson 1864

Osmia

(Melanosmia)

collinsiae

Robertson 1905

Osmia

(Melanosmia)

distincta

Cresson 1864

Osmia

(Melanosmia)

felti

Cockerell 1911

Osmia

(Melanosmia)

inermis

(Zetterstedt 1838)

Osmia

(Melanosmia)

inspergens


Osmia

(Melanosmia)

laticeps

Thomson 1872

Osmia

(Melanosmia)

nigriventris

(Zetterstedt 1838)

Osmia

(Melanosmia)

proxima

Cresson 1864

Osmia

(Melanosmia)

pumila

Cresson 1864

Osmia

(Melanosmia)

simillima

Smith 1853

Osmia

(Melanosmia)

subarctica

Mitchell 1962

Osmiini

Osmia

(Melanosmia)

tarsata

Provancher 1888

Osmia

(Melanosmia)

tersula

Cockerell 1912

Osmia

(Melanosmia)

aff. trevoris

Cockerell 1897

Osmia

(Melanosmia)

virga

Sandhouse 1939

Osmia

(Osmia)

cornifrons

(Radoszkowski 1887)

Osmia

(Osmia)

lignaria lignaria

Say 1837

Osmia

(Osmia)

taurus

Smith 1873 ......continued on the next page



159


Genus

Subgenus

Species

Authority

Macropis

(Macropis)

nuda

(Provancher 1882)

MELITTIDAE MELITTINAE


GIBBS ET AL.