Zootaxa, a Review of the Cleptoparasitic Bee Genus Triepeolus

ZOOTAXA

1710

A review of the cleptoparasitic bee genus Triepeolus (Hymenoptera: Apidae).—Part I

MOLLY G. RIGHTMYER

Magnolia Press Auckland, New Zealand MOLLY G. RIGHTMYER A review of the cleptoparasitic bee genus Triepeolus (Hymenoptera: Apidae).—Part I (Zootaxa 1710) 170 pp.; 30 cm. 22 Feb. 2008 ISBN 978-1-86977-191-1 (paperback) ISBN 978-1-86977-192-8 (Online edition)

FIRST PUBLISHED IN 2008 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

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ISSN 1175-5326 (Print edition) ISSN 1175-5334 (Online edition)

2 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Zootaxa 1710: 1–170 (2008) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ ZOOTAXA Copyright © 2008 · Magnolia Press ISSN 1175-5334 (online edition)

A review of the cleptoparasitic bee genus Triepeolus (Hymenoptera: Apidae).— Part I

MOLLY G. RIGHTMYER Department of Entomology, MRC 188, P. O. Box 37012, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20013-7012 USA [email protected]

Table of contents

Abstract ...... 5 Introduction ...... 6 Materials and methods ...... 7 Morphology ...... 9 Key to the females of North and Central America ...... 11 Key to the males and females of Eastern North America ...... 25 Key to the males and females of South America and the Caribbean...... 29 Taxonomy...... 30 Triepeolus antiguensis Cockerell 1949 ...... 31 Triepeolus antiochensis Rightmyer, n. sp...... 31 Triepeolus argentimus Rightmyer, n. sp...... 32 Triepeolus argus Rightmyer, n. sp...... 33 Triepeolus argyreus (Cockerell) 1907 ...... 35 Triepeolus atripes Mitchell 1962...... 36 Triepeolus aztecus (Cresson) 1878 ...... 37 Triepeolus balteatus Cockerell 1921 ...... 38 Triepeolus bimorulus Rightmyer, n. sp...... 39 Triepeolus brittaini Cockerell 1931 … ...... 40 Triepeolus californicus (Cresson) 1878...... 41 Triepeolus cameroni (Meade-Waldo) 1913 ...... 42 Triepeolus charlesi Rightmyer, n. sp...... 43 Triepeolus circumculus Rightmyer, n. sp...... 44 Triepeolus claytoni Rightmyer, n. sp...... 46 Triepeolus concavus (Cresson) 1878 … ...... 47 Triepeolus cressonii (Robertson) 1897 ...... 48 Triepeolus cruciformis Rightmyer, n. sp...... 49 Triepeolus cuabitensis Genaro 1999 ...... 50 Triepeolus dacotensis (Stevens) 1919 ...... 51 Triepeolus denverensis Cockerell 1910 ...... 51 Triepeolus diffusus Rightmyer, n. sp...... 52 Triepeolus dilutus Rightmyer, n. sp...... 54 Triepeolus distinctus (Cresson) 1878 ...... 55 Triepeolus diversipes Cockerell 1924 ...... 56 Triepeolus donatus (Smith) 1854 ...... 57 Triepeolus edwardi Rightmyer, n. sp...... 58

Accepted by L. Packer: 10 Dec. 2007; published: 22 Feb. 2008 3 Triepeolus eldoradensis (Cockerell) 1910 ...... 59 Triepeolus engeli Rightmyer, n. sp...... 60 Triepeolus epeolurus Rightmyer 2004 ...... 61 Triepeolus exilicurvus Rightmyer, n. sp...... 61 Triepeolus flavigradus Rightmyer, n. sp...... 62 Triepeolus fraserae Cockerell 1904 ...... 64 Triepeolus fulgidus Rightmyer, n. sp...... 64 Triepeolus georgicus Mitchell 1962 ...... 65 Triepeolus grandis (Friese) 1917 ...... 66 Triepeolus griswoldi Rightmyer, n. sp...... 67 Triepeolus helianthi (Robertson) 1897 ...... 68 Triepeolus heterurus (Cockerell & Sandhouse) 1924 ...... 70 Triepeolus interruptus Rightmyer, n. sp...... 71 Triepeolus intrepidus (Smith) 1879 ...... 72 Triepeolus isohedrus Rightmyer, n. sp...... 73 Triepeolus jennieae Rightmyer, n. sp...... 74 Triepeolus joliae Rightmyer, n. sp...... 74 Triepeolus lateralis Rightmyer, n. sp...... 75 Triepeolus laticaudus Cockerell 1921 ...... 77 Triepeolus laticeps (Friese) 1917...... 77 Triepeolus loomisorum Rozen 1989 ...... 79 Triepeolus lunatus (Say) 1824 ...... 80 Triepeolus margaretae Rightmyer, n. sp...... 82 Triepeolus martini (Cockerell) 1900 ...... 82 Triepeolus mauropygus Rightmyer, n. sp...... 83 Triepeolus medusa Cockerell 1917 …...... 84 Triepeolus melanarius Rightmyer, n. sp...... 85 Triepeolus micheneri Rightmyer, n. sp...... 87 Triepeolus michiganensis Mitchell 1962 ...... 88 Triepeolus micropygius Robertson 1903 ...... 88 Triepeolus mitchelli Hurd 1979 ...... 89 Triepeolus mojavensis Linsley 1939 ...... 90 Triepeolus monardae Mitchell 1962 ...... 91 Triepeolus nayaritensis Rightmyer, n. sp...... 91 Triepeolus nevadensis (Cresson) 1878 ...... 92 Triepeolus nigrihirtus Mitchell 1962 ...... 93 Triepeolus nisibonensis Genaro 2001 ...... 94 Triepeolus norae Cockerell 1907 ...... 94 Triepeolus occidentalis (Cresson) 1878 ...... 95 Triepeolus paenepectoralis Viereck 1905 ...... 96 Triepeolus parkeri Rightmyer, n. sp...... 97 Triepeolus partitus Rightmyer, n. sp...... 98 Triepeolus parvidiversipes Rightmyer, n. sp...... 100 Triepeolus parvus Rightmyer, n. sp...... 100 Triepeolus pectoralis (Robertson) 1897 ...... 101 Triepeolus penicilliferus (Brues) 1903 ...... 102 Triepeolus perpictus Rightmyer, n. sp...... 103 Triepeolus phaeopygus Rightmyer, n. sp...... 104 Triepeolus punctoclypeus Rightmyer, n. sp...... 106 Triepeolus quadratus Rightmyer, n. sp...... 107 Triepeolus quadrifasciatus atlanticus Mitchell 1962 ...... 108 Triepeolus quadrifasciatus quadrifasciatus (Say) 1823 ...... 109 Triepeolus remigatus (Fabricius) 1804 ...... 110 Triepeolus robustus (Cresson) 1878 ...... 111

4 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Triepeolus rohweri Cockerell 1911 ...... 112 Triepeolus rufithorax Graenicher 1928 ...... 113 Triepeolus rufoclypeus (Fox) 1891 ...... 114 Triepeolus rugosus Mitchell 1962 ...... 115 Triepeolus scelestus (Cresson) 1878 ...... 115 Triepeolus simulatus Rightmyer, n. sp...... 116 Triepeolus subalpinus Cockerell 1910...... 117 Triepeolus sublunatus Cockerell 1907...... 118 Triepeolus subnitens Cockerell & Timberlake 1929 ...... 119 Triepeolus tanneri Cockerell 1928 ...... 120 Triepeolus tepanecus (Cresson) 1878 ...... 120 Triepeolus texanus (Cresson) 1878 ...... 121 Triepeolus totonacus (Cresson) 1878 ...... 122 Triepeolus townsendi Cockerell 1907 ...... 123 Triepeolus tristis (Smith) 1854 ...... 123 Triepeolus ventralis (Meade-Waldo) 1913 ...... 124 Triepeolus vernus Rightmyer, n. sp...... 125 Triepeolus vicinus (Cresson) 1865...... 126 Triepeolus victori Genaro 1998 ...... 127 Triepeolus warriti Rightmyer, n. sp...... 128 Triepeolus wilsoni (Cresson) 1865 ...... 129 Triepeolus zacatecus (Cresson) 1878 ...... 130 Triepeolus simplex species group ...... 130 Triepeolus verbesinae species group ...... 131 Unplaced species names ...... 131 Acknowledgements ...... 132 References ...... 133 Appendix 1...... 138 Figures ...... 139

Abstract

The cleptoparasitic bee genus Triepeolus is a widespread, species-rich group that has never been revised throughout its geographic range. Herein I review 103 species of Triepeolus, including all but those that belong to the newly defined Triepeolus verbesinae and Triepeolus simplex species groups (these will be the topic of a subsequent paper). I present three keys to the species, which together cover the genus throughout its range in the New World; the keys are to the females of North and Central America, the males and females from Eastern North America, and the males and females from South America and the Caribbean. I propose the following 37 new species: Triepeolus antiochensis, T. argentimus, T. argus, T. bimorulus, T. charlesi, T. circumculus, T. claytoni, T. cruciformis, T. diffusus, T. dilutus, T. edwardi, T. engeli, T. exilicurvus, T. flavigradus, T. fulgidus, T. griswoldi, T. interruptus, T. isohedrus, T. jennieae, T. joliae, T. lateralis, T. margaretae, T. mauropygus, T. melanarius, T. micheneri, T. nayaritensis, T. parkeri, T. partitus, T. parvidiversipes, T. parvus, T. perpictus, T. phaeopygus, T. punctoclypeus, T. quadratus, T. simulatus, T. vernus, and T. warriti, and propose the new combinations Triepeolus laticeps (Friese), Triepeolus tepanecus (Cresson) and Triepeolus zacatecus (Cresson). I newly synonymize 45 of the 169 previously proposed Triepeolus names, for a total of 51 synonymies: T. alachuensis Mitchell under T. rufithorax Graenicher; Epeolus albopictus Cockerell, E. costaricensis Friese, and E. flavocinctus Friese under T. aztecus (Cresson); T. alpestris Cockerell, T. amandus Cockerell, and T. vandykei Cockerell and Sandhouse under T. paenepectoralis Viereck; E. bardus Cresson, T. mesillae Cockerell, and T. pimarum Cockerell under T. distinctus (Cresson); T. brunneus Cockerell under T. balteatus Cockerell; T. charlottensis Mitchell under T. brittaini Cockerell; T. cirsianus Mitchell under T. donatus (Smith); T. concinnus Cockerell under T. townsendi Cockerell; T. coquilletti Cocker- ell, T. helianthi arizonensis Cockerell, T. helianthi pacificus Cockerell, T. lineatulus Cockerell and Sandhouse, and T. maculiventris Cockerell under T. helianthi (Robertson); T. dichropus Cockerell, T. eldredi Cockerell, T. helianthi grandior Cockerell, T. pallidiventris Cockerell and Sandhouse, T. rectangularis Cockerell, and T. wyomingensis Cockerell under T. texanus (Cresson); T. digueti Cockerell and E. nobilis Friese under T. intrepidus (Smith); T. floridanus Mitchell

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 5 under T. georgicus Mitchell; T. fortis Cockerell, T. insolitus Cockerell, and T. trilobatus Cockerell under T. martini (Cockerell); T. foxii Cockerell under T. rufoclypeus (Fox); T. lestes Cockerell under T. subalpinus Cockerell; T. loganensis Cockerell and T. sandhousae Cockerell under T. fraserae Cockerell; T. nautlanus Cockerell under T. lunatus (Say); E. nigriceps Smith under T. robustus (Cresson); E. oswegoensis Mitchell under T. pectoralis (Robertson); T. perelegans Cockerell and T. trichopygus Cockerell and Timberlake under T. penicilliferus (Brues); T. signatus Hedicke under T. ventralis (Meade-Waldo); T. stricklandi Cockerell under T. subalpinus Cockerell; E. superbus Provancher and E. texanus nigripes Cockerell under T. remigatus (Fabricius); and E. utahensis Cockerell under T. heterurus (Cockerell and Sand- house).

Key words: Bee, Apoidea, Apidae, Nomadinae, Epeolini, brood parasite, cuckoo bee, taxonomic revision

Introduction

The cleptoparasitic bee genus Triepeolus (Apidae: Epeolini) is a relatively widespread, species-rich genus, much like the related genus Epeolus. Triepeolus ranges throughout much of the New World, from British Columbia and Nova Scotia, Canada, south through the Antilles and Central America. In South America, the genus is found from Colombia to Guárico, Venezuela, south to Río Negro, Argentina (but not including Chile), and Santa Catarina, Brazil. An additional two species are found in the Old World, with one species each in Eurasia [from Northern Italy east to the Volga (Privolzhsky) Federal District in Russia and western Kazakhstan] and eastern Asia (from Guangxi, China, north to the Khabarovsky Krai, Russia, and east to Japan) (Michener 2000, and herein). Unlike Epeolus, which only has Colletes as hosts, it has been documented to attack a wide range of host bees in several families (Rightmyer 2004, and references cited therein); further evidence suggests that Triepeolus may, in fact, parasitize members of every family within its geographic range (herein, and Rightmyer in prep.). The purpose of this and a forthcoming companion paper (Rightmyer in prep.) is to provide an overview of the species of Triepeolus worldwide. Herein I treat all Triepeolus species excluding those belonging to the newly defined T. verbesinae and T. simplex species groups (defined below, with lists of included taxa). For each species, I present a taxonomic history, description, geographic distribution, known bee and floral host data, tally of specimens examined, and their repositories. I provide three keys to the species. One is a key to the species in the regions where Triepeolus is most diverse—North and Central America. This key excludes species belonging to the T. verbesinae and the T. simplex species groups, and is restricted to females because the males have not yet been fully studied in these regions. The second and third keys are more comprehensive: they include both males and females of all species (to the extent that both genders are known), and include species of the T. verbesinae and T. simplex species groups. However, they are restricted to regions where Triepeolus is less diverse: the second key is limited to those species found in North America east of the Mississippi River, and the third to those species found in South America and the Caribbean. Thus, all female Triepeolus can be identified to species with the keys included herein except for western North and Central American species of the verbesinae and simplex species groups. Triepeolus is a relatively well-collected group of bees that is commonly confused with the related genus Epeolus. As a step towards gaining a better understanding of Triepeolus, the main focus of this paper is an examination of the females of this genus; however, where known, males are also included in these treatments. Indeed, the examination of male primary types was an important component of this study, and in most cases it was a straightforward matter to associate male type specimens with females. In only three cases, a name associated with an examined male holotype could not be associated with a female. These three unassociated names are T. cuneatus Cockerell, T. isocomae Cockerell, and T. sequior Cockerell (see “Unplaced Species Names,” p. 131). Also included in the list of “Unplaced Species Names” are five species whose primary type specimens or series could not be found (see below), and whose identities could not be determined based on the

6 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER original descriptions: Triepeolus scelestus tubercularis Brues, Triepeolus hopkinsi Cockerell, Triepeolus pomonalis Cockerell, Epeolus mercatus Fabricius, and Doeringiella nemoralis Holmberg. The majority of Triepeolus species were proposed by T.D.A. Cockerell, who also provided a number of keys to limited geographical areas, often including both Epeolus and Triepeolus within the same key [e.g., Cockerell 1898 (New Mexico species); 1904: 34–35 (mostly southwestern USA); 1905a: 313–314 (Smith types); 1907b: 248–249 (Boulder Colorado species); 1907c: 62 (misc. New Mexico species); 1921: 13–15 (Rocky Mountain species); and 1928a: 107–111 (Colorado species)]. Other, similarly limited keys, were provided by Robertson (1903: 284–287, primarily for Illinois species) and Brues (1903: 79–80, for Texas species). Unfortunately, confusion resulted from the failure of many of these authors to distinguish Epeolus from Triepeolus, and to associate males and females within each of the genera. Thus, a number of synonymous names were created for species based on different genders, or due to overlooking a name that was proposed in the incorrect genus. Strides towards correcting these problems were made by Linsley and Michener (1939: 299–301, plates xv–xviii), who redescribed the genus and provided a more accurate understanding of it. Moure (1955: 123–132) provided the first comprehensive treatment for a subset of the species (of South America), followed by Mitchell (1962: 459–485, for the Eastern U.S. species). Robertson (1928) listed a number of floral associations for many Triepeolus species, and Hurd et al. (1980) provided a list of species visiting Helianthus flowers.

Materials and methods

There have been 169 species names proposed for Triepeolus. Many are not clearly recognizable from the original descriptions, and subsequent publications and determinations have to be evaluated carefully. I have seen the primary type specimen or series for each of these names except for fifteen. Those names are: Epeo- lus penicilliferus Brues, E. scelestus tubercularis Brues, T. hopkinsi Cockerell, T. pomonalis Cockerell, E. luctuosus Eversmann, E. mercatus Fabricius, Melecta remigata Fabricius, T. cuabitensis Genaro, T. roni Genaro, T. signatus Hedicke, Doeringiella nemoralis Holmberg, T. atoconganus Moure, E. lunatus Say, E. quadrifasciatus Say, and E. osiriformis Schrottky. However, I had access to several high-quality photographs of the E. osiriformis Schrottky holotype, and was able to borrow a paralectotype specimen of T. atoconganus Moure and paratype specimens of the two Genaro species, T. cuabitensis and T. roni. Further comments regarding these species names can be found under the individual entries in the Triepeolus Descriptions section, below. Several Triepeolus species names are based on lectotype specimens. Cockerell frequently designated “type” and “cotype” specimens when describing new species; however it was not necessary to designate lectotype specimens for his species because he intended specimens labeled “type” to be considered holotypes, and those labeled “cotype” to be considered paratypes (C. D. Michener, personal communication, 2006). In the Taxonomy section, unless otherwise noted, all listed distributional, plant, and seasonal data are from labels on specimens that I personally examined. Host data were gathered from the labels of personally examined specimens as well as from published literature and communications with other researchers; where these host associations exist, the sources of the data are indicated under each species heading. Due to the general historical confusion surrounding the identity of many of the species of Triepeolus, published host data must generally be approached with some skepticism; however, many convincing associations have been established. I have not included published plant associations herein; they are both considerably more numerous than those of the host bees, and potentially less species-specific for cleptoparasitic bees such as Triepeolus than for other bees. In the following keys and Taxonomy section, measurements of total body size are given as approximate values, reflecting the fact that the relative positions of the head, mesosoma, and metasoma, as well as the extension of the different segments of the metasoma, were frequently variable. Rather than attempt to take

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 7 into account these variables by measuring each segment separately, total body size was taken as a single mea- surement, starting at the anterior margin of the head and ending at the posterior margin of the last visible tergum in dorsal view. Also in the following keys and Taxonomy section, emphasis is placed on the appearance of the specialized setae of the female fifth metasomal tergum, which together make up the pseudopygidial area (see below). In order to view these setae properly, it is necessary to use light originating dorso-anteriorly, and for the bee to be rotated such that the dorsal surface of the tergum is at an oblique angle to the viewer. It may be useful to compare the appearance of the pseudopygidial area of a well-known species with its image given at the end of the paper, as a means of standardizing the lighting and specimen orientation for unknown specimens. Special emphasis is also given to the relative length of the clypeus in the following keys; in order to determine this length, it is necessary to view to face such that the dorsal and ventral margins of the compound eyes are equally in focus. In the following keys, characters given in parentheses imply that the same features in the alternate half of the couplet are variable. Specimens used in this study were borrowed from the following individuals and institutions. The city where each institution and/or individual is located is used below to indicate where specimens are preserved. ANN ARBOR—University of Michigan Museum of Zoology (M. F. O’Brien); ANSFELDEN—Maximilian Schwarz, personal collection, Austria; AUSTIN—Central Texas Melittological Institute (J. L. Neff); BERKE- LEY—Essig Museum of Entomology, University of California (C. Barr); BERLIN—Zoologisches Museum, Humboldt-Universität (F. Koch); BOULDER—University of Colorado Museum (V. Scott); BRUSSELS—Insti- tut royal des Sciences naturelles de Belgique (J. Constant); CHAMELA—Estación de Biología, Universidad Nacional Autónoma de México, Jalisco (R. Ayala); CORVALLIS—Oregon State Arthropod Collection (J. Leathers); CURITIBA—Universidade Federal do Paraná, Brazil (G.A.R. Melo, D. Urban, and F. F. de Oliveira); DAVIS—Bohart Museum of Entomology, California (R. Thorp and L. Kimsey); FUKUOKA— Kyushu University, Japan (O. Tadauchi); GAINESVILLE—Florida State Collection of Arthropods (J. R. Wiley); HEREDIA—Instituto Nacional de Biodiversidad, Costa Rica (A. Lépiz); IOWA CITY—Chris Gienapp, personal collection, Iowa; ITHACA—Cornell University Insect Collection, New York (E. R. Hoebeke and B. N. Danforth); LAWRENCE—University of Kansas Natural History Museum and Biodiversity Research Center (C. D. Michener, M. S. Engel, and Z. Falin); LAWRENCE-BAKER—Donald and Madge Baker Collection, at the University of Kansas Natural History Museum and Biodiversity Research Center; LIMA—Museo Nacio- nal de Historia Natural de la Universidad Nacional Mayor de San Marcos, Peru (C. Rasmussen and G. Lamas); LOGAN—USDA-ARS Bee Biology and Systematics Laboratory, Utah (T. L. Griswold); LONDON— The Natural History Museum, U. K. (G. Else and C. Taylor); LOS ANGELES—Los Angeles County Museum of Natural History, California (R. Snelling); MILWAUKEE—Milwaukee Public Museum, Wisconsin (G. R. Noonan); NEW YORK—American Museum of Natural History (J. G. Rozen, Jr., J. S. Ascher, and V. Giles); NEW YORK-ASCHER—J. S. Ascher personal collection; OTTAWA—Canadian National Collection of Insects, Ontario (L. Dumouchel); PHILADELPHIA—Academy of Natural Sciences, Pennsylvania (J. Weintraub); PUE- BLA—Universidad de las Américas, México (C. Vergara); QUéBEC—Université Laval, Collection Pro- vancher (J.-M. Perron); RALEIGH—North Carolina State University Insect Collection (R. Blinn); RIVERSIDE—University of California Entomology Research Museum (D. Yanega); SAN DIEGO—San Diego Natural History Museum, California (D. Faulkner); SAN FRANCISCO—California Academy of Sciences (N. D. Penny, W. J. Pulawski, and R. Zuparko); STARKVILLE—Mississippi Entomological Museum (T. L. Schiefer); TEMPE—Arizona State University (M. E. Douglas); TERRE HAUTE—Robert P. Jean, personal collection, Indiana; TORONTO—Royal Ontario Museum (T. Romankova); TUCSON—University of Arizona (C. A. Olson); UNIVERSITY PARK—Frost Entomological Museum, Pennsylvania State University (R. A. Byers); URBANA—Illinois Natural History Survey, Illinois (C. Favret); WEST LAFAYETTE—Purdue Entomological Research Collections, Indiana—A. Provoncha; WASHINGTON D.C.—United States National Museum of Nat-

8 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER ural History (D. Furth, B. P. Harris, and M. Melo); ZILLAH—Eugene R. Miliczky, personal collection, Wash- ington. Specimens were examined, illustrated, and measured using an Olympus SZX9 dissection microscope, drawing tube, and ocular micrometer, ruler, or calipers. Photomicrographs were taken using MicrOptics and EntoVision Digital Imaging Systems.

Morphology

The morphological terminology used herein follows that proposed by Michener (1944; 2000), except for certain terminology proposed by Engel (2001) for wing veins. The following abbreviations are used in the text: flagellar segment (F), ocellar diameter (OD, measured from the median ocellus), intertegular width (ITW), metasomal tergum (T), metasomal sternum (S), and, for features on the terga of the metasoma, basal transverse band of pale setae (BTB), apical transverse band of pale setae (ATB), and lateral, longitudinal band of pale setae (LLB) (see Fig. 1a). Special descriptive terms and phrases used in this paper are as follows: The clypeal midline is a glabrous, sometimes elevated, longitudinal line at approximately the midpoint of the clypeus (Figs. 244–246). The midline is often more noticeable basally on the clypeus. The larger punctures of the clypeus are punctures that are distinctly larger than the other, generally more numerous, punctures found on the clypeus (Fig. 246). Thus, the term “larger punctures” is a relative one, indicating that two different sizes of punctures are present on the clypeus. These larger punctures are not found in all Triepeolus species, as sometimes only one size of puncture (generally rather small) is found. The paramedian bands are paired, longitudinal bands of pale setae on either side of the middle of the mesoscutum. The intertegular width is the distance between the inner margins of the tegulae, measured at the middle. The intercoxal area is a diamond-shaped region of integument found between the middle coxae, in ventral view. The midpoint of the scutellum is the approximate longitudinal middle of the scutellum, when viewed dorsally (not taking into account the posterior surface of the scutellum, which is typically not fully visible in dorsal view). The T1 discal patch is the region of dark brown or black setae found medially on that tergum, and which is typically partially or fully margined by bands of yellow or white setae (Fig. 1a). When referring to the bands of yellow or white (often more simply termed “pale”) setae on the metasoma, the transverse bands are those extending latitudinally across the tergum, while the longitudinal bands are those found laterally on the tergum, often connecting the transverse bands found apically and basally on the tergum (Fig. 1a). In dorsal view, on each lateral half of T2, the longitudinal band and apical transverse band of pale setae usually meet at the apicolateral margin, forming an angle. This angle is most commonly acute, although it can be nearly 90 degrees, or rarely even obtuse. In general, in the descriptions and keys below, a weakly acute angle is approximately between 65 and 80 degrees, while a strongly acute angle is approximately 65 degrees or less. The pseudopygidial area is the medioapical region of specialized setae on the female T5 (Fig. 1a). When describing the pseudopygidial area, the base or basal region is that region or margin of the pseudopygidial area basal on the tergum. In many species of Triepeolus, this basal region is differentiated by silvery setae. Sometimes the pseudopygidial area is distinctly triangular in shape (e.g., Figs. 2b, 24, 50, 97, 117); when this is the case, the base of the pseudopygidial area is the pointed end of the triangle (e.g., Fig. 117), and is not to be confused with what is sometimes considered the base, or flat edge, of the triangle. A pseudopygidial area that is described herein as semicircular resembles half of a circle (e.g., Figs. 32, 77), while an ovate pseudopygidial area is one that is round, but not perfectly circular (e.g., Figs. 10, 14, 81, 83, 130; for an example of a perfectly circular pseudopygidial area, see Fig. 2a). The most common pseudopygidial area shapes are quadrate (e.g., Figs. 6, 12, 17) and semicircular (e.g., Figs. 19, 32, 77), but these shapes can intergrade (e.g. Figs. 4, 54), and many other, more specialized forms also exist (e.g. Figs 8, 35, 71, 142).

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 9 FIGURES 1a and 1b. Triepeolus spp. 1a. Female metasoma, dorsal view. 1b. Male T7, dorsal view. See Morphology text for explanations of labeled features.

The T5 usually has a wide region of pale setae laterally on either side of the pseudopygidial area; this region of pale setae is termed the pale lateral setae of T5 (Fig. 1a). The transverse basal ridge of the male pygidial plate is a carina, sometimes rather weak and/or broken by punctures, which is found in some Triepeolus species, and which demarcates the transition between the apical downturned plate and the basal region of T5 (Fig. 1b). Occasionally, it is possible to differentiate an apical plate that is distinctly downturned from the base of T5, without the presence of a transverse basal ridge.

FIGURES 2a and 2b. Female pseudopygidial areas. 2a. Triepeolus verbesinae, from Arizona. 2b. Triepeolus simplex, from Kansas. Fig. 2c. Triepeolus kathrynae, from New Mexico, showing apical bristle-like setae on female S5, typical of the Triepeolus simplex species group.

1 Upper face with erect setae such that clypeus appears recessed (Fig. 190) [clypeus shining, with regularly-spaced small punctures and dense, minute, weakly impressed punctures; mesepisternum with erect, simple setae (Fig. 166); S5 strongly downcurved, Fig. 36]1...... 2 - Upper face with setae variable, but not such that clypeus appears recessed (e.g., Figs. 57, 58, 112, 143, 180, 182) ...... 3 2(1) Metasomal terga with bands of setae grading from pale yellow to grayish white posteriorly; T2 with LLB absent or forming very weakly acute angle with ATB ...... T. flavigradus Rightmyer, n. sp. (Fig. 74) - Metasomal terga with bands of setae the same shade of pale yellow throughout; T2 with LLB forming 90 degree angle with ATB ...... T. robustus (Cresson) (Fig. 188) 3(1) Pseudopygidial area with long, erect setae forming rounded, three-dimensional posterior structure (i.e., similar to that of T. concavus, Figs. 35, 36) (S5 strongly downcurved)...... 139 - Pseudopygidial area not similar to that of T. concavus (e.g., all pseudopygidial areas figured except Figs. 35, 36, 137, 169, 262a, 263a) ...... 4 4(3) Intercoxal area of mesosoma posteriorly forming clasper-like structure (i.e., with each lateral half apically incurved and laterally swollen, almost caliper-like in shape; Fig. 20) (preoccipital carina very strong, outcurved on gena; mesepisternum with erect, simple setae) ...... T. balteatus Cockerell (Fig. 18) - Intercoxal area mostly flat, sometimes slightly reflexed towards body at apex ...... 5 5(4) Pseudopygidial area with subapical transverse band of silvery, fine, appressed setae and apical band of long, stout, suberect setae (as in Fig. 71) ...... T. epeolurus Rightmyer (Fig. 70) - Pseudopygidial area variable, but lacking both transverse subapical band of silver setae and apical band of specialized setae as in Fig. 71 ...... 6 6(5) Pseudopygidial area with setae long and parted medially, apical margin concave (as in Fig. 89)...... T. heterurus (Cockerell & Sandhouse) (Figs. 88, 90) - Pseudopygidial area variable, but lacking long, medially-parted setae like those in Fig. 89...... 7 7(6) Preoccipital carina absent, sometimes with distinctly angled corner on posterior margin of gena, but not forming true, protruding carina; head in dorsal view globular, nearly as long as wide, and shining beneath erect setae (e.g., Fig. 45) (pseudopygidial area as in Figs. 46 and 133, with apical margin concave or straight, and with transverse apical band of sparse, stout setae; mesepisternum with dense, erect setae; mesoscutum with paramedian band of setae absent or poorly differentiated from erect setae present anteriorly; dorsum of mesosoma and metasoma black with bands of white setae) . . . . 8 - Preoccipital carina present at least on gena; head shape usually more conspicuously wider than long (e.g., Fig. 49) ...... 9 8(7) T1 with BTB present (Fig. 258); T2 usually with LLB present (Southwestern U.S.) ...... T. mojavensis Linsley (Fig. 132) - T1 lacking BTB (Fig. 249); T2 lacking LLB (Midwestern U.S. and Canada) ...... T. dacotensis (Stevens) (Fig. 45) 9(7) Scutellum with pale yellow setae encircling black or reddish brown region of setae on each biconvexity, axillar spine bordered on all sides with pale yellow setae (Fig. 205) (mesepisternum with erect, simple setae; pseudopygidial area with apical margin weakly convex or straight) ...... 10 - Scutellum lacking distinct regions of dark setae encircled by pale setae on biconvexities, axillar spine rarely bordered on all sides with pale setae ...... 11

1. Characters given in parentheses imply that the same features in the alternate half of the couplet are variable.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 11 10(9) Clypeus and mesepisternum shining, with setae sparse; mesepisternum with punctures separated by up to one to five puncture diameters; pseudopygidial area with basal crescent of silvery setae poorly differentiated from rest of setae ...... T. bimorulus Rightmyer, n. sp. (Fig. 21) - Clypeus and mesepisternum mostly covered with pale yellow, plumose setae; mesepisternum with punctures nearly contiguous to separated by up to one puncture diameter; pseudopygidial area with basal crescent well differentiated from rest of setae (California) ...... T. simulatus Rightmyer, n. sp. (Fig. 203) 11(9) T1 with ATB absent or greatly restricted laterally (mesepisternum lacking erect, simple setae). . 12 - T1 with ATB present, not especially restricted laterally...... 14 12(11) Pseudopygidial area with apical margin concave (e.g., Fig. 2b), laterally with stout, spine-like setae; mesoscutum lacking paramedian band of setae ...... T. mexicanus (Cresson)2 - Pseudopygidial area with apical margin weakly or strongly convex, lacking spine-like setae laterally; mesoscutum with paramedian band of setae...... 13 13(12) Pseudopygidial area circular, with apical margin strongly convex, and with suberect setae on disc and dense, stouter, slightly flattened setae along most of apical margin (e.g., Fig. 2a); paramedian band of setae tapering towards anterior margin of mesoscutum ...... T. bilineatus Cockerell3 - Pseudopygidial area subquadrate, covered with dense, fine, appressed, golden setae, with apical margin weakly convex (Fig. 28); paramedian band of setae ending abruptly, not reaching anterior margin of mesoscutum ...... T. cameroni (Meade-Waldo) (Fig. 27) 14(11) Pseudopygidial area rectangular, with setae uniformly very dense and short, apical margin straight, with thin region of silvery setae on apical margin (Fig. 56); T1 mostly covered with appressed yellow setae except for small, diamond, ovate, or rectangular patch of black setae mediobasally (axillar spine exceeding posterior margin of scutellum, incurved, often red) ...... T. distinctus (Cresson) (Fig. 55) - Pseudopygidial area variable but not so distinctly rectangular and with setae not so uniformly dense and short, shape of apical margin and amount of silvery setae variable; T1 with black discal patch variable, less commonly reduced to small mediobasal patch ...... 15 15(14) Scutellum red (rarely partially black), strongly contrasting with black mesoscutum and axilla (axilla small and triangular, lacking free apical spine; upper face, mesoscutum, and upper mesepisternum covered with dense, erect, golden setae; mesoscutum lacking distinct paramedian band of setae; pseudopygidial area poorly differentiated from rest of T5) . . . T. intrepidus (Smith) (Fig. 94) - Scutellum black or rarely red (if red then axilla also red)...... 16 16(15) Pseudopygidial area strongly triangular, distinctly longer than wide, composed of silvery setae except for small medial patch of darker setae (Fig. 50); dorsum of body entirely black (some specimens from Utah) or with banding of appressed, pale yellow setae (if with pale banding, then T2 with LLB reduced or forming 90 degree angle with ATB; mesepisternum with relatively short, sparse, erect, simple setae) (North Dakota and Montana south to Arizona, New Mexico, and Chi- huahua) ...... T. denverensis Cockerell (Figs. 47–49) - Pseudopygidial area variable in shape, if composed of silvery setae except for small medial patch of darker setae, then pseudopygidial area about as long as wide; dorsum of body with at least some bands of pale yellow setae ...... 17 17(16) Pseudopygidial area distinctly circular, with apical margin strongly convex, and with setae relatively sparse on disc, dense and stout on apical margin (Fig. 2a); S5 strongly downcurved (scutellum usually with transverse ridge on posterior surface; clypeus usually lacking midline) ......

2. Species belonging to the Triepeolus simplex species group; not treated further herein. 3. Species belonging to the Triepeolus verbesinae species group; not treated further herein.

12 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER ...... T. verbesinae species group - Pseudopygidial area variable but not distinctly circular, usually subquadrate (e.g., Figs. 6, 12, 17) or semicircular (e.g., Figs. 19, 32, 77) or an intergrade of the two, or less commonly triangular (e.g., Figs. 2b, 24, 50, 97), ovate (e.g., Figs. 10, 14, 81, 83, 130), or rarely another unique form, (if rounded then lacking combination of sparse setae on disc and dense setae at apical margin); S5 rarely strongly downcurved...... 18 18(17) Pseudopygidial area triangular, sometimes with stout, spine-like setae laterally, apical margin concave or straight (e.g., as in Fig. 2b); S5 straight in profile, apical margin with dense bristle-like setae (Fig. 2c) (clypeus often lacking midline) ...... T. simplex species group - Pseudopygidial area variable, but with apical margin straight or convex, and never with spine-like setae laterally; S5 shape in profile variable but lacking dense bristle-like setae on apical margin. 19 19(18) T5 with pale lateral setae absent (e.g., Fig. 140) or restricted to apical margin of tergum lateral to pseudopygidial area (e.g., Fig. 202) ...... 20 - T5 with pale setae bordering or adjacent to at least most of lateral margin of pseudopygidial area, these setae sometimes sparse (e.g., Fig. 207) ...... 27 20(19) Mesepisternum with long, erect, simple setae (e.g., Fig. 166) ...... 21 - Mesepisternum lacking long, erect, simple setae ...... 22 21(20) T1 lacking BTB...... T. texanus (Cresson), in part - T1 with BTB present...... T. edwardi Rightmyer, n. sp. (Figs. 63, 64) 22(20) Pseudopygidial area long and narrow (with apical semicircular region of coarser setae; Fig. 213) ...... T. tanneri Cockerell (Fig. 212) - Pseudopygidial area as wide as long, or wider ...... 23 23(22) Pseudopygidial area strongly triangular (Fig. 117) (paramedian band of pale setae joined laterally to pale yellow setae on anterior margin of mesoscutum) ...... T. martini (Cockerell) (Fig. 116) - Pseudopygidial area ovate to subrectangular ...... 24 24(23) Mesoscutum with dense yellow setae on entire anterior margin (Fig. 139); T2 with LLB forming 90 degree angle with ATB ...... T. nevadensis (Cresson) (Fig. 138) - Mesoscutum lacking pale setae on anterior margin; T2 with LLB absent or forming acute angle with ATB ...... 25 25(24) Mesoscutum and clypeus shining; pseudopygidial area relatively wide, forming band on apical margin of T5 (Fig. 211) (clypeus with distinct larger punctures as well as more numerous smaller punctures) ...... T. subnitens Cockerell & Timberlake (Fig. 210) - Mesoscutum and clypeus relatively matte; pseudopygidial area longitudinally ovate to subquadrate, not forming band on apical margin of T5 ...... 26 26(25) T2 with ATB medially interrupted; metasomal sterna lacking bands of pale yellow setae ...... T. scelestus (Cresson) (Fig. 201) - T2 with ATB continuous across disc of tergum; metasoma with apical transverse bands of pale setae laterally on at least S3 ...... T. helianthi (Robertson), in part [Fig. 86 (showing more common variety with pale setae on T5)] 27(19) Pseudopygidial area with setae uniform or nearly uniform in reflectance, texture, and density (i.e., when slightly rotating the specimen to different angles, strongly differentiated regions of different types of setae on the pseudopygidial area should not be detectable; e.g., Figs. 83, 87, 179, 184, 207, 209) ...... 28 - Pseudopygidial area not uniform in reflectance, texture, and/or density (e.g., Figs. 4, 6, 8, 10) . . 58 28(27) Mesoscutum with abundant pale yellow setae on anterior margin, joined laterally to paramedian band of setae (Fig. 186) (robust, body length greater than 10 mm and ITW greater than 2.3 mm) ...... T. remigatus (Fabricius) (Fig. 185)

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 13 - Mesoscutum lacking abundant pale yellow setae on anterior margin (if with sparse pale setae on anterior margin, then body length less than 10 mm and ITW less than 2 mm) ...... 29 29(28) Mesepisternum with erect, simple setae (Fig. 166; sometimes less than 0.25 OD in length, sparse, and difficult to see; if so, then these species additionally key from couplet 36)...... 30 - Mesepisternum lacking erect, simple setae...... 36 30(29) Pseudopygidial area triangular, with silvery reflecting setae ...... 31 - Pseudopygidial area rounded, with dark (or rarely pale) golden reflecting setae...... 32 31(30) Mesoscutum with paramedian band of setae ...... T. subalpinus Cockerell (Fig. 206) - Mesoscutum lacking paramedian band of setae ...... T. brittaini Cockerell (Fig. 23) 32(30) Mesepisternum with erect setae as long as 0.75 to 1 OD (axillar spine at least reaching, but usually surpassing, midpoint of scutellum; metasomal terga with relatively robust bands of pale setae) . 33 - Mesepisternum with erect setae only ca. 0.25 median ocellus diameter or less in length ...... 34 33(32) Metasomal terga with bands of pale yellow setae; legs red (rarely black); axillar spine triangular, reaching midpoint of scutellum, or less commonly sharply pointed and apically incurved, well-surpassing midpoint of scutellum (Southern México to Southwestern U.S.) ...... T. grandis (Friese) (Fig. 82) - Metasomal terga with bands of orange-yellow to yellow setae; legs black; axillar spine long, pointed, apically incurved (Southern México to Costa Rica) . . . .T. zacatecus (Cresson) (Fig. 239) 34(32) T1 discal patch widely rectangular with sinuate BTB and ATB; body length greater than 15 mm (integument entirely black except on mandible and F1; axillar spine pointed, reaching posterior margin of scutellum) (Southeastern U.S.) ...... T. monardae Mitchell (Fig. 134) - T1 discal patch transversely ovate or triangular; body length less than 14 mm (Southwestern U.S., México, and Central America) ...... 35 35(34) Legs red; T1 discal patch transversely ovate, BTB and ATB parallel; pseudopygidial area with pale golden brown setae ...... T. fulgidus Rightmyer, n. sp. (Fig. 78) - Legs black, or if partly red then T1 discal patch triangular; T1 discal patch triangular or transversely ovate; pseudopygidial area with setae dark brown in certain light, or reflecting dark golden ...... T. tepanecus group (Table 2) 36(29) Pseudopygidial area rounded apically, elongated basally into tear shape when viewed from oblique angle; S5 strongly downcurved; T1 discal patch very widely subovate to transversely diamond- shaped (Arizona, New Mexico, and Durango) ...... T. loomisorum Rozen (Fig. 108) - Pseudopygidial area variable but not elongated basally into tear shape; S5 straight or moderately downcurved in profile; T1 discal patch rarely transversely diamond-shaped ...... 37 37(36) T1 lacking BTB and LLB, or rarely with BTB reduced to small lateral patch; T2 lacking LLB (mesoscutum with paramedian band of setae absent or rarely reduced; integument of face entirely dark brown, or with limited orange-red regions on F1 and F2) (central México) ...... T. interruptus Rightmyer, n. sp. (Fig. 92) - T1 with BTB present, if reduced then LLB strongly present; T2 with LLB usually present, if absent, then T1 with BTB present...... 38 38(37) T1 with BTB and ATB of the same length, linear, and parallel to each other for most of width, thus T1 discal patch rectangular or transversely ovate (e.g., Figs. 86 and 234) ...... 39 - T1 with BTB and ATB not linear, not parallel to each other for most of width, or if parallel, then anterior margin of BTB and posterior margin of ATB not of the same length; thus T1 discal patch appearing ovate, trapezoidal, or triangular ...... 52 39(38) T1 lacking LLB...... 40 - T1 with LLB present ...... 41 40(39) Integument with red regions on antenna, labrum, pronotal lobe, tegula and legs; pseudopygidial

14 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER area widely semicircular, poorly differentiated basally from rest of T5 (Fig. 99) ...... T. jennieae Rightmyer, n. sp. (Fig. 98) - Integument entirely dark brown to black (excluding mandibles); pseudopygidial area ovate, with basal boundary well defined by change in setal morphology (Fig. 103) ...... T. lateralis Rightmyer, n. sp. (Fig. 102) 41(39) T1 with LLB forming 90 degree angle with ATB, thus T1 discal patch appearing distinctly rectangular (e.g., Fig. 235) ...... 42 - T1 with LLB forming acute angle with, or rounded connection between, BTB and ATB, thus T1 discal patch appearing transversely ovate or widely trapezoidal (e.g., Figs. 201, 210) ...... 46 42(41) Mesoscutum and clypeus shining; body length ca. 11–16 mm ...... 43 - Mesoscutum and clypeus relatively matte; body length ca. 7–11 mm ...... 44 43(42) T2 lacking LLB; axillar spine pointed, surpassing scutellar midpoint (Southeastern U.S.) ...... T. quadrifasciatus atlanticus Mitchell (Fig. 178) - T2 with LLB forming acute angle with ATB; axillar spine triangular, reaching scutellar midpoint (Western and Midwestern U.S., Northern México) T. subnitens Cockerell & Timberlake (Fig. 210) 44(42) Mesepisternum with pale, plumose setae restricted to small patches below scrobal groove and posterior to pronotal lobe; metanotum with pale setae restricted to lateral corner ...... T. lateralis Rightmyer, n. sp. (Fig. 102) - Mesepisternum with pale, plumose setae on much of dorsal half or more; metanotum mostly or entirely covered with pale setae ...... 45 45(44) Pseudopygidial area subrectangular to subtriangular, with setae suberect, and apical margin of integument straight but often appearing slightly concave due to longer setae laterally than medially (Fig. 173); T2 with ATB continuous medially; T1 discal patch not strongly resembling plus-sign ...... T. phaeopygus Rightmyer, n. sp. (Fig. 172) - Pseudopygidial area rounded, with setae mostly directed posteriorly, and apical margin of integument straight to weakly convex (Fig. 236); T2 with ATB interrupted medially (sometimes only barely); T1 discal patch somewhat resembling plus-sign (Fig. 235) ...... T. warriti Rightmyer, n. sp. (Fig. 234) 46(41) Mesoscutum with diffuse pale yellow setae along anterior margin, forming subtle anchor-shape with paramedian bands of setae (T2 with LLB forming 90 degree angle with ATB, or rarely forming acute angle; pseudopygidial area subrectangular to subtriangular, with setae suberect, and apical margin of integument straight, often appearing slightly concave due to longer setae laterally than medially; Fig. 173) ...... T. phaeopygus Rightmyer, n. sp. (Fig. 172) - Mesoscutum lacking diffuse pale yellow setae along anterior margin, with paramedian band of setae isolated from other pale setae...... 47 47(46) Pseudopygidial area forming transverse band on apical margin of T5 (Fig. 211) (clypeus shining, with distinct larger punctures, Fig. 246; mesoscutum shining)...... T. subnitens Cockerell & Timberlake (Fig. 210) - Pseudopygidial area longitudinally ovate (e.g., Figs. 10, 14, 81, 83, 130), or subquadrate (e.g., Figs. 6, 12, 17) to subtriangular (e.g., Figs. 135, 184), not forming transverse band on apical margin of T5 ...... 48 48(47) Body length ca. 15–18 mm; integument entirely black except on mandible and F1; axillar spine pointed, surpassing midpoint of scutellum (Southeastern U.S.) . . T. monardae Mitchell (Fig. 134) - Body length ca. 7–12 mm; integument entirely black or with red regions; axillar spine triangular, reaching only midpoint of scutellum ...... 49 49(48) Metasomal sterna lacking pale setae; pseudopygidial area dark, poorly differentiated from rest of T5 (Fig. 202) (T2 with LLB absent or reduced to small patch such that resulting angle with ATB is

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 15 not discernable) ...... T. scelestus (Cresson) (Fig. 201) - Metasomal sterna with white setae on at least S2 and S3; pseudopygidial area usually easily differentiated from rest of T5...... 50 50(49) Pseudopygidial area with apical margin of integument straight and setae pale golden brown (Fig. 79) (clypeus shining, with distinct larger punctures; S2–S4 with white setae) (Arizona, New Mex- ico) ...... T. fulgidus Rightmyer, n. sp. (Fig. 78) - Pseudopygidial area with apical margin of integument convex, setae dark brown to golden brown ...... 51 51(50) At least T1–T3 with ATBs medially interrupted; S2–S4 with white setae ...... T. mauropygus Rightmyer, n. sp. (Fig. 118) - Only T1 with ATB medially interrupted; only S3–S4 with white setae ...... T. helianthi (Robertson) (Fig. 86) 52(38) T1 in dorsal view with LLB enlarged, as wide as or wider than width of T1 discal patch (e.g., Fig. 16) ...... 53 - T1 with lateral area of pale setae not especially enlarged, narrower than width of T1 discal patch...... 55 53(52) Body length ca. 7–11 mm; T1 discal patch located basally on disk of tergum (Fig. 16) (Costa Rica) ...... T. aztecus (Cresson) (Figs. 15, 16) - Body length ca. 13–16 mm; T1 discal patch reduced to thin, longitudinal line or appearing as ovate or rectangular patch medially on disk of tergum (Texas and Kansas to East Coast of U.S.) . . . . . 54 54(53) Clypeus entirely red; scutellum and axilla often reddish; T1 mostly covered with pale yellow setae, except for median longitudinal line of dark brown setae (sometimes also with small diamond or ovate median area of dark setae) (Texas and Kansas to Louisiana) ...... T. q. quadrifasciatus (Say) (Fig. 183) - Clypeus at most red apically; scutellum and axilla black; T1 discal patch usually ovate to rectangular, sometimes as above (Eastern U.S.) ...... T. quadrifasciatus atlanticus Mitchell (Fig. 178) 55(52) T1 discal patch transversely ovate, with enlarged circular region medially (e.g., Fig. 25) (Western U.S.) ...... 56 - T1 discal patch subtriangular to ovate, if transversely ovate then lacking additional enlarged circular patch of dark setae medially...... 57 56(55) T2 with LLB forming strongly acute angle with ATB; dorsum of mesosoma and metasoma often with bands of bright yellow setae; pseudopygidial area with golden setae (Fig. 26) ...... T. californicus (Cresson) (Fig. 25) - T2 with LLB forming 90 degree angle with ATB; dorsum of mesosoma and metasoma with bands of pale yellow setae; pseudopygidial area with rust colored setae (Fig. 101) ...... T. joliae Rightmyer, n. sp. (Fig. 100) 57(55) Metasomal terga with bands of pale setae medially interrupted and slightly enlarged submedially, causing the metasoma to superficially resemble that of T. verbesinae; paramedian band of setae narrowed anteriorly, reaching anterior margin of mesoscutum; pseudopygidial area appearing slightly emarginate medioapically due to elongate setae laterally (Fig. 209) ...... T. sublunatus Cockerell (Fig. 208) - Metasomal terga with transverse bands of pale setae of similar width medially as laterally, not resembling those of T. verbesinae; paramedian band of setae either absent or if present, then truncate anteriorly and not reaching anterior margin of mesoscutum; pseudopygidial area evenly straight or convex on apical margin ...... T. tepanecus species group (Table 2) 58(27) Pseudopygidial area widely triangular, with suberect, dense, short, setae (Fig. 60)...... 59 - Pseudopygidial area variable, but lacking similarly dense, short, suberect setae as shown in Fig. 60

16 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER ...... 60 59(58) T2 with LLB forming 90 degree or weakly acute angle with ATB; mesepisternum with punctures very small, contiguous; body length ca. 10–13 mm (Western U.S., east to Colorado) ...... T. diversipes Cockerell (Fig. 59) - T2 with LLB forming acute angle with ATB; mesepisternum with small punctures separated by up to 0.5 puncture diameter; body length ca. 7–8 mm (Arizona) ...... T. parvidiversipes Rightmyer, n. sp. (Fig. 159) 60(58) Mesepisternum with erect setae ca. 0.5 OD in length or more (e.g., Fig. 166) ...... 61 - Mesepisternum lacking erect setae or with erect setae less than 0.25 OD in length...... 83 61(60) T2 with LLB forming 90 degree angle with ATB...... 62 - T2 with LLB absent or forming acute angle with ATB ...... 72 62(61) Clypeus extending below lower tangent of compound eyes by ca. 1.5 OD or more when face viewed with both dorsal and ventral margins of compound eyes in focus ...... 63 - Clypeus with apical margin ca. in line with lower tangent of compound eyes, or surpassing it by no more than 1 OD ...... 65 63(62) Apical third of hind wing distinctly darker than basal two-thirds; mesepisternum irregularly, deeply punctate (Fig. 199) (Eastern U.S.) ...... T. rugosus Mitchell (Fig. 198) - Hind wing uniformly transparent or only slightly darker at apical third; mesepisternum with small, relatively shallowly impressed punctures ...... 64 64(63) Dorsum of mesosoma and metasoma with bands of whitish gray setae; clypeus with larger punctures indistinct; mesepisternum shining between punctures, ventral half of lateral surface of mesepisternum with punctures mostly spaced approximately one puncture diameter apart (Eastern and Midwestern U.S.) ...... T. donatus (Smith) (Fig. 61) - Dorsum of mesosoma and metasoma with bands of pale yellow setae; clypeus with larger punctures distinct; mesepisternum not especially shining, with punctures nearly contiguous (sometimes with punctures spaced a puncture diameter apart in some areas) ...... T. texanus (Cresson) (Fig. 217) 65(62) Pseudopygidial area a longitudinally elongate triangle (i.e., clearly longer than greatest width) (with setae uniformly short, appressed, and dense, and with distinct basal and apical regions of silvery setae; Fig. 175) (Utah west to California)...... T. punctoclypeus Rightmyer, n. sp. (Fig. 174) - Pseudopygidial area variable, but if triangular then not so longitudinally elongate (i.e., at most as long as greatest width and often shorter) ...... 66 66(65) Pseudopygidial area triangular, mostly composed of shining golden setae but with area of shining silvery setae on basal third to fourth (golden and silvery setae similarly long and dense, Fig. 192) (Colorado) ...... T. rohweri Cockerell (Fig. 191) - Pseudopygidial area ovate to subtriangular, with basal crescent of setae distinctly denser and shinier than setae of rest of area, or with pseudopygidial area mostly dark, with vague basal silvery shining crescent ...... 67 67(66) Ventral half of lateral surface of mesepisternum shining, with punctures spaced well over three puncture diameters apart (mesepisternum with erect, simple setae especially long, Figs. 165 and 166) (clypeus with strong midline; T2 with LLB usually forming weakly acute angle with ATB) ...... T. pectoralis (Robertson) (Fig. 163) - Ventral half of lateral surface of mesepisternum variable, but with punctures separated by no more than two puncture diameters ...... 68 68(67) Axilla slightly longer than wide, with distinct apical spine curved medially, reaching or surpassing scutellar midpoint; pseudopygidial area with well defined region of silvery setae on basal third to half of area (paramedian band of setae clearly separated from other regions of pale setae on mesoscutum) ...... 69

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 17 - Axilla as long as wide or shorter, apical spine lacking or very small, reaching at most scutellar midpoint; pseudopygidial area with vague or distinct basal region of silvery setae confined to basal fourth or less of area ...... 70 69(68) T1–T4 with ATB medially interrupted (California: San Luis Obispo)...... T. quadratus Rightmyer, n. sp. (Fig. 176) - At least T3–T4 with ATB continuous medially (California: Antioch) ...... T. antiochensis Rightmyer, n. sp. (Fig. 5) 70(68) Mesepisternum entirely, though somewhat sparsely, covered with pale yellow, appressed, plumose setae; preoccipital carina strong, outcurved on entire gena (e.g., Fig. 144, but on gena only, not also on posterior margin of head behind vertex, as figured); T1 discal patch forming rectangular region of dark setae nearly reaching lateral margin of tergum (as seen in dorsal view) (mesepisternum with integument beneath plumose setae rugose and shining; paramedian band joined laterally to pale yellow setae on anterior third to fourth of mesoscutum; vertex and mesoscutum with dense, golden, erect setae) (Utah, Texas) ...... T. griswoldi Rightmyer, n. sp. (Fig. 84) - Mesepisternum with pale yellow, appressed, plumose setae restricted dorsally, or if covering entire mesepisternum, then these setae very dense such that integument beneath poorly visible; preoccipital carina strong or weak on gena, but rarely strongly outcurved; T1 discal patch reduced or forming rectangular region, but rarely so wide as to nearly reach lateral margin of tergum (as seen in dorsal view) ...... 71 71(70) Pseudopygidial area dark brown, with vague basal crescent of shining setae (Fig. 65); T5 with patch of white setae lateral to pseudopygidial area usually reduced or absent (California, Oregon) ...... T. edwardi Rightmyer, n. sp. (Figs. 63, 64) - Pseudopygidial area with distinct basal crescent of silvery shining setae; T5 with large patch of white setae lateral to pseudopygidial area...... T. paenepectoralis species group (Table 1) 72(61) Pseudopygidial area well-defined, longitudinally ovate, with region of golden setae at apical margin (Fig. 142)...... T. nigrihirtus Mitchell (Fig. 141) - Pseudopygidial area variable, but if longitudinally ovate then not so well defined on basal margin and lacking such highly differentiated region of golden setae at apical margin ...... 73 73(72) Pseudopygidial area longitudinally ovate, entirely covered with dense, shining setae, apically with depressed, circular region (Fig. 8); S5 strongly downcurved (clypeus lacking midline) (California) ...... T. argentimus Rightmyer, n. sp. (Fig. 7) - Pseudopygidial area variable but lacking apical, depressed, circular region; S5 straight or only weakly downcurved in profile...... 74 74(73) Ventral half of lateral surface of mesepisternum shining, with punctures spaced well over three puncture diameters apart (Fig. 165) (clypeus with strong midline; mesepisternum with very long erect, simple setae, Fig. 166)...... T. pectoralis (Robertson) (Fig. 163) - Ventral half of lateral surface of mesepisternum variable, but with punctures separated not more than two puncture diameters apart ...... 75 75(74) Clypeus extending below lower tangent of compound eyes by ca. 1.5 OD or more when face viewed with both dorsal and ventral margins of compound eyes in focus ...... 76 - Clypeus with apical margin ca. in line with lower tangent of compound eyes, or surpassing it by no more than 1 OD ...... 77 76(75) T1 lacking BTB; mesepisternum with integument relatively smooth, with fine punctures (Washing- ton) ...... T. texanus (Cresson), in part - T1 with BTB present; mesepisternum with integument rugose, with relatively deeply impressed, larger punctures (Fig. 199) (Eastern U.S.) ...... T. rugosus Mitchell (Fig. 198) 77(75) Pseudopygidial area relatively poorly differentiated from rest of T5 except for silvery shining setae

18 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER at apical margin, resembling that of Epeolus (Fig. 123) (mesepisternum with erect setae mostly restricted to anterior margin; clypeus lacking midline) (California)...... T. melanarius Rightmyer, n. sp. (Fig. 122) - Pseudopygidial area relatively well-defined, especially basally on T5, lacking apical band of silvery setae resembling that of Epeolus ...... 78 78(77) Pseudopygidial area longitudinally ovate and usually darkly shining, with a vague basal crescent of denser setae (Fig. 83); T1 discal patch triangular to trapezoidal; body length ca. 11–15 mm; T1–T2 often with ATBs medially interrupted and slightly broadened submedially (Southern and Midwest- ern U.S., México) ...... T. grandis (Friese) (Fig. 82) - Pseudopygidial area rarely both longitudinally ovate and nearly uniformly composed of darkly shining setae; T1 discal patch variable but rarely triangular; body usually shorter than 13 mm; T1– T2 with ATBs nearly the same width medially as laterally ...... 79 79(78) Clypeus flat, shining, black, usually strongly contrasting with red antennal scape and labrum (pseudopygidial area triangular, with distinct, narrow basal crescent of shining setae, Fig. 158; T1 with BTB and ATB parallel, surrounding wide ovate discal patch) (Northern México, Southwestern U.S.) ...... T. partitus Rightmyer, n. sp. (Fig. 157) - Clypeus at least weakly convex in profile, usually not strongly contrasting with scape and labrum in color...... 80 80(79) Venter of body entirely covered with dark brown or black setae; T1–T2 with ATBs uninterrupted medially...... T. edwardi Rightmyer, n. sp. (Figs. 63, 64) - Venter of body with at least some pale yellow or white setae; T1–T2 with ATBs usually interrupted medially (rarely entire) ...... 81 81(80) T1 with ATB medially interrupted by less than 1 OD (pseudopygidial area longitudinally ovate, with distinct basal shining crescent, Fig. 77) (North Dakota and Montana south to California and New Mexico)...... T. fraserae Cockerell (Fig. 76) - T1 with ATB medially interrupted by more than 1 OD ...... 82 82(81) Mesepisternum with both simple, erect setae and minutely-branched, erect setae, especially dorsally; antenna usually with at least F1 orange; pseudopygidial area relatively long, subquadrate in outline (Fig. 4)...... T. antiguensis Cockerell (Fig. 3) - Mesepisternum with only simple, erect setae (with minutely-branched setae appressed or weakly suberect); antenna entirely dark brown; pseudopygidial area relatively wide, semicircular in outline (Fig. 121) (México) ...... T. medusa Cockerell (Fig. 120) 83(60) Pseudopygidial area strongly triangular, with straight apical margin weakly shining silvery to golden, contrasting with dark brown setae on rest of area (Fig. 117) (paramedian band joined laterally to pale yellow setae on anterior margin of mesoscutum, forming anchor-shape; T2 with LLB forming acute angle with ATB) (Midwest and Southwest U.S., Northern México) ...... T. martini (Cockerell) (Fig. 116) - Pseudopygidial area variable, but not so perfectly triangular and nearly uniformly dark brown. . 84 84(83) At least lower two-thirds of mesepisternum intensely shining, black, almost entirely glabrous (T1 with ATB medially interrupted by at least 1 OD; axillar spine black) (México) ...... 85 - Ventral half of lateral surface of mesepisternum not so distinctly shining, more conspicuously covered with setae ...... 86 85(84) T2 with LLB forming acute angle with ATB; mesepisternum with punctures usually separated by as much as 4–5 puncture diameters (only 2–3 puncture diameters in some specimens), with white, appressed, plumose setae covering much of upper third, excluding hypoepimeron (Fig. 126); axillar spine pointed, slightly incurved apically, surpassing scutellar midpoint ...... T. micheneri Rightmyer, n. sp. (Fig. 124)

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 19 - T2 lacking LLB; mesepisternum with punctures usually separated by up to two puncture diameters, with white, appressed, plumose setae usually limited to just below scrobal groove (Fig. 216); axilla usually triangular, reaching scutellar midpoint (axillar spine sometimes surpassing scutellar midpoint) ...... T. tepanecus (Cresson) (Fig. 214) 86(84) Scutellum entirely or partially red...... 87 - Scutellum black...... 88 87(86) Paramedian band absent or short, not reaching anterior margin of mesoscutum; mesoscutum usually red but sometimes partially or entirely black; body length ca. 10–14 mm (Southeastern U.S.) ...... T. rufithorax Graenicher (Figs. 193, 194) - Paramedian band reaching anterior margin of mesoscutum, often joined to pale yellow setae on anterior margin of mesoscutum; integument of mesoscutum usually black but sometimes with red areas; body length ca. 7–9 mm (Colorado west to California and Arizona)...... T. perpictus Rightmyer, n. sp. (Fig. 170) 88(85) Clypeus extending below lower tangent of compound eyes by ca. 1.5 OD when face viewed with both dorsal and ventral margins of compound eyes in focus ...... 89 - Clypeus with apical margin ca. in line with lower tangent of compound eyes, or surpassing it by no more than 1 OD ...... 91 89(88) Dorsum of mesosoma and metasoma with bands of very pale gray to white setae (mesepisternum usually with short, erect setae) ...... T. donatus (Smith) (Fig. 61) - Dorsum of mesosoma and metasoma with bands of pale yellow setae ...... 90 90(89) Paramedian band joined laterally to dense band of pale yellow setae on anterior margin of mesoscutum (Fig. 186); T1 discal patch triangular to semicircular, often only as wide as width of LLB; pseudopygidial area with basal and apical setae not strongly differentiated, but with apical setae slightly stouter and longer (Fig. 187) ...... T. remigatus (Fabricius) (Fig. 185) - Paramedian band clearly separated from other pale yellow setae on mesoscutum; T1 discal patch triangular to transversely ovate, wider than width of LLB in dorsal view; pseudopygidial area with basal and apical setae distinctly differentiated (Fig. 30) . . . . . T. charlesi Rightmyer, n. sp. (Fig. 29) 91(88) T1 discal patch strongly rectangular in shape (i.e., BTB and ATB parallel, and LLB forming 90 degree angle with transverse bands) ...... 92 - T1 discal patch shape variable but not strongly rectangular...... 99 92(91) Pseudopygidial area triangular to subquadrate, with apical margin straight or weakly convex, and with basal and apical setae strongly differentiated; T1 discal patch not forming plus-sign shape, with BTB and ATB often weakly interrupted medially (mesepisternum usually with sparse, short erect setae)...... 93 - Pseudopygidial area ovate, with apical margin strongly to weakly convex, and with basal and apical setae variable; T1 discal patch often forming plus-sign shape (with BTB and ATB medially interrupted for about same width as length of T1 discal patch, e.g., Fig. 235) (mesepisternum lacking erect, simple setae) ...... 96 93(92) Pseudopygidial area longitudinally elongate and triangular, with setae uniformly short, appressed, and dense, but color variable: with silvery setae on all margins enclosing median triangular region of dark setae (Fig. 175) (Utah west to California) . . . T. punctoclypeus Rightmyer, n. sp. (Fig. 174) - Pseudopygidial area not longitudinally elongate and triangular, setae variable ...... 94 94(93) Axilla slightly longer than wide, with distinct apical spine curved medially, reaching or surpassing scutellar midpoint; pseudopygidial area with well defined region of silvery setae on basal third (Fig. 6) to half (Fig. 177) of area (paramedian band of setae clearly separated from other regions of pale setae on mesoscutum) ...... 95 - Axilla as long as wide or shorter, lacking or with very small free apical spine, reaching or not

20 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER attaining scutellar midpoint; pseudopygidial area with vague or distinct basal region of silvery setae confined to basal fourth or less of area (Fig. 154). . . . . T. paenepectoralis Viereck (Figs. 151–153) 95(94) T1–T4 with ATBs interrupted medially (California: Mono Co., San Luis Obispo Co.)...... T. quadratus Rightmyer, n. sp. (Fig. 176) - At least T3–T4 with ATBs continuous (California: Antioch Co.) ...... T. antiochensis Rightmyer, n. sp. (Fig. 5) 96(92) S5 strongly downcurved; pseudopygidial area with well differentiated basal and apical setae . . 97 - S5 only moderately or not downcurved in profile; pseudopygidial area with poorly differentiated basal and apical setae ...... 98 97(96) T1–T4 with ATBs interrupted medially; legs black or dark brown (Puebla) ...... T. cruciformis Rightmyer, n. sp. (Fig. 41) - Only T1 and sometimes T2 with ATBs interrupted medially, T3 and T4 with bands uninterrupted; legs red (Texas and Northern México) ...... T. dilutus Rightmyer, n. sp. (Fig. 53) 98(96) Metanotum entirely covered with pale yellow setae, or with small patch of dark setae medially less than 1 OD in width (Southwestern U.S. and Northern México) ...... T. warriti Rightmyer, n. sp. (Fig. 234) - Metanotum with pale setae restricted to lateral corners, such that median patch of dark setae much wider than 1 OD ...... T. lateralis Rightmyer, n. sp. (Fig. 102) 99(91) T1 with BTB absent or nearly so; T2–T4 with ATBs uninterrupted medially; paramedian band usually reduced or absent; body length ca. 7–10 mm (México south to Costa Rica)...... 100 - T1 with BTB present, or if reduced then at least T2–T3 with ATBs medially interrupted; T2–T4 with apical banding variable; paramedian band usually strongly present; body length usually ca. 9– 14 mm ...... 101 100(99) T1 with ATB interrupted medially; clypeus lacking or with faint midline (México) ...... T. interruptus Rightmyer, n. sp. (Fig. 92) - T1 with ATB continuous medially; clypeus with strong midline (Costa Rica) ...... T. parkeri Rightmyer, n. sp. (Fig. 155) 101(99) Pseudopygidial area strongly triangular (i.e., apical and basal margins straight, and basal margins forming inverted V-shape at basal-most point), with distinct basal region of silvery setae (Fig. 97); axillar spine often with reddish tinge (paramedian band laterally contiguous with band of pale yellow setae on anterior margin of mesoscutum) (Southwestern and Midwestern U.S.) ...... T. isohedrus Rightmyer, n. sp. (Fig. 96) - Pseudopygidial area variable but not with the combination of strongly triangular shape and with distinct basal region of silvery setae; axillar spine usually lacking reddish tinge ...... 102 102(101) Paramedian band laterally contiguous with relatively dense band of pale yellow setae on anterior margin of mesoscutum (Fig. 186) (T1 discal patch reduced, ovate or triangular; T2 with LLB forming 90 degree or obtuse angle with ATB, resulting in basal brown region appearing rectangular or semicircular) ...... T. remigatus (Fabricius) (Fig. 185) - Paramedian band clearly separated from other yellow setae on mesoscutum, or if contiguous with other yellow setae, then T2 with LLB forming acute angle with ATB ...... 103 103(102) T1 discal patch triangular ...... 104 - T1 discal patch ovate or trapezoidal ...... 107 104(103) Pseudopygidial area 1.5 to 2 times as long as wide, or nearly so, with poorly differentiated region of silvery setae on basal third (Fig. 197) (T2–T4 with ATBs medially continuous, rarely T1 with ATB minutely interrupted medially) (Texas, Nuevo León, Tamaulipas, Jamaica, Grenada) ...... T. rufoclypeus (Fox) (Fig. 196) - Pseudopygidial area about as long as wide, or at most 1.5 times as long as wide, with silvery setae

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 21 distinct or poorly differentiated, restricted to crescent at base of pseudopygidial area ...... 105 105(104) Pseudopygidial area longitudinally ovate, with vague basal crescent of shining setae (Fig. 107) (Southwestern U.S. south to El Salvador and Guatemala) ...... T. laticeps (Friese) (Fig. 106) - Pseudopygidial area subquadrate, with distinct basal crescent of silvery setae ...... 106 106(105) Mesoscutum with at least some red integumental coloration (Southeastern U.S.)...... T. rufithorax Graenicher (Figs. 193, 194) - Mesoscutum black (U.S. and Northern México)...... T. lunatus (Say) (Figs. 110, 111) 107(103) Pseudopygidial area resembling that of Epeolus (i.e., basally poorly differentiated from rest of T5, with transverse band of silvery setae on apical margin, Fig. 123) (axillae short and rounded, with minute free apical spine) ...... T. melanarius Rightmyer, n. sp. (Fig. 122) - Pseudopygidial area variable but lacking transverse band of silvery setae on apical margin resembling that of Epeolus ...... 108 108(107) Pseudopygidial area wider than long, with medioapical tuft of long, silvery setae (Fig. 69) (Texas) ...... T. engeli Rightmyer, n. sp. (Fig. 68) - Pseudopygidial area usually at least as long as wide, lacking medioapical tuft of long, silvery setae ...... 109 109(108) Pseudopygidial area entirely with uniformly fine, dense setae, but with basal half silvery and apical half pale golden (S5 straight in profile) ...... 110 - Pseudopygidial area either with at least some coarse, sparse setae, and with apical setae dark brown rather than pale golden or if apical setae pale golden, then coarse setae indistinct and pseudopygidial area nearly entirely with pale golden setae ...... 111 110(109) Pseudopygidial area longitudinally subovate, rounded at apical margin (Fig. 34) (Northern México) ...... T. claytoni Rightmyer, n. sp. (Fig. 33) - Pseudopygidial area quadrate, truncate at apical margin (Fig. 115) (Texas) ...... T. margaretae Rightmyer, n. sp. (Fig. 114) 111(109) Integument of body entirely black (excluding mandible and antenna); dorsum of mesosoma and metasoma with bands of pale gray setae, resembling those of T. donatus (pseudopygidial area round; S5 moderately to distinctly downcurved) ...... 112 - Body coloration variable, but if legs black, then dorsum of mesosoma and metasoma with bands of yellowish setae ...... 114 112(111) T5 and S5 distinctly narrowed posteriorly, thus T5 with apical margin bearing correspondingly small, subquadrate pseudopygidial area, width at widest point ca. one fourth or less the width of T1 in dorsal view (Fig. 130); ventral half of lateral surface of mesepisternum with punctures minute, nearly contiguous (Mississippi) ...... T. micropygius Robertson, in part (Fig. 129) - T5 and S5 not so distinctly narrowed posteriorly, thus pseudopygidial area at its widest point ca. one third width of T1; in some places, ventral half of lateral surface of mesepisternum with punctures spaced a puncture diameter apart or more ...... 113 113(112) Ventral half of lateral surface of mesepisternum with integument raised between punctures; pseudopygidial area with basal shining crescent clearly distinct from apical coarse setae (Fig. 14) (New York and North Carolina west to Illinois and Texas) ...... T. atripes Mitchell (Fig. 13) - Ventral half of lateral surface of mesepisternum with integument flat between punctures; pseudopygidial area with basal shining crescent less clearly delineated from apical coarse setae (Fig. 81) (Florida, Georgia) ...... T. georgicus Mitchell (Fig. 80) 114(111) Pseudopygidial area longitudinally subovate, with apical margin convex, basal third to half with fine, silvery setae strongly contrasting with circular apical region of coarse setae (Figs. 10 and 130) AND S5 strongly downcurved (e.g., Fig. 36) (Midwestern and Western U.S.) ...... 115 - Pseudopygidial area variable, but if basal half to third with fine, silvery setae, then pseudopygidial

22 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER area quadrate with apical margin truncate; IF pseudopygidial area as above, THEN S5 slightly or not downcurved in profile (e.g., Fig. 2c)...... 116 115(114) Pseudopygidial area at widest point only ca. one fourth or less the width of T1, with shining setae only on basal third (Fig. 130) ...... T. micropygius Robertson (Fig. 129) - Pseudopygidial area at widest point one third the width of T1, with area of basal silvery setae occu- pying one half length of entire pseudopygidial area (Fig. 10) . . . T. argus Rightmyer, n. sp. (Fig. 9) 116(114) All legs black or dark brown (sometimes paler brown on tarsi) ...... 117 - At least hind leg orange or red, or at least tibiae of legs orange or red ...... 121 117(116) T1 with BTB and ATB parallel to each other, enclosing widely ovate to subrectangular region, with bands of pale setae uninterrupted medially or broken by less than 0.5 OD (California to Washing- ton, east to Colorado) ...... 118 - T1 with BTB and ATB variable, enclosing triangular or ovate region, with bands of pale setae medially interrupted by 1 OD or more (Arizona south throughout Central America) ...... 119 118(117) T2 with LLB forming 90 degree or weakly acute angle with ATB, or absent; pseudopygidial area with distinct basal crescent of silvery setae (Fig. 154) (mesepisternum usually with at least some short, erect, simple setae) ...... T. paenepectoralis Viereck (Figs. 151–153) - T2 with LLB forming acute angle with ATB; pseudopygidial area lacking distinct basal crescent of silvery setae (Fig. 87) (mesepisternum lacking erect, simple setae)...... T. helianthi (Robertson) (Fig. 86) 119(117) Pseudopygidial area with distinct region of silvery setae on basal third to half (Fig. 220) (T1 discal patch ovate) (Jalisco to Chiapas)...... T. totonacus (Cresson) (Fig. 219) - Pseudopygidial area lacking or with poorly differentiated basal region of silvery setae ...... 120 (T. tepanecus species group) 120(119) T1 discal patch ovate; T1–T4 with ATBs medially interrupted or nearly so; body length ca. 8–10 mm (Arizona and Sinaloa) ...... T. mauropygus Rightmyer, n. sp. (Fig. 118) - T1 discal patch triangular to quadrate; T2–T4 with ATBs usually uninterrupted medially; body length ca. 9–14 mm in length (Arizona to El Salvador) ...... T. laticeps (Friese) (Fig. 106) 121(116) Pseudopygidial area with setae nearly uniformly pale brown or golden, with apical margin straight (Fig. 79) (T1 discal patch transversely ovate, with BTB and ATB interrupted medially; clypeus shining, with distinct larger punctures; body length ca. 10–12 mm) (Arizona, New Mexico)...... T. fulgidus Rightmyer, n. sp. (Fig. 78) - Pseudopygidial area variable, but if setae nearly uniform then appearing dark brown, or if nearly uniform pale golden brown then apical margin of pseudopygidial area rounded ...... 122 122(121) Pseudopygidial area longitudinally elongate and triangular to subquadrate; with apical margin straight; basally with narrow, inverted V-shaped region of shining setae (Fig. 158) (clypeus distinctly flat in profile, shining, black, and often contrasted with red scape, pedicel, F1, and labrum) (Arizona and New Mexico south to Durango and Baja California) ...... T. partitus Rightmyer, n. sp. (Fig. 157) - Pseudopygidial area variable, but if triangular then either not longer than wide, or with silvery setae on basal third ...... 123 123(122) Pseudopygidial area triangular, with length ca. equal to width, with relatively long, coarse setae on most of area except at base (Fig. 162) (paramedian band joined laterally to pale yellow setae on anterior margin of mesoscutum; Arizona, Colorado, New Mexico) ...... T. parvus Rightmyer, n. sp. (Fig. 161) - Pseudopygidial area variable but if triangular, then not nearly the same length as width, and lacking long, coarse setae on apical half ...... 124 124(123) Pseudopygidial area distinctly longer than wide, quadrate to subtriangular, with silvery setae on

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 23 basal third to half forming triangular shape (posterior margin of region of silvery setae mostly straight) (Fig. 148); clypeus shining, black (often with red apical margin), strongly contrasting with bright orange scape (mesepisternum with short, sparse, erect, simple setae) (Southwestern and Mid- western U.S. and Northern México) ...... T. norae Cockerell (Fig. 147) - Pseudopygidial area variable, but if with silvery setae on basal third to half, then these setae forming semicircular crescent rather than nearly triangular region; clypeus variable (sometimes exactly as described above) ...... 125 125(124) Pseudopygidial area with silvery setae strongly differentiated from dark, coarse setae, with silvery setae extending along lateral margins of pseudopygidial area, enclosing or nearly enclosing medioapical circular region of dark, coarse setae (Figs. 32 and 73)...... 126 - Pseudopygidial area with setae variable but if with silvery and dark setae segregated into distinct regions, then region of dark setae not so perfectly circular (sometimes with silvery setae on all margins of pseudopygidial area) ...... 127 126(125) Clypeus shining, with distinct larger punctures (sometimes sparsely covered with white setae); scape bright orange (rarely brown); T1–T4 with all transverse bands of pale setae uninterrupted medially, or T1–T2 with all transverse bands interrupted by 0.5 OD or less (Southwestern U.S. and Northern México) ...... T. circumculus Rightmyer, n. sp. (Fig. 31) - Clypeus mostly obscured by white setae; scape brown; T1–T2 with all transverse bands of pale setae medially interrupted, those of T1 interrupted by nearly 1 OD (Coahuila)...... T. exilicurvus Rightmyer, n. sp. (Fig. 72) 127(125) Pseudopygidial area ca. as long as wide, bordered on all margins by silvery setae enclosing median patch of dark, coarse setae (Fig. 150) ...... 128 - Pseudopygidial area variable but not distinctly bordered on all margins with silvery setae . . . . 129 128(127) T1–T4 with all transverse bands of pale setae relatively thick (ca. 2 OD in length), uninterrupted or barely interrupted medially on T1; axillar spine almost reaching posterior margin of scutellum and apically incurved; mesepisternum with punctures minute and nearly contiguous (Colorado and Idaho to Kansas and North Dakota) ...... T. occidentalis (Cresson) (Fig. 149) - T1–T4 with transverse bands of pale setae relatively narrow (ca. 1.5 OD or less), interrupted medially on T1; axillar spine reaching midpoint of scutellum or nearly so; mesepisternal punctures separated by up to 0.5 puncture diameter (Arizona and New Mexico) ...... T. circumculus Rightmyer, n. sp., in part (Fig. 31) 129(127) Dorsum of mesosoma and metasoma with bands of white setae ...... T. michiganensis Mitchell (Fig. 127) - Dorsum of mesosoma and metasoma with bands of yellow to pale yellow setae ...... 130 130(129) All metasomal terga with transverse bands of pale setae uninterrupted medially ...... 131 - At least T1 with ATB medially interrupted...... 132 131(130) Clypeus shining, with distinct larger punctures; T2 with LLB forming strongly acute angle with ATB; paramedian band joined laterally to sparse, diffuse pale setae on anterior margin of mesoscutum, or mesoscutum entirely covered with sparse, diffuse pale setae ...... T. diffusus Rightmyer, n. sp. (Fig. 51) - Clypeus matte, lacking or with vague larger punctures; T2 with LLB forming weakly acute angle with ATB; paramedian band clearly separated or (rarely) joined laterally to pale setae on anterior margin of mesoscutum ...... T. helianthi (Robertson) (Fig. 86) 132(130) Mesepisternum mostly asetose except for patches of pale setae below scrobal groove and near pronotal lobe (mesepisternum very densely punctate) ...... T. cressonii (Robertson) (Fig. 39) - Mesepisternum with pale setae more widely distributed across dorsal third to half of sclerite . . 133 133(132) Hypoepimeron mostly covered with pale setae (sometimes this region of pale setae slightly sparser

24 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER than those on rest of dorsal half of mesepisternum) ...... 134 - Hypoepimeron mostly covered with dark brown or black setae ...... 136 134(133) Paramedian band joined laterally to patch of pale setae on anterior margin of mesoscutum ...... T. diffusus Rightmyer, n. sp., in part (Fig. 51) - Paramedian band clearly separated from other pale setae on mesoscutum ...... 135 135(134) Mesepisternum with medioventral spot of brown, branched setae...... T. townsendi Cockerell (Fig. 221) - Mesepisternum with ventral half entirely covered with sparse, pale yellow setae ...... T. vernus Rightmyer, n. sp. (Fig. 227) 136(133) All metasomal terga with ATBs interrupted medially (antennae entirely brown; legs orange except front leg with brown patches; mesepisternum usually with sparse, short, erect, simple setae) ...... T. medusa Cockerell (Fig. 120) - Metasomal terga with ATBs uninterrupted medially on at least T4 ...... 137 137(136) Pseudopygidial area quadrate, with weakly differentiated region of shining setae forming relatively narrow, inverted U shape along margin of ca. basal fourth or less of area (mesepisternum shining beneath pale setae) ...... T. laticaudus Cockerell (Fig. 104) - Pseudopygidial area rounded, often slightly longitudinally ovate, with weakly differentiated region of shining setae more broadly covering basal third or half of area ...... 138 138(137) T1 with BTB and ATB parallel along most of width, enclosing relatively short discal patch (ca. 2 OD in length or little more)...... T. helianthi (Robertson) (Fig. 86) - T1 with BTB and ATB not parallel or only parallel medially, enclosing longer, widely quadrate or subtriangular discal patch (ca. 3–4 OD in length) ...... T. laticeps (Fig. 106) 139(3) Mesepisternum with erect, simple setae ...... T. nayaritensis Rightmyer, n. sp. (Fig. 136) - Mesepisternum lacking erect, simple setae ...... 140 140(139) Mesepisternum with minute, contiguous punctures; mesoscutum with anterior fourth covered by dense, yellow, appressed, plumose setae, mostly indistinctly separated from paramedian band of setae (Fig. 37); T1 discal patch usually reduced to small rectangular or circular black region; T2 with wide LLB, with remaining basal region of black setae semicircular (sometimes rectangular) ...... T. concavus (Cresson) (Frontispiece, Figs. 37, 38) - Mesepisternum with small punctures, mostly separated by one puncture diameter or more, integument between punctures shining; mesoscutum lacking dense, appressed, plumose, pale setae along anterior margin, paramedian band of setae clearly separated from other pale setae on mesoscutum; T1 discal patch widely rectangular to subrectangular; T2 with LLB relatively narrow, forming acute angle with ATB ...... T. penicilliferus (Brues) (Figs. 167, 168).

Key to the males and females of the United States east of the Mississippi River and Eastern Canada

1 Dorsum of mesosoma and metasoma with pale gray to white banding ...... 2 - Dorsum of mesosoma and metasoma with pale to bright yellow banding ...... 11 2(1) Preoccipital carina strong on gena and dorsally behind vertex (Fig. 144); mesoscutum lacking paramedian band of pale setae (mesoscutum anteriorly and pronotal collar with sparse, erect white setae)...... T. nigrihirtus Mitchell (Fig. 141) - Preoccipital carina absent behind vertex, but at least partially present on gena; mesoscutum rarely lacking paramedian band of pale setae ...... 3 3(2) Mesepisternum with long (at least 1 OD in length), erect, simple setae (e.g., Fig. 166) ...... 4 - Mesepisternum lacking erect, simple setae or with such setae less than 0.5 OD in length ...... 6

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 25 4(3) Clypeus with apical margin surpassing lower tangent of compound eyes by at least 1.5 OD when face viewed with both dorsal and ventral margins of compound eyes in focus (Fig. 58); female pseudopygidial area with basal crescent of shining silver setae; mesoscutum with paramedian band present in both sexes ...... 5 - Clypeus with apical margin below lower tangent of compound eyes by less than 1 OD; female pseudopygidial area uniformly shining silver (Fig. 24); female mesoscutum lacking paramedian band, male mesoscutum with paramedian band narrow and short . .T. brittaini Cockerell (Fig. 23) 5(4) Mesepisternum with integument between punctures rugose, tuberculate (Fig. 199); body length ca. 8.5–10 mm ...... T. rugosus Mitchell (male unknown) (Fig. 198) - Mesepisternum with integument between punctures relatively flat; body length ca. 10–13 mm ...... T. donatus (Smith) (Fig. 61) 6(3) Pseudopygidial area with apical margin slightly concave (e.g., Fig. 2b); clypeus lacking midline and distinct larger punctures; male pygidial plate relatively wide and triangular, lacking transverse basal ridge (axillar spine reaching or exceeding scutellar midpoint, pointed apically) ...... T. obliteratus Graenicher4 - Pseudopygidial area with apical margin weakly or strongly convex; male at least with distinct clypeal midline or with distinct transverse basal ridge on pygidial plate (often with both) ...... 7 7(6) Clypeus lacking midline; apex of axilla often rounded apically, not or barely reaching scutellar midpoint; pseudopygidial area with apical margin strongly convex, entire area distinctly circular (e.g., Fig. 2a); female S5 strongly downcurved; male pygidial plate narrow (ca. 1 OD in width), almost parallel-sided ...... Triepeolus n. sp. 15 - Clypeus with weak to strong midline; apex of axilla pointed apically, nearly reaching or surpassing scutellar midpoint; pseudopygidial area with apical margin weakly to moderately convex, entire area not so distinctly circular; female S5 not or moderately downcurved; male pygidial plate not as narrow (ca. 1.5 OD in width or more), but sometimes almost parallel-sided ...... 8 8(7) Legs distally and tegula usually orange; body length ca. 8–9 mm; clypeus with strong midline; pseudopygidial area subquadrate to subovate, apical margin weakly convex (Fig. 128); male pygidial plate nearly parallel-sided ...... T. michiganensis Mitchell (Fig. 127) - Legs and tegula usually brown but sometimes red; body length ca. 10–13 mm; clypeus with strong to moderate midline; pseudopygidial area longitudinally ovate, apical margin moderately convex; male pygidial plate nearly parallel-sided to subtriangular ...... 9 9(8) Clypeus flattened, with strong midline; mesepisternum with punctures separated by only 0.5 puncture diameter or less, integument between punctures raised, somewhat tuberculate; female S5 moderately downcurved; pseudopygidial area with elongate basal silvery region (Fig. 14); female paramedian band not reaching anterior margin of mesoscutum ...... T. atripes Mitchell (Fig. 13) - Clypeus convex in profile, with weak midline; mesepisternum with punctures separated by up to one puncture diameter, integument relatively flat between punctures; female S5 weakly downcurved; pseudopygidial area with less elongate basal silvery region; female paramedian band reaching anterior margin of mesoscutum ...... 10 10(9) T1 discal patch transversely ovate ...... T. georgicus Mitchell (Fig. 80) - T1 discal patch subtriangular ...... T. mitchelli Hurd (female unknown) (Fig. 131) 11(1) Mesepisternum with long (ca. 1 OD or greater), erect, simple setae (e.g., Fig. 166) ...... 12 - Mesepisternum lacking erect, simple setae or with such setae shorter than 0.5 OD ...... 13

4. Species belonging to the Triepeolus simplex species group; not treated further herein. 5. Species belonging to the Triepeolus verbesinae species group, to be described in forthcoming publication (Right- myer, in prep.).

26 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER 12(11) Clypeus with relatively weak midline; female mesoscutum lacking paramedian band, male mesoscutum with paramedian band narrow and short; pseudopygidial area uniformly shining silver, small and triangular (Fig. 24); female mesepisternum with punctures separated by up to two puncture diameters ...... T. brittaini Cockerell (Fig. 23) - Clypeus with strong midline; mesoscutum with paramedian band present in both sexes; pseudopygidial area subovate to subquadrate, with distinct basal silvery crescent (Fig. 164); female mesepisternum with punctures separated by up to 4–5 puncture diameters (Fig. 165) ...... T. pectoralis (Robertson) (Fig. 163) 13(11) Paramedian band indistinctly separated from diffuse yellow setae on anterior third or fourth of mesoscutum (Fig. 37); mesoscutum and mesepisternum with fine, nearly contiguous punctures; T2 with basal black region semicircular to slightly quadrate (Fig. 38); female S5 very strongly downcurved; pseudopygidial area with long, erect setae forming rounded, three-dimensional structure (Fig. 35) (male metasoma ventrally entirely black or occasionally with white setae laterally on S3) ...... T. concavus (Cresson) (Frontispiece, Figs. 37, 38) - Paramedian band of pale setae usually clearly separated from other pale setae on mesoscutum, sometimes connected laterally to diffuse yellow setae on anterior of mesoscutum [if paramedian band continuous with other pale setae, then mesoscutum shining, with punctures spaced up to two puncture diameters apart in females, up to one puncture diameter apart in males (T. remigatus)]; T2 with basal black region variable but not semicircular; female S5 rarely strongly downcurved; pseudopygidial area variable but lacking long, erect setae forming rounded, three-dimensional posterior structure ...... 14 14(13) T1 mostly covered with appressed yellow setae except for small diamond-shaped, ovate, or rectangular patch of black setae mediobasally (pseudopygidial area rectangular, with setae uniformly very dense and short, setae at apical margin reflecting silver, apical margin straight, Fig. 56; axillar spine exceeding posterior margin of scutellum, apically incurved, often red) ...... T. distinctus (Cresson) (Fig. 55) - T1 discal patch not reduced as above, or if reduced to small patch of black setae then either paramedian bands connected laterally to pale setae on anterior margin of mesoscutum forming distinct anchor pattern on mesoscutum (T. remigatus) or pronotal collar nearly 2 OD in length (T. quadrifasciatus atlanticus) ...... 15 15(14) Pronotal collar with dorsal length ca. 2 OD (Fig. 181); paramedian band of pale setae clearly separated from other pale setae on mesoscutum, often reduced in width and length (pseudopygidial area lacking distinctly differentiated basal region of silvery shining setae) ...... T. quadrifasciatus atlanticus Mitchell (Fig. 178) - Pronotal collar with dorsal length only ca. 1 OD (if nearly 2 OD, then paramedian band laterally contiguous with yellow setae on anterior margin of mesoscutum) ...... 16 16(15) Pseudopygidial area triangular, with apical margin concave (e.g., Fig. 2b); female S5 straight in profile, with bristle-like setae on apical margin (e.g., Fig. 2c); male pygidial plate lacking distinct transverse basal ridge (clypeus of both sexes lacking midline) ...... 17 - Pseudopygidial area variable, but with apical margin weakly to strongly convex; S5 shape in profile variable but lacking dense bristle-like setae on apical margin; male pygidial plate with distinct transverse basal ridge (however, male of T. micropygius unknown) ...... 18 17(16) T2 with LLB absent or reduced, if reduced then LLB forming 90 degree angle with ATB; T1 discal patch rectangular to subtriangular; male S4 only with apical fringe of setae, S5 apical setae distinctly shorter than those on S4 ...... T. simplex Robertson6

6. Species belonging to the Triepeolus simplex species group; not treated further herein.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 27 - T2 with LLB forming acute angle with ATB; T1 discal patch transversely ovate; male S4–S5 with apical fringes of setae...... T. rhododontus Cockerell7 18(16) Mesoscutum with dense yellow setae on entire anterior margin (Fig. 139); female T5 usually lacking pale setae lateral to pseudopygidial area or with these setae restricted to apical margin of T5 (Fig. 140) (both sexes with mesoscutum shining and; T2 with LLB forming 90 degree angle with ATB) ...... T. nevadensis (Cresson) (Fig. 138) - Mesoscutum usually lacking dense yellow setae on anterior margin of mesoscutum, or if present then at least space between paramedian bands of pale setae lacking such yellow setae; female T5 with pale setae lateral to pseudopygidial area along most of pseudopygidial area length ...... 19 19(18) Mesoscutum with paramedian bands of setae joined laterally to pale yellow setae on anterior margin, forming well-defined anchor-shape of black setae on disc (Fig. 186) (T1 discal patch often reduced to diamond or ovate region; T2 with LLB forming obtuse angle with ATB; pseudopygidial area setae uniformly shining golden brown, lacking distinct basal region of differentiated setae, Fig. 187) ...... T. remigatus (Fabricius) (Fig. 185) - Mesoscutum with paramedian band of setae clearly separated from other pale setae on mesoscutum, not delimiting anchor-shaped patch of black setae on disc ...... 20 20(19) T1 discal patch widely rectangular with sinuate BTB and ATB; body length ca. 15–18 mm; integument entirely black except on mandible and F1; (axillar spine pointed, reaching posterior margin of scutellum; pseudopygidial area ovate, with uniformly darkly shining setae on entire area, Fig. 135)...... T. monardae Mitchell (Fig. 134) - T1 discal patch transversely ovate or triangular; body length less than 16 mm, usually ca. 9–15 mm; integument usually with at least some orange or red areas, especially on labrum, antennae, and/or legs ...... 21 21(20) Scutellum entirely or partially red; mesoscutum at least partly red; paramedian band absent or relatively short, not reaching anterior margin of mesoscutum ...... T. rufithorax Graenicher (Figs. 193, 194) - Scutellum and mesoscutum black; paramedian band variable, very rarely absent ...... 22 22(21) T1 discal patch triangular...... T. lunatus (Say) (Figs. 110, 111) - T1 discal patch transversely ovate ...... 23 23(22) Pseudopygidial area round and small, basal third with fine, silvery setae strongly contrasting with circular apical region of coarse setae (Fig. 130); S5 strongly downcurved ...... T. micropygius Robertson (male unknown) (Fig. 129) - Pseudopygidial area subtriangular to subquadrate, with basal silvery setae absent or present; S5 not or only weakly downcurved ...... 24 24(23) Female T1 and T2 (rarely also other terga) with transverse bands of pale setae medially interrupted; pseudopygidial area with setae mostly uniform in reflectance, density, and texture; shape of area typically longitudinally ovate (Fig. 87); male clypeus with apical fourth asetose...... T. helianthi (Robertson) (Fig. 86) - Female T1–T4 usually with transverse bands of pale setae medially interrupted; pseudopygidial area with distinct basal crescent of shining setae; shape of pseudopygidial area typically more longitudinally elongate, subtriangular to subquadrate (Fig. 40); male clypeus entirely covered with white setae ...... T. cressonii (Robertson) (Fig. 39).

7. Species belonging to the Triepeolus simplex species group; not treated further herein.

1 Body length ca. 16 mm or more; mesepisternum with long, erect, simple setae (axillar spine surpassing posterior margin of scutellum, sharply pointed, incurved) (Caribbean) ...... 2 - Body length ca. 14 mm or less; mesepisternum lacking or with very short, erect, simple setae (except T. atoconganus, from Ecuador and Peru, with long setae) ...... 3 2(1) T1 lacking BTB, with ATB uninterrupted or only narrowly interrupted medially; T2–T4 with ATBs uninterrupted medially (Dominican Republic) ...... T. victori Genaro (Fig. 232) - T1 with BTB present and both BTB and ATB interrupted medially; T2–T4 with ATBs interrupted medially (Cuba) ...... T. wilsoni (Cresson) (Fig. 237) 3(1) At least scutellum bright red, shining (T2–T3 with ATBs interrupted medially) (Caribbean, South America) ...... T. osiriformis (Schrottky)8 - Scutellum black (or, if red, then T2–T3 with ATBs not interrupted medially) ...... 4 4(3) T1 lacking ATB ...... 5 - T1 with ATB present ...... 6 5(4) Paramedian band reaching anterior margin of mesoscutum, tapering anteriorly; T2 with ATB yellow, at least slightly interrupted medially; male S3 with pale apical setae surpassing apical margin (female unknown) (Colombia) ...... T. flavipennsis (Friese) - Paramedian band of mesoscutum well separated from anterior margin, not tapering anteriorly; T2 with ATB not or very slightly interrupted medially; male S3 with pale apical setae not surpassing apical margin (female pseudopygidial area concave apically, with stout lateral setae) (Brazil) ...... T. alvarengai Moure9 6(4) At least T2–T3 with ATBs not interrupted medially (Caribbean, México) ...... T. rufoclypeus (Fox) (Fig. 196) - At least T2–T3 with ATBs interrupted medially ...... 7 7(6) Vertex and upper mesepisternum with dense, erect setae (Peru) ...... T. atoconganus Moure10 - Upper mesepisternum lacking erect setae; vertex with erect setae mostly limited to small patch posterior to ocelli ...... 8 8(7) Axilla triangular, with free apical point only ca. 0.25 total length...... 9 - Axilla more strongly pointed, with free apical point ca. 0.5 or more total length ...... 10 9(8) T1 and T2 usually with LLB present; vertex between lateral ocellus and compound eye with punctures nearly contiguous (Ecuador, Peru) ...... 14 - T2 lacking LLB, T1 with LLB absent (Caribbean) or sometimes present (Colombia, Venezuela); vertex between lateral ocellus and compound eye with punctures separated by up to one puncture diameter ...... T. rufotegularis (Ashmead)11 10(8) T1 discal patch strongly rectangular; pseudopygidial area with elevated basal arch, apical margin weakly concave, often appearing to have small median notch on apical margin formed by lack of setae medially and apparently by weakly impressed, longitudinal line on integument (Caribbean) ...... T. roni Genaro12 - T1 discal patch transversely ovate, lacking distinctly angulate corners; pseudopygidial area lacking elevated subapical arch, apical margin straight or convex ...... 11

8. Species belonging to the Triepeolus verbesinae species group; not treated further herein. 9. Species belonging to the Triepeolus simplex species group; not treated further herein. 10. Species belonging to the Triepeolus verbesinae species group; not treated further herein. 11. Species belonging to the Triepeolus verbesinae species group; not treated further herein. 12. Species belonging to the Triepeolus simplex species group; not treated further herein.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 29 11(10) Mesepisternum mostly covered with dense, yellow or white, plumose setae, except for circular dark region medioventrally (Caribbean) ...... 12 - Mesepisternum with dense, white, plumose setae dorsally, mostly asetose ventrally, lacking distinct circular dark region (Venezuela) ...... Triepeolus n. sp. 213 12(11) Dorsal margin of head with strong median notch (Fig. 146) (female unknown) ...... T. nisibonensis Genaro (Fig. 145) - Dorsal margin of head evenly, weakly concave, lacking median notch ...... 13 13(12) Anterior surface of antenna (when directed dorsally) entirely orange; body length 6–8 mm; pseudopygidial area subrectangular, with apical tuft of shining setae (Fig. 44) ...... T. cuabitensis Genaro (Fig. 43) - Antenna orange or reddish only basal to F2, remainder brown; body length ca. 9–12 mm; pseudopygidial area subtriangular, with vague basal crescent of shining setae (Fig. 231) ...... T. vicinus (Cresson) (Fig. 230) 14(9) T1 with BTB medially interrupted by more than 2.25 OD; body length ca. 9–13 mm, ITW 1.9–2.3 mm ...... T. buchwaldi (Friese)14 - T1 with BTB medially interrupted by less than 2 OD; body length ca. 7.5–10 mm, ITW 1.6–1.8 mm ...... T. aguilari Moure15

Taxonomy

In this section, I treat all named and newly described species of Triepeolus, excluding those currently known to belong to the T. verbesinae and T. simplex species groups (defined below), in alphabetical order. The T. verbesinae and T. simplex species groups and their described species are listed starting on page 130. In the taxonomic histories below, details of the primary type specimens are presented in brackets. Addi- tional information regarding the types is found in parentheses within the brackets; for example, if plant data, or a more precise collecting date, is given in the original publication but is not found on the physical type label itself, then such data are provided in parentheses. Descriptions of new species refer to the holotype specimen, with variation found in other specimens presented in parentheses. The descriptions of previously described species refer to both genders, unless otherwise specified. In the descriptions, if the male of a certain species is stated to be unknown, it is not necessarily an indication of the rarity of that gender; rather it is the result of the fact that I focused primarily on females in this study and did not always discriminate males of very similar species. For each species, the listing of specimens examined and their repositories includes the primary type for that species unless I indicate that I have not seen it. Specimen repositories are listed by the city in which the repository is found. A complete listing of these repositories is in the Materials and Methods section. The following names were either proposed in Triepeolus, or were subsequently placed in Triepeolus, but are herein confirmed to belong to other genera: Epeolus scutellaris Say (correct generic placement), Epeolus flavofasciatus Smith and Triepeolus agaricifer Cockerell (new synonymy) [= Epeolus flavofasciatus (Smith)], Triepeolus banksi Cockerell [= Epeolus banksi (Cockerell)], Triepeolus fazi Cockerell [= Doeringiella gayi (Spinola)], Triepeolus minimus Robertson [= Epeolus minimus (Robertson)], and Triepeolus pruinosus Cock- erell [= Doeringiella holmbergi (Schrottky)]. The taxonomic histories of these names are in Appendix 1.

13. Species belonging to the Triepeolus verbesinae species group, to be described in a forthcoming paper (Rightmyer, in prep.). 14. Species belonging to the Triepeolus verbesinae species group; not treated further herein. 15. Species belonging to the Triepeolus verbesinae species group; not treated further herein.

Triepeolus antiguensis Cockerell 1949: 460 [Holotype: U. S. National Museum of Natural History No. 58537; %, Anti- gua (Sacatepéquez), Guatemala; 26 December].

Description.—Length ca. 9–11 mm; ITW 1.6–2.5 mm. Integument black, with red on apical half of mandible; entirely or mostly orange on F1, tibiae, and tarsi, excluding spurs, sometimes on labrum, scape, pedicel, and part of legs basal to tibiae (specimen from México); pale orange-brown on tegula; dorsum of mesosoma and metasoma with bands of orange-yellow setae. Clypeus with elevated midline, with very faint larger punctures laterally, sometimes covered with sparse white setae (much denser in males). Face dorsally and mesepisternum with erect, simple setae; erect setae on most of head and mesosoma. Paramedian band clearly separated from other pale setae on mesoscutum. Scutellum weakly to moderately bigibbous; axillar spine not attaining midpoint of scutellum. Mesepisternum covered with long, pale, minutely-branched, suberect to erect setae, sparser medioventrally and on hypoepimeron; punctures deeply impressed, nearly contiguous to separated by up to a puncture diameter. T1 sometimes lacking LLB (especially in specimens from Guatemala and Costa Rica); T1 discal patch transversely ovate to trapezoidal, ATB widely interrupted medially; T2 with LLB absent or forming acute angle with ATB. Female: Pseudopygidial area semicircular, basally with distinct crescent of finer, denser setae; S5 straight in profile. Male: Pygidial plate keyhole shaped, with distinct transverse basal ridge and apical downturned plate; S2–S3 with apical bands white, grading into sparser white setae basally (S3 with white setae slightly surpassing apical margin); S4–S5 with apical fringes of setae golden brown medially, white laterally. Comments.—This species has two female forms. One is found from Chiapas south to Costa Rica, and lacks the LLB on T1 (although the male holotype is from this region and has the LLB present, though weak); the other form is found from Chiapas north to Hidalgo, and has the LLB present on T1. In addition, these two forms appear to be active in different seasons. In all other respects the two forms are very similar, including the erect, simple and branched setae of the mesepisternum, the medially interrupted ATBs on T1 and T2, the isolated paramedian band, the coloration of the legs, and the type of pseudopygidial area. Further evidence may support segregating these two forms into distinct species. Triepeolus antiguensis is very similar to T. medusa; however, the two can be separated by the presence of long, erect, branched and simple setae on the mesepisternum of T. antiguensis (as opposed to only shorter, erect, simple setae on the mesepisternum of T. medusa). Distribution.—COSTA RICA: Heredia, Puntarenas, San José; GUATEMALA: Sacatepéquez; MéXICO: Chi- apas, Durango, Hidalgo, Michoacán, Morelos, Puebla. Seasonal Records.—20 July to 18 September (northern form); 16 December to 12 March (southern form). Specimens examined.—13 &, 1 % (BERKELEY, HEREDIA, LAWRENCE, LOGAN, WASHINGTON D.C.)

TRIEPEOLUS ANTIOCHENSIS new species (Figs. 5, 6)

Diagnosis.—This species can be distinguished by the following combination of features: the relatively short clypeus; the dense punctation of the ventral half of the lateral surface of the mesepisternum; the slightly longer than wide, incurved axillar spine; the rectangular T1 discal patch; the LLB on T2, which forms a 90 degree angle with the ATB on that tergum; the uninterrupted ATBs of T3–T4; the presence of pale setae lateral to the pseudopygidial area on T5; and the well-differentiated silvery setae of the pseudopygidial area. Trie- peolus antiochensis strongly resembles T. quadratus, but in that species the ATBs are broadly interrupted

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 31 medially on T3 and T4, and the integument of the labrum and legs is red. This species less strongly resembles T. edwardi, which can be distinguished due to the absent or reduced pale setae lateral to the pseudopygidial area, and the poorly differentiated basal silvery setae of the pseudopygidial area. Description of holotype female.—Length ca. 10 mm (7–10 mm); ITW 2.2 mm (1.6–2.2 mm). Integu- ment black, with red on apical half of mandible and outer surface of F1 (rarely lacking); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint midline and distinct larger punctures; face with transverse area of white setae at level of antennal sockets, excluding supraclypeal area, (covered with dense, white setae in males). Paramedian band robust and clearly separated from other pale setae on mesoscutum (surrounded by diffuse pale setae on entire or anterior half of mesoscutum in males). Scutellum strongly bigibbous; axillar spine triangular, slightly exceeding midpoint of scutellum, apical point slightly incurved. Mesepisternum with sparse, short, erect, simple setae (sometimes apparently lacking these erect setae); dorsal third, excluding hypoepimeron, with dense, pale yellow, branched setae; ventrally with short, black, branched setae (entire mesepisternum covered with dense, pale yellow, branched setae in males); punctation very small and dense, punctures in a few places separated by ca. half puncture diameter, with surrounding integument appearing raised, somewhat tuberculate (slightly more exaggerated in other female specimens). T1 discal patch narrowly rectangular; T2 with LLB forming 90 degree (or weakly acute) angle with ATB; all metasomal terga with ATBs not or only slightly medially interrupted. Pseudopygidial area subquadrate (or triangular), with basal setae fine, dense, and silvery, extending apicolaterally and partially surrounding apicomedial area of coarser, longer setae; (S4 sometimes with small, apicolateral patch of white setae); S5 straight in profile. (Male: Pygidial plate keyhole shaped, with distinct transverse basal ridge and moderately downturned apical plate; S2–S3 with apical bands of white setae usually restricted to laterally; S4–S5 with apical fringes of dark brown setae.) Comments.—This species is noteworthy for its apparent endemism to Antioch, California; I have not seen any specimens of this species collected in the last 30 years, suggesting that T. antiochensis may be scarce or possibly even extinct. More effort should be made to determine the status of this enigmatic species. It strongly resembles T. quadratus, which is also noteworthy for its apparently limited range in Mono Co. and San Louis Obispo Co., California. Distribution.—USA: California (Antioch). Floral Records.—Eriogonum sp. Seasonal Records.—23 August to 17 October. Holotype.—“Antioch Cal [California], IX [September]-9-40 [1940] // Presented by E. C. VanDyke Col- lector // M. G. Rightmyer Triepeolus Database Specimen No. 3538 // HOLOTYPE & Triepeolus antiochensis Rightmyer 2007” [California Academy of Sciences #18246] (SAN FRANCISCO). Paratypes.—USA: CALIFORNIA: Contra Costa Co., Antioch, 23 August 1892, E. C. VanDyke (1&, SAN FRANCISCO); 23 August 1936, E. C. VanDyke (7%, SAN FRANCISCO); 26 August 1955, W. R. Kellen (1%, NEW YORK); 9 September 1940, E. C. VanDyke (1&, SAN FRANCISCO); 10 September 1947, U. N. Lanham (1%, ANN ARBOR); 18 September 1938, E. C. VanDyke (1&, SAN FRANCISCO); 18 September 1977, R. J. McGinley, Eriogonum sp. (1%, WASHINGTON D.C.); 25 September 1951, J. G. Rozen (1&, NEW YORK); Sep- tember 1936 (1&, RIVERSIDE); 17 October 1937, E. C. VanDyke (1&, SAN FRANCISCO). Etymology.—This species is named after its collection locality, Antioch, California.

TRIEPEOLUS ARGENTIMUS new species (Figs. 7, 8)

Diagnosis.—This species can be primarily distinguished by its unusual pseudopygidial area, which is entirely covered with dense, silvery, shining setae, and which also has a distinct apical, circular depression; in addi-

32 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER tion, the mesepisternum has long, erect, simple setae; the LLB on T2 forms an acute angle with the ATB; T5 has pale lateral setae; and S5 is strongly downcurved in lateral view. Triepeolus argentimus resembles T. heterurus and T. melanarius in the banding pattern of the metasoma, but females can be easily distinguished from them by the entirely unique form of the pseudopygidial area. Males can be separated by the plentiful, long, erect, simple setae in T. argentimus (the simple setae shorter and less abundant in T. heterurus and T. melanarius) and the more abundant pale setae on the metasomal sterna of T. argentimus. Description of holotype female.—Length ca. 11 mm (8–11 mm); ITW 2.0 mm. Integument black, with reddish orange on basal half of mandible, labrum, and apical margin of clypeus; reddish brown on pronotal lobe and legs basal to tibiae; orange on F1, tegula, and apical to femora (these features darker in males); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus lacking midline, with faint larger punctures. Paramedian band clearly separated from other pale setae on mesoscutum (all females and most males, or laterally contiguous with pale yellow setae on apical margin of mesoscutum in some males). Scutel- lum weakly bigibbous; axillar spine triangular, not attaining midpoint of scutellum. Mesepisternum and most of mesosoma with long, erect, simple setae; dorsal half and ventral margin covered with long, pale yellow, branched setae, sparser on hypoepimeron; medioventrally with wide, circular region of sparse, black, branched setae (male with white, erect, branched setae covering most of mesepisternum, sometimes with small medioventral patch of dark, branched setae); punctures nearly contiguous. T1 discal patch subovate, enlarged medially by indentations in BTB and ATB; T2 with LLB forming acute angle with ATB. Pseudopy- gidial area tear-shaped, entirely with dense, fine, silvery setae, apically with circular region of integument strongly depressed; S5 strongly downcurved. (Male: Pygidial plate relatively long and slender; S2–S3 with apical bands of white setae; S4 with white, and S5 with brown apical fringes of setae.) Comments.—This species appears to be endemic to Riverside Co., California, and has apparently not been collected since 1950, although other material may exist that I have not examined. A male specimen of this species bears an anonymous manuscript name that suggests the ground or a low position. Distribution.—USA: California (The Gavilan, Riverside Co.). Floral Records.—Coreopsis sp., Eriastrum sp., Eriogonum fasciculatum Benth., Helianthus gracilentus Gray. Seasonal Records.—12 May to 24 June. Holotype.—“Riverside Cal. [California], May 12, 28 [1928] // Timberlake Coll. // on garden Coreopsis // Triepeolus argentiopygus Timb. MS. // M. G. Rightmyer Triepeolus Database Specimen No. 6670 // HOLO- TYPE & Triepeolus argentimus Rightmyer 2007” [University of California, Berkeley #69999] (BERKELEY). Paratypes.—USA: CALIFORNIA: (Riverside Co.), Gavilan, 31 May 1937, Timberlake (1%, BERKE- LEY); 9 June 1950, Timberlake, Helianthus gracilentus (1&, LOGAN; 2%, BERKELEY); Eriogonum fasciculatum (1%, BERKELEY); 24 June 1938, Timberlake, Hugelia nigata (1%, BERKELEY). Etymology.—This name is in reference to the remarkable, silvery pseudopygidial area of the female. I follow Timberlake, who placed a manuscript label with a similar name on a specimen of this species. From the Latin, “argenteus” (silvery) and “imus” (having the quality of).

TRIEPEOLUS ARGUS new species (Figs. 9, 10)

Diagnosis.—Females of this species can be distinguished by the distinct pseudopygidial area, which is longer than wide and tear-shaped, and which has basal silvery setae that strongly contrast with a circular apical region of coarse, dark setae. It is additionally recognizable by the following combination of features: the relatively short clypeus; the isolated paramedian band of pale setae, the orange-colored legs, the lack of erect,

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 33 simple setae on the mesepisternum; the quadrate T1 discal patch; the presence of pale setae laterally on T5; and the strongly downturned female S5. Description of holotype female.—Length ca. 11 mm (10–15 mm); ITW 2.4 mm (2.0–2.7 mm). Integu- ment black, with clear red medially and reddish brown apically on mandible; reddish orange on outer surface of F1, outer margins of labrum, and outer surfaces of scape and pedicel; orange on legs, excluding bases of coxae and tibial spurs; pale brownish orange on tegula; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint midline and very faint larger punctures. Paramedian band isolated from (or sometimes laterally contiguous with) pale setae on anterior margin of mesoscutum. Scutellum moderately (sometimes strongly) bigibbous; axillar spine triangular, surpassing scutellum midpoint, apical point somewhat curved inward. Mesepisternum lacking erect, simple setae; with dense, pale yellow, branched setae on small region below scrobal groove and dorsally on anterior surface of mesepisternum; ventrally mostly asetose; punctures small, deeply impressed, and nearly contiguous. T1 discal patch widely trapezoidal (sometimes ovate), BTB and ATB slightly interrupted medially; T2 with LLB mostly on lateral surface of T2, forming weakly acute angle with ATB. Pseudopygidial area elongate, longitudinally ovate; basal half with dense, silvery, fine, pilose setae; this region of silvery setae with basal margin notched mediobasally (not as noticeable in some specimens); apical half with rounded region of coarse, sparse setae; S5 strongly downcurved. (Male: Unknown.) Distribution.—USA: Arizona, Colorado, Idaho, Kansas, Nebraska, New Mexico, North Dakota, Ore- gon, Utah, Washington. Host Records.—Melissodes (Eumelissodes) pallidisignata Cockerell? (1 specimen from nest site, Yakima Co., Washington); “going into Nomia, Melissodes, etc. nest holes” (1 specimen from Boulder Co., Colorado); “flying around ground where Melissodes nests” (1 specimen from Boulder Co., Colorado). Floral Records.—Chrysothamnus nauseosus [= Ericameria nauseosa (Pallas ex Pursh) Nesom & Baird], Grindelia squarrosa (Pursh) Dunal, Helianthus sp., Heterotheca villosa (Pursh) Shinners, Melilotus sp. Seasonal Records.—27 July to October (day unspecified). Holotype.—“Rhone N.D. [North Dakota], Aug. [August] 8, 1941, O. A. Stevens // Grindelia squarrosa // 15136 // M. G. Rightmyer Triepeolus Database Specimen No. 3576 // HOLOTYPE & Triepeolus argus Right- myer 2007” (NEW YORK). Paratypes.—USA: COLORADO: Boulder Co., 27 August 1925, C. P. Custer (1&, BOULDER); 2 Sep- tember 1925, C. H. Hicks (2&, BOULDER); 4 September 1926, C. H. Hicks (2&, BOULDER); 6 September 1925, C. P. Custer (3&, BOULDER); 8 September 1926, C. P. Custer (5&, BOULDER); 13 September 1926, C. H. Hicks (1&, BOULDER); 17 September 1925, C. P. Custer (2&, BOULDER); 18 September 1925, C. P. Custer (1&, BOULDER); Boulder, 5 September 1976, U. N. Lanham, 5800 ft., Heterotheca (1&, BOULDER); El Paso Co., Foster Ranch, F. M. Brown, 5700 ft., Heterotheca villosa (1&, BOULDER); Melilotus sp. (1&, BOULDER); Weld Co., 4 mi SW Hudson, 13 September 1964, L. R. Ertie (1&, ITHACA); Pawnee Grasslands, 13 August 1994, R. Parks (1&, SAN DIEGO); KANSAS: Phillips Co., 30 August 1912, F. X. Williams, 1940 ft. (1&, LAWRENCE); Riley Co., October, Marlatt (1&, WASHINGTON D.C.); NEBRASKA: Sioux Co., 7 mi N Harri- son, 13 August 1962, J. G. & B. L. Rozen, Helianthus sp. (1&, NEW YORK); 30 mi S Harrison, 9 August 1971, J. G., B. L., & K. C. Rozen (1&, NEW YORK). Additional specimens.—USA: ARIZONA: Cochise Co., Ft. Huachuca Garden Canyon, 31°48’N 110°32’W, 1–9 September 1997, B. Harris, (1&, LOS ANGELES); (Coconino Co.), Flagstaff, Sunset Crater, 18 August 1932, E. L. Bell, 7000 ft. (1&, NEW YORK); NEW MEXICO: Hidalgo Co., 27–32 mi S Animas, 24 August 1994, J. G. Rozen, J. S. Rozen (1&, NEW YORK); Otero Co., White Sands National Monument, 10 September 1983, L. D. French (1&, DAVIS); OREGON: (Umatilla Co.), Echo, 15 September 1924, E.S.G. Titus (1&, WASHINGTON D.C.); UTAH: Emery Co., 24 km NW Green River, 7 September 1990, T. Griswold (1&, LOGAN); Garfield Co., Tropic, 30 August 1985, D. K. Broemeling (1&, LOGAN); T. L. Griswold (1&, LOGAN); WASHINGTON: Benton Co., 4 km NE Prosser, 29 September 1988, E. Miliczky, Chrysothamnus nauseosus (1&, ZILLAH); 15.5 km N Prosser, 24 September 1990, E. Miliczky, C. nauseosus (1&, ZILLAH);

34 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Yakima Co., 11 km SE Granger, 1 September 1993, E. Miliczky, C. nauseosus (1&, ZILLAH); 25 September 1993, E. Miliczky, C. nauseosus (1&, ZILLAH); 1 mi N Zillah, 13 September 1992, E. Miliczky, at Melissodes pallidisignata nest site (1&, ZILLAH). Etymology.—The specific name is in reference to the pattern of setae on the pseudopygidial area, which might fancifully be compared to the spot of a peacock’s tail. From the Latin, “argus” (the hundred-eyed guardian of Io, whose eyes became the spots in the peacock’s tail).

TRIEPEOLUS ARGYREUS Cockerell (Figs. 11, 12)

Triepeolus argyreus Cockerell 1907c: 60–61 [Holotype: U. S. National Museum of Natural History No.100019; %, North Yakima, Washington; 4 August 1903].

Description.—Length ca. 9–12 mm; ITW 1.5–2.5 mm. Integument black, with orange usually on F1, sometimes on labrum and part or entire legs, excluding basal halves of coxae and spurs; dorsum of mesosoma and metasoma with relatively broad bands of very pale yellow setae. Clypeus lacking or with midline dorsally, with distinct larger punctures. Paramedian band intermixed with sparse white setae on anterior mesoscutum, sometimes poorly distinguished from them. Scutellum weakly bigibbous; axillar spine triangular, not attaining midpoint of scutellum, sometimes with apical point rounded. Mesepisternum with long, erect, simple setae; dorsal half with dense, pale yellow, branched setae; ventral half with similar, but less dense setae, or with brown, branched setae; medially with punctures nearly contiguous to separated by 0.5 puncture diameter, with integument between punctures raised, tuberculate. T1 discal patch widely rectangular, sometimes reduced to smaller black region medially; T2 with LLB forming 90 degree angle with ATB. Female: Pseudopygidial area semicircular to subquadrate, with distinct basal crescent of silvery setae; S5 slightly downcurved apically. Male: Unknown.

TABLE 1. Characterization of the species within the Triepeolus paenepectoralis species group.

T. eldoradensis T. argyreus T. paenepectoralis

Leg color pale orange brown to black (rarely orange) brown to black Mesepisternum covered with pale yellow, covered with pale yellow, pale yellow, appressed, plu- appressed, plumose setae appressed, plumose setae or mose setae usually restricted with pale setae restricted dor- dorsally, rarely covering most sally of mesepisternum Mesoscutum mostly covered with pale paramedian band of setae paramedian band of setae dis- yellow setae, obscuring mostly distinct, but often sur- tinct, often reduced, or some- paramedian band of setae rounded by diffuse pale yellow times connected to lateral setae setae, esp. anteriorly Scutellum mostly covered with pale sometimes covered with pale not covered with pale setae yellow setae yellow setae T1 transverse bands continuous medially continuous medially continuous or interrupted medi- of pale setae ally Pseudopygidial area basal and apical setae usu- basal and apical setae usually basal and apical setae usually ally poorly differentiated well differentiated well differentiated Distribution Colorado, Idaho, Utah, California, Idaho, Nevada, Alberta, British Columbia, Cal- Wyoming Oregon, Washington ifornia, Colorado, Idaho, Nevada, Oregon, Utah, Wash- ington, Wyoming

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 35 Comments.—This species resembles T. eldoradensis and T. paenepectoralis due to the presence of long, erect, simple setae on the mesepisternum, the 90 degree angle formed by the LLB and ATB on T2, the distinct basal shining crescent of setae on the pseudopygidial area, and the relatively short clypeus. Due to these similarities, T. paenepectoralis, T. eldoradensis, and T. argyreus are assembled in a group here termed the “T. paenepectoralis species group.” See Table 1 for differentiating characteristics among these species. Distribution.—USA: California (as far south as San Bernardino Co.), Idaho, Nevada, Oregon, Washing- ton. Host Records.—Melissodes (Eumelissodes) pallidisignata Cockerell (1 female specimen reared from cell by E. Miliczky; 11 females captured from nest site, in Yakima Co., Washington). Floral Records.—Chrysothamnus nauseosus [= Ericameria nauseosa (Pallas ex Pursh) Nesom & Baird], Hemizonia pugens (Hook. and Arn.) Torrey and Gray, Machaeranthera canescens (Pursh) Gray, Melilotus sp. Seasonal Records.—4 July to 20 September. Specimens examined.—25 & (CORVALLIS, DAVIS, GAINESVILLE, LOS ANGELES, SAN FRANCISCO, WASHINGTON D.C., ZILLAH).

TRIEPEOLUS ATRIPES Mitchell (Figs. 13, 14)

Triepeolus micropygius atripes Mitchell 1962: 474, Fig. 112 [Holotype: U. S. National Museum of Natural History No. 75245; &, Holly Shelter (Pender Co.), North Carolina; 18 October 1952].

Description.—Length ca. 10–13 mm; ITW 2.0–2.5 mm. Integument black, with red on apical half of mandible and sometimes F1; dorsum of mesosoma and metasoma with bands of pale gray to white setae. Clypeus relatively flat, with strong midline; integument entirely covered with white, medially-directed setae, or partially or entirely asetose, with no or very vague larger punctures. Paramedian band clearly separated from other pale setae on mesoscutum (females and some males) or connected laterally to diffuse white setae on anterior margin of mesoscutum (some males). Scutellum moderately to strongly bigibbous; axillar spine triangular, reaching or surpassing midpoint of scutellum, sometimes weakly incurved apically. Mesepisternum lacking erect, simple setae; punctures irregular and fairly dense, in places separated by up to 2, sometimes 3, puncture diameters; integument between punctures raised and shining. T1 discal patch transversely ovate; T2 with LLB forming weakly acute angle with ATB. Female: Pseudopygidial area longitudinally ovate, with elongate, somewhat elevated basal crescent of silvery shining setae; S3–S4 with lateral bands of white setae on apical margins; S5 weakly to moderately downcurved in profile. Male: Pygidial plate of moderate size, weakly keyhole shaped, with weak basal transverse ridge; S2–S3 with apical bands of white setae; S4–S5 with apical fringes of brown setae (S4 sometimes with small patch of white setae on apicolateral margin). Comments.—This species strongly resembles T. georgicus, but in T. atripes the clypeus is flattened and has a robust midline; the mesepisternum is more densely punctured and the integument between the punctures is raised and slightly tuberculate in appearance; and in the females, the paramedian band does not reach the anterior margin of the mesoscutum, the pseudopygidial area has an enlarged basal crescent of shining setae, and S5 is slightly more downcurved. Triepeolus atripes is also similar to T. donatus and T. rugosus, but unlike the latter two species, T. atripes has a relatively short clypeus and lacks the long, erect, simple setae on the mesepisternum. There are two female specimens from Pennsylvania, Coleman Park, that were identified as T. atripes by Mitchell (one is a paratype); however, I tentatively identify these specimens as T. cressonii, albeit with somewhat paler than usual yellow banding. Triepeolus atripes can be distinguished from T. cressonii by the pale gray to white colored bands, and the nearly entirely black integument (as opposed to pale yellow bands, usually with at least some areas of red integument on the legs and face in T. cressonii).

36 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Distribution.—USA: Arkansas, Georgia, Illinois, Maryland, Mississippi, Missouri, North Carolina, New York, Pennsylvania, Texas, Virginia. Floral Records.—Aster lateriflorus [= Symphyotrichum lateriflorum (L.) A.& D. Löve var. lateriflorum], A. pilosus [= Symphyotrichum pilosum (Willd.) Nesom var. pilosum], Chrysopsis mariana (L.) Ell., C. microcephala [= Pityopsis graminifolia (Michx.) Nutt. var. tenuifolia (Torr.) Semple & Bowers], Helianthus sp., and Solidago sp. Seasonal Records.—2 July to 4 November. Specimens examined.—44 &, 31 % (AUSTIN, DAVIS, ITHACA, LAWRENCE, LAWRENCE-BAKER, NEW YORK, RALEIGH, SAN FRANCISCO, STARKVILLE, URBANA, WASHINGTON D.C.).

TRIEPEOLUS AZTECUS (Cresson) (Figs. 15–17)

Epeolus aztecus Cresson 1878: 89–90 [Lectotype: Academy of Natural Sciences No. 2238; &, México]; Cresson 1916: 113 [designation of lectotype]. Epeolus flavocinctus Friese 1917 [1916]: 336 [Neotype: Zoologisches Museum, Humboldt-Universität; &, San Mateo (Alajuela), Costa Rica; 1904 (See Comments below)]. new synonymy, new neotype designation Epeolus costaricensis Friese 1925: 32–33 [Lectotype: Zoologisches Museum, Humboldt-Universität; &, San José, Costa Rica; 11 April 1903 (See Comments below)]. new synonymy, new lectotype designation Epeolus albopictus Cockerell 1949: 459 [Holotype: U. S. National Museum of Natural History No. 58535; %, San Fran- cisco finca, near Zamorano (El Paraíso), Honduras; 3 November]. new synonymy Triepeolus aztecus; Brumley 1965: 73.

Description.—Length ca. 7.5–11 mm; ITW 1.9–2.2 mm. Integument black, with red on apical half of mandible; orange on F1; dorsum of mesosoma and metasoma with bands of yellow to pale yellow setae. Clypeus lacking or with very faint midline, with very faint larger punctures. Paramedian band absent or somewhat sparse, short, and close together medially on anterior margin of mesoscutum. Scutellum moderately bigibbous; axillar spine triangular, reaching or slightly surpassing midpoint of scutellum. Mesepisternum lacking erect, simple setae; dorsally with dense, pale, branched setae below scrobal groove and along posterior margin; ventrally with sparse, brown, branched setae; with punctures mostly small and nearly contiguous, but also with some areas of impunctate, raised integument ca. 0.5 to 1 puncture width in size. T1 with pale yellow setae almost entirely covering dorsal surface, except for black longitudinal line and semicircular black spot medially on basal margin; T2 lacking LLB. Female: Pseudopygidial area subquadrate, with mostly uniform setae, with slight region of paler reflectance basally. Male: Pygidial plate keyhole shaped, with distinct transverse basal ridge and apical downturned plate; S2–S3 with apical bands of white setae, S3 with white setae slightly surpassing apical margin; S4–S5 with apical fringes of brown to pale golden setae. Comments.—This species is identified by the unusual pattern of pale setae on T1 and relatively small body size. At the Berlin museum there are two specimens (a male and a female) with Friese “Type” labels for Epeo- lus flavocinctus. The male specimen is from the same collection locality as in the original description (San José), but was apparently collected in 1923 and identified by Friese in 1925 (well past the publication date of 1917); in addition, the original description was of a female, and the description of T1 does not match that of the male specimen (which is actually T. parkeri). Instead, I have designated the female specimen as the neotype, despite being from the wrong (but nearby) locality, because it is the correct gender and because the distinctive T1 matches that of the original description. The female specimen cannot be part of the original type series, as the species was described based on only one female from a different locality; thus the specimen is designated as a neotype. The complete label data for this specimen are as follows: “Costa Rica, San Mateo, 1904 // Epeolus flavocinctus Fr. & 1915 Friese det // Type [red label] // Neotype & Epeolus flavocinctus Friese 1916 des. Rightmyer 2005.” Designation of a neotype is essential in order to stabilize the use of the name.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 37 Also at the Berlin Museum, there is a series of one female and six males identified by Friese as Epeolus costaricensis, all from the original type locality. As the holotype specimen was not designated by Friese in the original publication, both males and females are described therein, the female specimen bears a type label, and females are typically more readily identifiable, I am designating the female specimen as the lectotype. The complete label data for the lectotype of Epeolus costaricensis are as follows: “Costa Rica, San José, 4.11.1903 [the number ‘21’ appears to be handwritten over the date 1903] // Epeolus costaricensis Fr. & 1910 Friese det. // Type [red label] // Lectotype & Epeolus costaricensis Friese 1925 des. M. Rightmyer 2005.” Distribution.—COSTA RICA: Alajuela, Guanacaste, San José; HONDURAS: El Paraíso; MéXICO (state unspecified). Floral Records.—Hibiscus rosasinensis L. Seasonal Records.—9 November to 25 December. Specimens examined.—5 &, 2 % (BERLIN, HEREDIA, LOGAN, PHILADELPHIA, WASHINGTON D.C.).

TRIEPEOLUS BALTEATUS Cockerell (Figs. 18–20)

Triepeolus balteatus Cockerell 1921: 5 [Holotype: American Museum of Natural History No. 25084; %, Denver, Colo- rado; 28 August 1919]. Triepeolus brunneus Cockerell 1921: 7 [Holotype: American Museum of Natural History No. 25087; &, Lawn Lake, Rocky Mountain National Park, Colorado; 10,000 ft; 22 August 1919]. new synonymy

Description.—Length ca. 8.5–11.5 mm; ITW 1.8–2.2 mm. Integument black, with red to orange on apical half of mandible and F1, entire tegula, and legs, excluding basal halves of coxae and spurs, sometimes on part of labrum; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint midline and larger punctures (females), or entirely covered with dense, white, appressed setae (males). Preoccipital carina very strongly developed along entire length of gena, outcurved. Paramedian band laterally contiguous with diffuse apical pale setae (medially, between paramedian bands, mostly asetose in females, more densely setose in males). Scutellum weakly bigibbous; axillar spine triangular, not or barely reaching midpoint of scutellum. Mesepisternum with erect, simple setae; dorsally with dense, pale yellow, branched setae, grading to somewhat sparser setae ventrally; punctation irregular, integument between punctures somewhat raised and shining (females), or entirely covered by dense, white, appressed setae (males). Intercoxal area developed into clasper-like structure. T1 discal patch widely rectangular; T2 with LLB forming 90 degree angle with ATB. Female: Pseudopygidial area semicircular, apical margin convex, with distinct, somewhat pilose, basal silvery setae; S5 moderately downcurved. Male: Pygidial plate keyhole shaped, with transverse basal ridge and downturned apical plate; S2–S3 with apical bands of white setae; S4–S5 with brown and golden brown to white apical fringes of setae. Comments.—Both sexes of this species are easily recognized by the clasper-like intercoxal area, the protruding preoccipital carina, and the erect, simple setae on the mesepisternum. The distribution of this species seems to be centered on the range of the Rocky Mountains, although it is also found in states such as Oregon and Nebraska. The names balteatus and brunneus were published simultaneously. As first reviewer, I have selected the former name, which also has page priority. Distribution.—CANADA: Alberta; USA: Colorado, Idaho, Montana, Nebraska, New Mexico, Oregon, Utah, Wyoming. Host Records.—“Nomia, Melissodes, etc. holes” (2 female specimens from Boulder, Colorado). Floral Records: Chrysopsis villosa [= Heterotheca villosa (Pursh) Shinners var. villosa], Chrysothamnus viscidiflorus (Hook.) Nutt., Grindelia squarrosa (Pursh) Dunal, Kochia prostrata [= Bassia prostrata (L.) A.J. Scott], Solidago missouriensis Nutt.

TRIEPEOLUS BIMORULUS new species (Figs. 21, 22)

Diagnosis.—This species primarily can be distinguished by the scutellum, in which extensive pale yellow setae encircle one black or reddish brown region of setae on each dorsal convexity. The only other species with this type of scutellum is T. simulatus, but the two species can be differentiated by the clypeus and ventral half of the mesepisternum, which are shining and sparsely punctate in T. bimorulus (densely punctate and not shining in T. simulatus), and by the pseudopygidial area, the setae of which transition relatively smoothly from relatively dense, fine, and silvery basally to coarse, sparse, and dark brown apically in T. bimorulus (rather than with a relatively abrupt, clear boundary between the two setal types in T. simulatus). Description of holotype female.—Length ca. 11 mm (7.5–12 mm); ITW 2.3 (1.4–2.4 mm). Integument black to reddish brown, with red grading to orange on basal half of mandible, labrum, and clypeus (darker in some specimens); orange on scape, pedicel, F1, legs excluding spurs, and tegula (brown in some specimens) (some specimens with orange on apex of axillar spine); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus shining, with midline dorsally and larger punctures (mostly covered with white setae in males). Paramedian band clearly separated from other pale setae on mesoscutum, reaching anterior margin and tapering anteriorly (some males with paramedian band contiguous with lateral setae on mesoscutum). Scutellum moderately (sometimes weakly) bigibbous, mostly covered with appressed setae so that only biconvexity lacks setae, forming distinctive ovate dark spot laterally on scutellum; axillar spine triangular, slightly surpassing midpoint of scutellum (shorter in some specimens). Mesepisternum with sparse, relatively short, erect, simple setae; dorsally with dense, pale, branched setae; ventrally mostly asetose; integument shining, with small, weakly impressed punctures, separated by a puncture width up to five puncture widths (slightly closer spaced in some female specimens; males with sparse, branched setae covering most of mesepisternum). T1 discal patch subtriangular (to transversely ovate), BTB very slightly interrupted medially (sometimes uninterrupted), ATB continuous medially; T2 with LLB forming acute angle with ATB; apical bands of all terga continuous. Pseudopygidial area subquadrate, with basal crescent of silvery setae; S5 straight in profile. (Male: Pygidial plate of moderate size, lacking strongly delineated apical downturned plane; S2–S3 with apical bands of white setae; S4–S5 with apical fringes of golden to pale golden setae.) Distribution.—MéXICO: Baja California Sur; USA: Arizona, New Mexico, Texas. Host Records.—Svastra (Idiomelissodes) duplocincta Cockerell (1 specimen collected as larva from nest, Pima Co., Arizona; reared to adult by J. G. Rozen, Jr.). Floral Records.—Aplopappus gracilis [= Machaeranthera gracilis (Nutt.) Shinners], Bahia absinthifolia var. dealbata (Gray) Gray, Eriogonum deflexum Torr., Ferocactus sp., Melilotus alba [= Melilotus officinalis (L.) Lam.]. Seasonal Records.—25 June to 10 October. Holotype.—“S.W.R.S. [Southwestern Research Station], 5 mi W Portal, 5400 ft., Cochise Co., Ariz. [Ari- zona], Aug. [August] 2, 1956 // Collected on Melilotus alba Desr. // C. & M. Cazier Collectors // M. G. Right- myer Triepeolus Database Specimen No. 1253 // HOLOTYPE & Triepeolus bimorulus Rightmyer 2007” (NEW YORK). Paratypes.—MÉXICO: BAJA CALIFORNIA SUR: Puerto Escondido, 25 June–10 July 1990, R. Shaver (1&, DAVIS); 11 mi NE Todos Santos, 16 September 1983, R. R. Snelling (1&, LOS ANGELES); La Paz, 10–12 October 1954, F. X. Williams (1&, SAN FRANCISCO); USA: ARIZONA: Cochise Co., 1 mi E Dou- glas, 16 August 1974, Rozens, Favreau (1&, NEW YORK); 1 mi N Douglas, 16 August 1962, E. G. Linsley,

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 39 Bahia absinthifolia dealbata (1&, DAVIS); 2 mi NE Portal, 30 July 1959, M. Statham (1&, NEW YORK); 5 mi W Portal, Chiricahua Mts., 1 August 1958, R. E. Rice (2&, DAVIS); 5 mi W Portal, S.W.R.S., 22 July 1956, M. A. Cazier (1&, 1%, YORK); 23 July 1956, C. & M. Cazier, 5400 ft., Melilotus alba (1%, NEW YORK); 24 July 1956, C. & M. Cazier, 5400 ft., Melilotus alba (2&, 1%, NEW YORK); 25 July 1956, C. & M. Cazier, 5400 ft., Melilotus alba (1&, NEW YORK); 2 August 1956, C. & M. Cazier, 5400 ft., Melilotus alba (1&, NEW YORK); 8 August 1956, C. & M. Cazier, 5400 ft., Melilotus alba (1&, NEW YORK); 4 mi E Willcox, 24 August 1992, J. G. Rozen (1&, NEW YORK); 26 August 1994, J. G. Rozen, J. S. Ascher (1&, NEW YORK); 28 August 1992, J. G. Rozen (1&, NEW YORK); 64 mi E Willcox, 16 August 1992, J. G. Rozen (1&, NEW YORK); Maricopa Co., 11 September 1957, Timberlake, Haplopappus gracilis (1&, WASHINGTON D.C.); Phoenix, 9 July 2003, K. C. Rozen, 33°36’26”N 111°59’12”W (1&, NEW YORK); 13 July 1996, K. C. Rozen (1&, NEW YORK); 7 August 1994, J. G. & K. C. Rozen (2&, NEW YORK); Ferocactus (1&, NEW YORK); Pima Co., 4 mi N Continental, 11 August 1964, M. E. Irwin (1&, RIVERSIDE); 8 km E Robles Jct, 32°4’N 111°15’W, 23 August 1997, R. L. Minckley (1&, LAWRENCE); Silver Bell Bajada Int. Biol. Program, J. L. Neff (2&, 2%, AUSTIN); 6 km E Silver- bell, 18 August 1998, R. Minckley, Eriogonum deflexum (1&, LAWRENCE); 22 mi WSW Tucson, 11 August 1983, J. G. & B. L. Rozen (1&, NEW YORK); 23 mi S Tucson, 1 August 1961, D. W. Ribble (1&, URBANA); Pinal Co., Red Rock, 8 July 1952, R. H. & L. D. Beamer et al. (2&, LAWRENCE); NEW MEXICO: Dona Ana Co., 5 mi N Anthony, 16 July 1958, W. F. Barr (1&, LOGAN); Las Cruces, 15 August 1967, R. M. Bohart (2&, DAVIS); Hidalgo Co., Animas, 26 August 1975, R. J. McGinley (1&, WASHINGTON D.C.); TEXAS: Brewster Co., Castolon, 19 July 1950, R. F. Smith (1&, NEW YORK); (El Paso Co.), El Paso, 18 August 1908, F. C. Pratt (1&, WASHINGTON D.C.). Etymology.—This name is in reference to the two ovate regions of dark setae on the scutellum. From the Latin, “bi” (two) and “morulus” (black, like a mulberry).

TRIEPEOLUS BRITTAINI Cockerell (Figs. 23, 24)

Triepeolus brittaini Cockerell 1931: 279 [Holotype: Canadian National Collection No. 3358; %, Kings Co., Nova Scotia, Canada; 31 July 1930; Epilobium]. Triepeolus charlottensis Mitchell 1962: 462–463, Fig. 112 [Holotype: Canadian National Collection No. 15270; &, Char- lotte Co., New Brunswick, Canada; 10 September 1956]. new synonymy

Description.—Length ca. 10 mm; ITW 1.9 mm. Integument black, with red on distal half of mandible; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint midline, faint large punctures present (female) or lacking, covered with setae (male). Paramedian band absent (female) or diffuse, narrow (male). Axillar spine triangular, relatively small (almost reaching midpoint of scutellum in females, reaching midpoint in males); scutellum weakly bigibbous (female), or moderately bigibbous (male), somewhat extended posteriorly and sloping ventrally, rugose. Mesepisternum with sparse, long, erect, simple setae; dorsally with sparse, pale yellow, branched setae (female), or mostly covered with pale, branched setae (male); punctation fine and somewhat irregular, punctures separated by up to two puncture widths in some spots, these areas elevated, shining (punctures denser in male). T1 discal patch widely rectangular to ovate, T1 with LLB mostly lacking, especially basally; T2 with LLB absent, or reduced and forming 90 degree angle with ATB; banding relatively narrow on metasoma. Female: Pseudopygidial area small, triangular, with uniformly silver setae; S5 not downcurved. Mesosoma and metasoma venter lacking pale setae. Male: Pygidial plate keyhole shaped (not notched subapically), with distinct apical downturned plate and weak basal transverse ridge; S2–S3 with apical bands of white setae (very slightly surpassing apical margin on S3); S4–S5 with apical fringes of setae white or a mixture of white and brown.

40 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Comments.—Triepeolus brittaini is extremely similar to T. subalpinus, but the latter species has the paramedian band present on the mesoscutum in both sexes, the metasomal banding is broader, and the mesepisternal punctation is denser. None of these characters are particularly strong, but I hesitate to synonymize the two with so few specimens available for examination. Triepeolus brittaini is currently known from New Brun- swick and Nova Scotia, Canada, while T. subalpinus is known from Alberta and Saskatchewan, Canada, south to eastern California, Arizona, and Kansas. Triepeolus brittaini is also similar to T. pectoralis in the erect, simple setae on mesepisternum and pale yellow banding on metasoma, but T. brittaini females can be recognized by the punctation of the mesepisternum, which is more closely spaced; the lack of the paramedian band on the mesoscutum; and the pseudopygidial area, which is uniformly silver. Potential characters to separate the males of these two species include the clypeal midline, which tends to be strong in T. pectoralis and weak in T. brittaini, and the leg coloration, which tends to be more orange in T. pectoralis. Distribution.—CANADA: New Brunswick, Nova Scotia. Floral Records.—Epilobium sp. Seasonal Records.—31 July to 10 September. Specimens examined.—2 &, 1 % (OTTAWA, RALEIGH).

TRIEPEOLUS CALIFORNICUS (Cresson) (Figs. 25, 26)

Epeolus californicus Cresson 1878: 86 [Lectotype: Academy of Natural Sciences No. 2219; &, California]; Cresson 1916: 114 [lectotype designation]. Triepeolus californicus; Hurd 1979: 2092.

Description.—Length ca. 8–11 mm; ITW 1.9–2.5 mm. Integument black, with red on distal half of mandible; orange on at least tibiae and sometimes on F1; dorsum of mesosoma and metasoma with bands of yellow- orange to yellow setae (paler yellow in males). Clypeus with midline and faint larger punctures, entirely covered with dense white setae in males. Paramedian band rather elongate, clearly separated from other pale setae on mesoscutum. Scutellum moderately bigibbous; axillar spine triangular, not or barely reaching midpoint of scutellum. Mesepisternum lacking erect, simple setae; densely covered with yellow, branched setae. T1 discal patch entirely enclosed by yellow setae, forming very broad, parallel-sided region with rounded lateral sides and circular medial spot (rarely not present); T2 with LLB forming strongly acute angle with ATB. Female: Pseudopygidial area subquadrate, composed of uniformly golden setae; S5 straight to very slightly downcurved in profile, sometimes with lateral patches of yellow setae. Male: Pygidial plate elongate, either lacking transverse basal ridge or with this ridge so basal on plate as to be obscured by preceding tergum; S2– S3 with apical bands of white setae (S3 with white setae slightly exceeding apical margin); S4–S5 with apical fringes of setae (brown on S5, white with brownish tinge on tips on S4). Comments.—Triepeolus californicus resembles T. joliae; however the two species can be separated based on the angle formed by the LLB and ATB on T2 (acute in T. californicus, obtuse or absent in T. joliae), by the paramedian bands (long and distinct in T. californicus, shorter and often connected laterally to diffuse pale setae on anterior margin of the mesoscutum in T. joliae), and by the color of the setae on the pseudopygidial area (golden in T. californicus, coppery in T. joliae). In addition, the flight season of T. californicus is in the summer and early fall, while that of T. joliae is in the spring. Distribution.—USA: California (Mendocino Co. and Calaveras Co. south to San Diego Co. and River- side Co.). Floral Records.—Eriogonum nudum Doug. ex Benth., Bigelovia sp. (=Ericameria or Machaeranthera sp.?), Hemizonia fasciculata (DC.) Torr. & Gray.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 41 Seasonal Records.—10 June to 21 September. Specimens examined.—20 &, 8 % (BOULDER, DAVIS, LAWRENCE, LOGAN, LOS ANGELES, PHILADEL- PHIA, RIVERSIDE, SAN FRANCISCO, WASHINGTON D.C.).

TRIEPEOLUS CAMERONI (Meade-Waldo) (Figs. 27, 28)

Epeolus bifasciatus Cameron 1907: 136 [nec. Cresson, nec. Jörgensen] [Lectotype: The Natural History Museum, Lon- don No. 17B.516; % (not &), México (see Comments, below)]. new lectotype designation Epeolus cameroni Meade-Waldo 1913: 97–98 [replacement name]. Triepeolus cameroni (Meade-Waldo); Brumley 1965: 73.

Description.—Length ca. 8.5–10 mm; ITW 1.6–2.3 mm. Integument black, with red to orange on most of mandible, scape, pedicel, F1, and tegula, sometimes on outer margins of labrum, pronotal lobe, and small patches on tibiae; dorsum of mesosoma and metasoma with bands of orange-yellow to yellow setae. Clypeus lacking or with weak midline, lacking larger punctures, sometimes covered with sparse pale setae. Pronotal collar often narrowed submedially. Paramedian band clearly separated from other pale setae on mesoscutum, not attaining or, uncommonly, barely reaching anterior margin of mesoscutum. Scutellum moderately to weakly bigibbous; axillar spine triangular, reaching or slightly surpassing midpoint of scutellum. Mesepister- num lacking erect, simple setae; dorsally with patch of dense, branched, pale yellow setae below scrobal groove and pronotal lobe; ventrally mostly asetose, integument shining, with punctures nearly contiguous to separated by up to two puncture diameters. T1 with bands of yellow setae restricted to lateral margin or basolateral corner of tergum, sometimes with narrow ATB laterally; T2 lacking LLB. Female: Pseudopygidial area subrectangular, with uniformly golden setae or with weakly differentiated basal region of denser, shinier setae; S5 straight in profile. Male: Pygidial plate keyhole shaped, with weak transverse basal ridge and apical downturned plate; metasomal sternal setae uniformly golden; S3–S5 with well-developed apical fringes of setae. Comments.—Triepeolus cameroni is easily confused with T. rufoclypeus; the two may be synonymous but I am keeping them distinct until more complete information on their distributions is obtained. The two species can be distinguished by the relative reduction of the ATB on T1 (more reduced in T. cameroni, less reduced in T. rufoclypeus), and, usually, by the collection locality (the two species appear to only overlap in eastern Mexico, in Tamaulipas; otherwise, T. cameroni appears to be found primarily in southern Mexico, Bel- ize, Honduras, and Costa Rica; while T. rufoclypeus is found as far north as Texas, as well as on the islands of Jamaica and Grenada). Triepeolus cameroni also resembles T. mexicanus; females are easily distinguished from those of T. mexicanus by the pseudopygidial area, which is apically concave in that species. Males of these species are less readily separated, but may be distinguished by the lack of pale paramedian bands in T. mexicanus, contrasted with the presence of pale paramedian bands in T. cameroni. In addition, in dorsal view, the pale setal area on the pronotal collar is uniformly broad in T. mexicanus, in contrast to its often medially or submedially narrowed condition in T. cameroni. In addition, T. cameroni resembles T. bilineatus, but can be separated from that species by the pseudopygidial area, which is circular, with a distinct basal crescent of shining setae in T. bilineatus. Males of the two species can be distinguished by the golden apical fringe of setae strongly surpassing the apical margin of S3 in T. cameroni, contrasted with the apical setae only slightly surpassing the apical margin of S3 in T. bilineatus. In addition, the paramedian band does not typically reach the anterior mesoscutal margin and is not anteriorly tapering in T. cameroni (but sometimes does in the related species T. rufoclypeus; reaching anterior margin and tapering anteriorly in T. bilineatus).

42 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER In The Natural History Museum, London, there are two male specimens labeled by Cameron as Epeolus bifasciatus types. Although Cameron’s original description of this species was headed “Epeolus bifasciatus sp. nov. &,” the remainder of the description is likely that of a male, based on given details of the pygidial plate. I have designated the better preserved male cotype as the lectotype specimen. The label data for the lectotype specimen are as follows: “Type // B. M. Type Hym. 17B.516 // Epeolus bifasciatus Cam. Type México // Cameron Coll. 1909-182. // In B. M. 1965 Under E. cameroni M-W. // % Sex of type // Lectotype % Epeolus bifasciatus Cameron = E. cameroni Meade-Waldo des. M. Rightmyer 2005.” Distribution.—BELIZE: Cayo; COSTA RICA: Alajuela, Guanacaste, Puntarenas, San José; HONDURAS: El Paraíso; MéXICO: Campeche, Chiapas, Quintana Roo, Tamaulipas, Veracruz, Yucatán. Floral Records.—Acacia tenuifolia (L.) Willd., Anoda cristata (L.) Schlecht., Lantana sp., “white Mimo- saceae”. Seasonal Records.—January (day unspecified) to 14 October. Specimens examined.—66 &, 21 % (BERKELEY, BOULDER, HEREDIA, ITHACA, LAWRENCE, LOGAN, LON- DON, NEW YORK, WASHINGTON D.C.).

TRIEPEOLUS CHARLESI new species (Figs. 29, 30)

Diagnosis.—This species is readily recognizable by the female pseudopygidial area, which has an apical region of exceptionally coarse and sparse setae that is clearly distinct from the basal region of fine and dense setae. The pseudopygidial area is almost reminiscent of the apical setae found in T. mojavensis or T. dacotensis, but T. charlesi can be separated from them by the combination of the relatively long clypeus; the paramedian band, which is clearly separated from other pale setae on the mesoscutum; and the relatively wide, triangular to transversely ovate T1 discal patch. Description of holotype female.—Length ca. 12 mm; ITW 3.0 mm (3.0–3.1 mm). Integument black, with orange on basal half of mandible, labrum, scape, pedicel, F1, and legs excluding coxae and spurs; (sometimes also on apical margin of clypeus); brown on tegula; dorsum of mesosoma and metasoma with bands of yellow setae. Clypeus slightly elongate, with elevated midline and faint larger punctures, apical margin with sparse white setae (less apparent in other specimens). Paramedian band clearly separated from other pale setae on mesoscutum. Scutellum moderately (to weakly) bigibbous; axillar spine triangular, relatively flat, reaching midpoint of scutellum. Mesepisternum lacking erect, simple setae; dorsal half with dense, yellow, branched setae, sparser on hypoepimeron; anterior surface with sparse, yellow, branched setae; punctation nearly contiguous to separated by one puncture diameter, integument between flat, matte. T1 discal patch transversely ovate (or triangular); T2 with LLB forming weakly acute angle with ATB. Pseudopygidial area subquadrate, basally with relatively long region of silvery setae, apically with wide region of very coarse, sparse, suberect setae; S5 straight in profile. (Male: Unknown.) Distribution.—USA: Texas (Val Verde Co., Terrell Co.). Seasonal Records.—22 September to 15 October. Holotype.—“TEXAS Val Verde Co., Devils R. [River] Dolan Falls, 12–15 Oct. [October] 1995, coll. A. W. Hook // DR 1 // M. G. Rightmyer Triepeolus Database No. 1305 // HOLOTYPE & Triepeolus charlesi Rightmyer 2007” (LAWRENCE). Paratypes.—USA: TEXAS: Val Verde Co., Devils River, Dolan Falls Preserve, 24–25 September 1994, A. W. Hook (1&, AUSTIN); Terrell Co., 10 mi SE Dryden, 22 September 1972, W. Hanson, J. Poff (1&, LOGAN). Etymology.—This species is named in honor of my father, Charles W. Rightmyer, for his guidance, cre- ativity, and optimism.

Diagnosis.—This species can be distinguished by the combination of its relatively small size; the pseudopygidial area, in which the predominantly basal, silvery setae almost entirely encloses an apical, circular region of coarser, darker setae; the relatively short and shining clypeus; the mesepisternum, which lacks erect, simple setae; the paramedian band, which is clearly separated from, or barely contiguous with, other pale setae on the mesoscutum; the acute angle formed by the LLB and ATB of T2; and the apical bands of the metasomal terga, which are uninterrupted medially or only interrupted by 0.5 OD or less. The form of the pseudopygidial area is similar to that of T. argus, in that there is a nearly enclosed, apical, circular region of coarser, darker setae present; however, in T. argus, the basal region of silvery setae that enclose the apical, darker setae is much longer than in T. circumculus. Triepeolus circumculus is also similar to T. diffusus, T. exilicurvus, T. laticaudus, and T. occidentalis. The clearly isolated, or only faintly contiguous, paramedian band helps to separate it from T. diffusus and T. exilicurvus, in which this band of setae is clearly laterally contiguous with other pale setae on the mesoscutum. The well-separated regions of basal silvery shining setae and apical coarse, dark setae of the pseudopygidial area also help distinguish T. circumculus from T. diffusus and T. occidentalis, in which these two setal types more gradually intergrade. Triepeolus circumculus can be differentiated from T. laticaudus by the mesepisternum, which has sparser punctures and more tuberculate-like integument between the punctures in T. laticaudus. Conversely, T. circumculus can be separated from T. occidentalis by the more densely punctate mesepisternum in T. occidentalis. Description of holotype female.—Length ca. 7 mm (7–10 mm); ITW 1.5 mm (1.5–2.0 mm). Integument black, with orange on basal half of mandible, labrum, apical margin of clypeus, scape (most specimens), pedicel (most specimens), F1, medially on pronotal lobe (most specimens), tegula, and legs excluding basal halves of coxae and spurs; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus shining, with very faint (or no) midline, with distinct larger punctures. Paramedian band very faintly contiguous with pale setae on anterolateral margin of mesoscutum (clearly separated from other pale setae in most specimens). Scutellum moderately bigibbous; axillar spine triangular, almost reaching (or reaching) midpoint of scutellum. Mesepisternum lacking erect, simple setae; dorsally and on lateral and ventral margins with dense, pale yellow, branched setae, sparser on hypoepimeron and ventrally on margins; medioventrally with black, circular, asetose area (sometimes with sparse, pale, branched setae medially); punctures relatively small, irregular, almost contiguous to separated by 0.5 puncture diameter in some places. T1 discal patch transversely ovate (to trapezoidal); T2 with LLB forming acute angle with ATB. Pseudopygidial area semicircular, basally and along lateral margins with silvery, pilose setae, mostly enclosing circular, apical area of coarse setae; S5 straight in profile (or slightly downcurved apically). (Male: Unknown.) Comments.— A specimen of this species was labeled “PCAM 25” by D. Yanega. Distribution.—MéXICO: Chihuahua; USA: Arizona, Colorado (Boulder Co.), New Mexico, Texas (Yoa- kum Co.). Host Records.—Xenoglossodes (= Tetraloniella) (1 specimen flying in nesting area, Cochise Co., Ari- zona). Floral Records.—Aster sp. (= Symphyotrichum), Baccharis glutinosa [=Baccharis salicifolia (Ruiz & Pavón) Pers.], Baileya multiradiata Harv., B. pleniradiata Harvey & Gray ex Gray, Chrysothamnus sp., Eri- ogonum sp., Heliopsis sp., Heterotheca subaxillaris (Lam.) Britt. & Rusby, Kallstroemia grandiflora Torr. ex Gray, Machaeranthera tanacetifolia (Kunth) Nees, Poliomintha incana (Torr.) Gray, Sphaeralcea sp., Verbe- sina encelioides (Cav.) Benth. & Hook. f. ex Gray. Seasonal Records.—7 May to 13 June, and 20 July to 13 October. Holotype.—“USA, Arizona: Cochise Co., Willcox, Sept. [September] 21, 1976, J. G. Rozen // M. G. Rightmyer Triepeolus Database Specimen No. 1301 // HOLOTYPE & Triepeolus circumculus Rightmyer 2007” (NEW YORK).

44 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Paratypes.— USA: ARIZONA: Cochise Co., Apache, 16 August 1972, J. G. & K. C. Rozen, R. McGinley (2&, NEW YORK); 17 August 1961, M. A. Cazier, Baccharis glutinosa (1&, NEW YORK); 19 August 1972, Rozen, McGinley (1&, NEW YORK); 20 August 1972, Rozen, Favreau (1&, NEW YORK); 25 August 1986, R. M. Bohart; 2 mi NE Apache, 20 August 1966, J. G. & B. G. Rozen (1&, NEW YORK); 3.2 km E Apache, 24 August 1988, A. Roig-Alsina, flying on nesting area of Xenoglossodes (1&, LAWRENCE); 3.3 km E Apache, 17 August 1992, B. N. Danforth, Eriogonum sp. (1&, WASHINGTON D.C.); 4 mi S Apache, 18 August 1982, R. M. Bohart (1&, DAVIS); 6 mi SE Apache, Skeleton Canyon, 1 September 1958, E. G. Linsley (1&, DAVIS); 8 mi SW Apache, 20 August 1994, J. G. Rozen, J. S. Ascher (1&, NEW YORK); 9 mi N Apache, 14 August 1959, E. G. Linsley (1&, DAVIS); 14 mi SW Apache, 20 August 1996, A. Pence, J. G. & B. L. Rozen (1&, NEW YORK); 21 August 1986, J. G. Rozen (1&, NEW YORK); 15 mi SW Apache, 22 August 1970, J. G. Rozen, Heterotheca (1&, NEW YORK); 18 mi SW Apache, 18 August 1994, J. G. Rozen, J. S. Ascher, Kallstroemia (4&, NEW YORK); 20 August 1994, J. G. Rozen, J. S. Ascher (3&, NEW YORK); 22–23 August 1994, J. G. Rozen, J. S. Ascher (1&, NEW YORK); Benson, 27 September 1960, G. E. Bohart, (3&, LOGAN); Aster sp. (2&, LOGAN); Bowie, 31 July 1986, B. Cutler (1&, LAWRENCE); Chiricahua Mountains, 28 Septem- ber 1960, G. E. Bohart, 8,000 ft., Heliopsis sp. (1&, LOGAN); Chiricahua Mountains, West Turkey Creek, 2 September 2003, J. G. Rozen et al. (1&, NEW YORK); Chiricahua National Monument, 23 August 1951, L. Martin (1&, LOS ANGELES); 14 September 1985, A. F. Shinn, 1554 m, Heterotheca subaxillaris (1&, LAWRENCE); Double Adobe, 23 August 1990, J. G. Rozen, J. Krieger (2&, NEW YORK); 1 mi E Douglas, 20 August 1968, Rozen, Favreau (1&, NEW YORK); 23 August 1967, J. G. Rozen, G. Krueger (1&, NEW YORK); 29 August 1967, J. H. & J. M. Davidson, M. A. Cazier (3&, TEMPE); Rozen, Favreau (2&, NEW YORK); 1 mi N Douglas, 16 August 1962, J. M. Linsley, Verbesina encelioides (1&, DAVIS); 4–5 mi NE Douglas, 26 August 1995, J. G. Rozen, S. A. Budick (1&, NEW YORK); 16 mi NE Douglas, 25 August 1962, J. G. Rozen, M. Statham, S. J. Hessel (1&, NEW YORK); 31 August 1962, J. G. Rozen, M. Statham, S. J. Hessel (1&, NEW YORK); 7 mi N Elfrida, 3 August 1961, E. G. Linsley, Kallstroemia grandiflora (1&, NEW YORK); 4 August 1961, E. G. Linsley, Baileya multiradiata (1&, NEW YORK); 9 mi N Elfrida, 5 August 1961, J. G. Rozen (3&, NEW YORK); Guadalupe Canyon, 18 August 1967, E. I. Schlinger (1&, RIVERSIDE); 3 mi E Palominas, 5 Sep- tember 1986, A. S. Menke (1&, WASHINGTON D.C.); near Portal, 8 August 1984, W. J. Hanson (1&, LOGAN); 30 August 1967, W. J. Hanson (1&, LOGAN); 2 mi NE Portal, 20 August 1989, E. Brewster, J. Rozen, R. S. Foster (1&, NEW YORK); 2.9 mi NE Portal, 24 August 1959, M. Statham (1&, NEW YORK); 3 mi E Portal, 23 August 1971, J. D. Pinto (1&, RIVERSIDE); 5 mi NE Portal, Southwestern Research Station, 12 August 1994, J. S. Ascher (1&, NEW YORK); 7 mi E Portal, 18 September 1962, J. Willcox (1&, RIVERSIDE); 8 mi NE Portal, 17 August 1990, J. G. Rozen, J. Krieger (1&, NEW YORK); 23 August 1990, J. G. Rozen, J. Krieger (1&, NEW YORK); 23–25 August 1989, J. G. Rozen, R. L. Foster (1&, NEW YORK); 28 August 1990, J. G. & B. L. Rozen (1&, NEW YORK); Skeleton Canyon, 4 October 1958, H. V. Weems, 4–5000 ft. (1&, GAINESVILLE); 8 km W Sunset, 25 August 1985, E. S. Ross (1&, SAN FRANCISCO); Willcox, 18 August 1985, J. G. Rozen (1&, NEW YORK); Baileya pleniradiata (2&, NEW YORK); 21 September 1976, J. G. Rozen (1&, NEW YORK); 4 mi E Willcox, 19 August 1989, J. G. Rozen, R. L. Foster (1&, NEW YORK); 22 August 1985, J. G. & B. L. Rozen (1&, NEW YORK); 23 August 1996, J. G. & B. L. Rozen, A. Pence (1&, NEW YORK); 30 August 1993, J. G. Rozen (2&, NEW YORK); Willcox Dry Lake, 25 August 1967, E. I. Schlinger (2&, RIVERSIDE). Additional specimens.— MÉXICO: CHIHUAHUA: 9 mi S Hidalgo del Parral, 19 July 1967, R. C. Gardner, C. R. Kovacic, K. Lorenzea (1&, DAVIS); 9 August 1967, R. C. Gardner, C. R. Kovacic, K. Lorenzea (2&, DAVIS); 73 km W C. Juárez, 30 August 1991, R. Ayala, 1160 m (1&, LAWRENCE); JALISCO: Chamela, 1–8 October 1985, F. D. Parker, T. L. Griswold (1&, LOGAN); USA: ARIZONA: Cochise Co., 2 mi E Apache, 7 May 1989, J. G. Rozen (1&, NEW YORK); 9 May 1986, J. G. Rozen (4&, NEW YORK); 11 May 1987, J. G. Rozen (2&, NEW YORK); 3.2 km E Apache, 17 May 1987, B. N. Danforth (1&, LAWRENCE); Coconino Co., 2 mi E Moenkopi, 13 June 1966, J. M. Davidson, M. A. Cazier, Poliomintha incana (1&, TEMPE); Grant Co., 0.5 km N Huachita, 31°55’24”N 108°19’33”W, 23 August 2001, R. Brooks, 1440 m, Machaeranthera

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 45 tanacetifolia (1&, LAWRENCE); Pima Co., Santa Catalina Mountains, Molino Basin, 8 May 1978, J. G. & B. L. Rozen (1&, NEW YORK); Santa Rita Mountains, 18 August 1968, R. M. Bohart (1&, DAVIS); 29 August 1976, W. J. Hanson (2&, LOGAN); Santa Cruz Co., Santa Rita Mountains, 5 September 1937, W. Benedict (1&, LAWRENCE); COLORADO: Boulder Co., 20 July 1926, C. H. Hicks (1&, BOULDER); 15 September 1926, C. P. Custer (1&, BOULDER); NEW MEXICO: (Dona Ana Co.), Las Cruces, C. F. Baker (1&, WASH- INGTON D.C.); (Eddy Co.), Loving, 8 May 1975, J. W. MacSwain (1&, SAN FRANCISCO); Hidalgo Co., S of Animas, 28 August 1977, R. J. McGinley (1&, WASHINGTON D.C.); 9 mi W Animas, 22 August 1977, R. J. McGinley (1&, WASHINGTON D.C.); 12 mi E Animas, 17 August 1967, R. R. Snelling (1&, LOS ANGELES); 20 mi S Animas, 8 September 1977, J. G. Rozen (2&, NEW YORK); 20 km W Animas, 6 September 1982, L. D. French (1&, DAVIS); 21 mi S Animas, Middle Well, 23 August 1973, Rozen, Favreau (1&, NEW YORK); 22 mi S Animas, 23 August 1973, Rozen, Favreau (1&, NEW YORK); 25 August 1973, J. G. & K. C. Rozen, M. Favreau, R. McGinley (1&, NEW YORK); 22–26 mi S Animas, 10 September 1976, R. J. McGinley (1&, WASHINGTON D.C.); 27–32 mi S Animas, 24 August 1994, J. G. Rozen, J. S. Ascher (1&, NEW YORK); Rodeo, 22 August 1958, D. D. Linsdale (1&, DAVIS); 7 September 1973, M. Favreau (1&, NEW YORK); 9 September 1973, M. Favreau (1&, NEW YORK); 1 mi N Rodeo, 13 August 1974, M. & T. M. Favreau (1&, NEW YORK); J. G. & K. C. Rozen (1&, NEW YORK); 19 August 1974, Rozen, Favreau (1&, NEW YORK); 25 August 1970, J. G. Rozen (1&, NEW YORK); 28 August 1970, J. G. Rozen (1&, NEW YORK); 28 August 1987, J. G. Rozen (1&, NEW YORK); 28–29 August 1987, J. G. & B. L. Rozen (1&, NEW YORK); 31 August 1987, J. G. & B. L. Rozen (1&, NEW YORK); 2 September 1972, M. & T. M. Favreau (3&, NEW YORK); 2 mi N Rodeo, 18 August 1962, H. A. Scullen, 4100 ft. (1&, CORVALLIS); 22 August 1996, J. G. & B. L. Rozen, A. Pence (1&, NEW YORK); 2.5 km N Rodeo, 12–20 August 1991, Sphaeralcea (1&, LAWRENCE); 4 mi SW Rodeo, 18 August 1977, R. J. McGinley (1&, WASHINGTON D.C.); 5 mi N Rodeo, 22 August 1988, Rozen, Quinter, Brewster (1&, NEW YORK); 26 August 1985, J. G. & B. L. Rozen; 7 mi N Rodeo, 18 August 1986, J. G. Rozen, H. Henter (1&, NEW YORK); 18 mi N Rodeo, 7 September 1976, R. M. Bohart (1&, DAVIS); 10 October 1964, R. R. Snelling (1&, LOS ANGELES); 31.5 mi SW Rodeo, 16 August 1986, R. J. McGinley, B. M. Norden (1&, WASHINGTON D.C.); Luna Co., 20 mi Deming (no direction given), 30 August 1977, R. M. Bohart (1&, DAVIS); Sandoval Co., 22 mi SW Santa Fe, Peña Blanca, 6 September 1983, Rozen, Favreau (1&, NEW YORK); TEXAS: Yoa- kum Co., 6 mi NW Plains, 18 September 1977, Hanson, Hsiao (2&, LOGAN). Etymology.—This name is in reference to the silvery setae of the pseudopygidial area, which mostly encloses a circular dark region of coarse setae. From the Latin, “circum” (around, on all sides) and “culus” (rump).

TRIEPEOLUS CLAYTONI new species (Figs. 33, 34)

Diagnosis.—This species is known by the unique form of the pseudopygidial area, which is entirely formed by fine, dense setae. These setae are segregated into two regions: a basal region with silvery reflectance, and an apical region with pale golden reflectance. This type of pseudopygidial area is similar to that found in T. margaretae and T. rohweri. However, in T. margaretae the apical margin is more truncate, giving the pseudopygidial area a quadrate appearance, while in T. claytoni the apical margin is rounded, giving it a more elongated, ovate shape; also in T. margaretae the line separating the basal, silvery setae and the apical, golden setae is indented medially, while in T. claytoni this line is straight. Triepeolus claytoni can be separated from T. rohweri by the much longer (ca. 1 OD vs. less than 0.5 OD) and much denser erect setae of the mesepisternum in T. rohweri; by the paramedian band, which is laterally contiguous with pale setae on the mesoscutum in T. rohweri, and clearly isolated from such pale setae in T. claytoni; and by the angle formed by the LLB and ATB on T2 (90 degrees in T. rohweri and acute in T. claytoni).

46 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Description of holotype female.—Length ca. 10.5 mm (7–10.5 mm); ITW 1.8 mm (1.5–1.8) mm. Integ- ument black, with dark red on mandible, scape, pedicel, F1, and pronotal lobe; orange on tegula and legs excluding basal halves of coxae, and parts of foretrochanter and forefemur; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint midline and larger punctures. Paramedian band clearly separated from pale setae on anterior of mesoscutum. Scutellum moderately bigibbous; axillar spine triangular, almost reaching scutellum midpoint. Mesepisternum apparently lacking or under very close magnification with short, sparse, erect, simple setae; dorsal half with dense, pale yellow to white, branched setae, except hypoepimeron with such setae much sparser; ventral half of mesepisternum with sparse pale setae along margins, medioventrally asetose or with sparse, branched, pale setae; punctures nearly contiguous to separated by a puncture diameter, with integument between punctures raised, tuberculate. T1 discal patch subquadrangular (or widely ovate); T2 with LLB forming acute angle with ATB. Pseudopygidial area longitudinally ovate, almost tear shaped, entirely composed of very dense, pilose setae, basal half with silvery setae contrasted with apical area of golden setae; S5 straight in profile (or faintly downcurved apically). (Male: Unknown). Distribution.—MéXICO: Chihuahua, Durango. Seasonal Records.—19 August to 11 September. Holotype.—“Yerbanis, Cuencame Dist., Dgo. [Durango] Mex. [México] 6700 ft., Aug. [August] 19, 1947 // D. Rockefeller Exp., Schramel // M. G. Rightmyer Triepeolus Database Specimen No. 1313 // HOLO- TYPE & Triepeolus claytoni Rightmyer 2007” (NEW YORK). Paratype.—MÉXICO: CHIHUAHUA: 10 mi W Jiménez, 11 September 1950, R. F. Smith (1&, NEW YORK). Etymology.—This species is named in honor of my uncle and sushi chef extraordinaire, Clayton S. Depue, for his friendship and support.

TRIEPEOLUS CONCAVUS (Cresson) (Frontispiece, Figs. 35–38, 263)

Epeolus concavus Cresson 1878: 85 [Lectotype: Academy of Natural Sciences No. 2241; &, California]; Brues 1903: 80 [illustration of female T5 and S5]; Cresson 1916: 115 [designation of lectotype]. Triepeolus concavus; Robertson 1901: 231; Linsley and Michener 1939: plates xv, xviii [illustration of female S6, anterior wing]; Mitchell 1962: 464–465, Figs. 111, 112 [redescription, floral records, illustrations of male genitalia, scutellum, axillae]; Bohart 1970: Fig. 19 [photograph of egg in cell wall of host]; Hurd et al. 1980: 28, 90, 91 [habitus illustration, visitation records for Helianthus species]; Rozen 1989b: 15, Figs. 24, 25 [description, illustrations of first instar]. Triepeolus concava; Minckley et al. 1994: 1415.

Description.—Length ca. 10–16 mm; ITW 2.2–3.1 mm. Integument entirely black or dark brown, except often ferruginous apically on mandibles, basolaterally on labrum, and laterally on F1; dorsum of mesosoma and metasoma with bands of yellow setae. Clypeus with weak midline and distinct larger punctures (partially obscured in specimens with brown/black setae on clypeus, especially males). Paramedian band indistinct from dense yellow setae covering anterior third to fourth of mesoscutum. Scutellum strongly to moderately bigibbous; axillar spine triangular, reaching midpoint of scutellum. Mesepisternum lacking erect, simple setae; with distinct region of yellow, branched setae on dorsal third to fourth; remainder black, covered with minute, contiguous punctures and black or dark brown, branched setae. T1 discal patch strongly rectangular to subovate (very reduced in some specimens from Arizona); T2 completely covered with pale yellow setae except for semicircular to trapezoidal basal black region. Female: Pseudopygidial area with long, stout setae forming posterior “plate”; S5 strongly downcurved; ventral meso- and metasomata entirely dark brown. Male: Pygidial plate relatively wide, with distinct basal transverse ridge; metasomal sterna with setae dark

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 47 brown except sometimes with white setae laterally on S3; S3 with apical setae slightly surpassing apical margin; S4–S5 with dark brown apical fringes. Comments.—Males of T. concavus might be mistaken for those of T. remigatus; however, in T. remigatus the paramedian bands and yellow setae on the anterior margin of the mesoscutum form a strong anchor pattern, while in T. concavus the paramedian bands are not distinct from the region of dense yellow setae on the anterior third or fourth of the mesoscutum. Males might also be mistaken for those of T. nevadensis; however, in T. concavus the metasomal sterna usually lack white setae, while in T. nevadensis there is an apical band of white setae on S3. Also, the clypeus of T. nevadensis is shining, with distinct larger punctures, while that of T. concavus is relatively matte, and covered with setae, thus partly obscuring the larger punctures. The mesoscutum is shinier in T. nevadensis (with punctures separated by up to one puncture diameter) and the scutellum is somewhat flattened and extended posteriorly; in T. concavus the mesoscutum is matte (with punctures nearly contiguous) and the scutellum is bigibbous, not extended posteriorly. Distribution.—USA: Alabama, Arizona, Arkansas, California (the Central Valley, and extending further south into San Bernardino Co. and Riverside Co.), Colorado, Florida, Georgia, Idaho, Illinois, Indiana, Kan- sas, Louisiana, Michigan, Minnesota, Mississippi, Missouri, New Mexico, North Carolina, Oklahoma, Texas, Utah, Washington, Washington D.C., Wisconsin. Host Records.—Svastra (Epimelissodes) obliqua (Say) (Custer 1928, observations of adults entering nests, larvae in cells; Bohart 1970, egg in cell; Rozen 1989b, larva from nest). Floral Records.—Helenium sp., Helianthus annuus L., H. tuberosus L., Heliopsis helianthoides (L.) Sweet, Lepandena marginata (= Euphorbia marginata Pursh), Prionopsis ciliata (= Grindelia papposa Nesom & Suh), Ratibida pinnata (Vent.) Barnh., Verbesina sp., Vernonia baldwinii Torr., “star thistle” (= Cen- taurea). Seasonal Records.—4 June to 21 October. Specimens examined.—193 &, 53 % (AUSTIN, BOULDER, CORVALLIS, DAVIS, GAINESVILLE, ITHACA, LAWRENCE, LOGAN, LOS ANGELES, NEW YORK, PHILADELPHIA, RIVERSIDE, STARKVILLE, TUCSON, URBANA, WASHINGTON D.C.).

TRIEPEOLUS CRESSONII (Robertson) (Figs. 39, 40, 248)

Epeolus cressonii Robertson 1897: 344 [Lectotype: Illinois Natural History Survey No. 3282a; &, Illinois, Macoupin Co., Carlinville; 15 September 1886; Coreopsis (=Bidens) aristosa]; Webb 1980: 108 [lectotype designation (by W. E. LaBerge)]. Triepeolus cressonii; Robertson 1901: 231. Triepeolus cressonii cressonii; Mitchell 1962: 466, Fig. 112 [redescription, floral records, illustrations of scutellum, axillae].

Description.—Length ca. 8–10.5 mm; ITW 1.6–2.3 mm. Integument black, often with at least partially red mandible, labrum, scape, pedicel, F1, tegula, and legs, and occasionally with red pronotal lobe and tip of axillar spine; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus sometimes elongate, entirely or partly asetose (female) or entirely covered with dense white setae (male), with moderate to strong midline, and weak larger punctures. Paramedian band clearly separated from other pale setae on mesoscutum in females; usually laterally contiguous with diffuse pale setae on anterior margin of mesoscutum in males. Scutellum moderately to strongly bigibbous; axillar spine triangular, reaching or slightly exceeding scutellar midpoint, apex sometimes apically incurved. Mesepisternum lacking erect, simple setae; with dorsal region of pale, branched setae often restricted to below scrobal groove (sometimes more extensive); lower pleuron asetose, with dense, small punctures (females and few males); or entirely covered with white, branched setae (most males). T1 with very wide, parallel-sided (i.e., BTB and ATB parallel), ovate or quadrate discal patch.

48 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER T2 with LLB forming weakly to strongly acute angle with ATB. Female: Pseudopygidial area subovate to subquadrate, with distinct basal crescent of shining setae; S5 not or very slightly downcurved apically. Meso- somal and metasomal venter black except for (often sparse) apical bands of pale setae on S2–S4. Male: Pygidial plate relatively narrow, keyhole shaped, with distinct basal transverse ridge; S2–S3 usually with uninterrupted bands of white setae; S4–S5 with apical fringes of brown setae (S4 often with white setae on basal margin of fringe). Comments.—This species very closely resembles T. helianthi and T. laticaudus. Females of T. cressonii and T. helianthi can be separated by the pseudopygidial area (with a distinct basal crescent of shining setae in T. cressonii, entire area nearly uniformly shining in T. helianthi), the mesepisternum (with dense pale setae more or less restricted to beneath the scrobal groove in T. cressonii, dorsally with more such pale setae in T. helianthi); and by the amount of pale setae laterally on the sterna (present on S2–S4 in T. cressonii, S3–S4 in T. helianthi). The males are extremely similar, but may be separated based on characters of the clypeus (usually entirely covered with white setae in T. cressonii, apically asetose in T. helianthi), paramedian bands (usually laterally contiguous with pale setae on the anterior margin of the mesoscutum in T. cressonii, usually distinct in T. helianthi), and white apical bands of S2–S3 (usually entire in T. cressonii, usually medially interrupted in T. helianthi). Females of T. cressonii can be differentiated from those of T. laticaudus by the mesepisternum (in T. laticaudus the dorsal third mostly covered by dense, pale setae, and medioventrally the integument between the punctures is distinctly raised, appearing tuberculate; while in T. cressonii the pale setae is more restricted dorsally, and the integument is much flatter between punctures medioventrally). Also, the basal shining setae of the pseudopygidial area are longer in T. laticaudus than in T. cressonii. Distribution.—USA: Alabama, Georgia, Illinois, Indiana, Kansas, Louisiana, Maryland, Minnesota, Mississippi, Missouri, Nebraska, New Jersey, North Carolina, North Dakota, Oklahoma, Pennsylvania, Ten- nessee, Texas, South Dakota, Wisconsin. Floral Records.—Aster paniculatus [= Symphyotrichum lanceolatum (Willd.) Nesom ssp. lanceolatum var. lanceolatum], A. pilosus [= Symphyotrichum pilosum (Willd.) Nesom var. pilosum], Bidens aristosa (Michx.) Britt., Brauneria pallida [= Echinacea pallida (Nutt.) Nutt.], Coreopsis tinctoria Nutt., Gaillardia pulchella Foug., Helianthus annuus L., H. atrorubens L., H. petiolaris Nutt., H. tuberosus L., Monarda sp., Nepeta cataria L., Physostegia parviflora Nutt. ex Gray, Prionopsis ciliata (= Grindelia papposa Nesom & Suh), Rudbeckia laciniata L., Silphium sp., Solidago altissima L., S. serotina (= S. gigantea Ait.), Vernonia baldwinii interior (Small) Faust, V. m is su r ic a Raf., V. noveboracensis (L.) Michx. Seasonal Records.—1 June to 14 October. Specimens examined.—154 &, 48 % (BERLIN, DAVIS, GAINESVILLE, ITHACA, LAWRENCE, LOS ANGELES, NEW YORK, PHILADELPHIA, STARKVILLE, UNIVERSITY PARK, URBANA, WASHINGTON D.C.).

TRIEPEOLUS CRUCIFORMIS new species (Figs. 41, 42)

Diagnosis.—This species can be distinguished by the combination of the plus-sign shaped T1 discal patch; the medially interrupted ATBs on T1–T4; the black or dark brown legs; the distinct basal region of silvery setae on the pseudopygidial area (from which it can be separated from the similar species T. warriti, whose pseudopygidial area is entirely dark brown or has a very poorly differentiated basal area of silvery setae); the relatively short clypeus; the absence of erect, simple setae on the mesepisternum; the white lateral setae on the female T5; and the strongly downturned apical margin of S5. Description of holotype female.—Length ca. 8 mm; ITW 1.6 mm. Integument black, with red on apical half of mandible; orange-brown on F1; dorsum of mesosoma and metasoma with bands of pale yellow setae.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 49 Clypeus lacking midline and larger punctures. Paramedian band clearly separated from other pale setae on mesoscutum. Scutellum weakly bigibbous; axillar spine triangular, not attaining midpoint of scutellum. Mesepisternum lacking erect, simple setae; only small area posterior to pronotal lobe with pale, branched setae (or entire dorsal fourth, excluding hypoepimeron, covered with such setae); ventrally with integument shining, with small punctures relatively evenly spaced ca. 0.5–1 puncture width apart. T1 discal patch strongly rectangular, forming plus-shaped sign with medially interrupted ATB and BTB; T2 with LLB absent. Pseudopygidial area longitudinally ovate, with basal area of silvery, pilose setae, these silvery setae not extended apically along lateral margin of pseudopygidial area, apically with relatively long, coarse setae; S5 conspicuously downcurved. (Male: Unknown.) Comments.—This species was labeled “PCAM 29” by T. Griswold. Distribution.—MéXICO: Puebla. Floral Records.—Viguiera dentata (Cav.) Spreng. Seasonal Records.—16 September to 3 November. Holotype.—“MÉXICO Puebla, 12 km SW Tehuacán, 1470 m, 3 Nov. [November] 1991, T. Griswold // Viguiera dentata // USDA-ARS Bee Biol. & Syst. Lab, Logan Utah, Faunal Survey No. 000 036 073 // female // Triepeolus sp. 29 & T. Griswold det 93 // Triepeolus PCAM-29 // Triepeolus PCAM 29 det. T. Griswold // SM032446 KUNHM-ENT // M. G. Rightmyer Triepeolus Database Specimen No. 1226 // HOLOTYPE & Triepeolus cruciformis Rightmyer 2007” (LAWRENCE). Paratype.—MÉXICO: PUEBLA: Zapotitlán, Salinas, 16 September 1995, Y. Cardel, 18°19’45”(N), 97°27’15”(W) (PUEBLA). Etymology.—This name is in reference to the cross-shaped region of dark setae on T1. From Latin “cru- cis” (of the cross).

TRIEPEOLUS CUABITENSIS Genaro (Figs. 43, 44)

Triepeolus cuabitensis Genaro 1999: 217–218, Figs 1b, 3d [Holotype: Museo Nacional de Historia Natural de Cuba; &, Cuabitas, Santiago de Cuba, Cuba; December 1948]. (paratype only viewed)

Description.—Length ca. 6–8 mm; ITW 1.4 mm. Integument black with dark reddish brown on mandible, labrum, apical clypeus, and venter of metasoma; orange on entire antenna, pronotal lobe, tegula, and legs including spurs but excluding basal portions of coxae; dorsum of mesosoma and metasoma with bands of yellow setae. Clypeus with absent to faint midline, lacking larger punctures. Paramedian band tapering anteriorly, reaching anterior margin of mesoscutum, barely laterally contiguous with narrow strip of pale setae on anterior margin of mesoscutum. Scutellum moderately bigibbous, with posterior surface slightly extended posteriorly, and covered by yellow setae; axillar spine strongly pointed, incurved, reaching posterior margin of scutellum. Mesepisternum with erect, simple setae; mostly covered by pale yellow, branched setae, but medially with circular region of dark brown, branched setae (this region reduced in male); with punctures nearly contiguous to separated by 0.5 puncture diameter. T1 discal patch transversely subovate; T2 with LLB forming weakly acute angle with ATB. Female: Pseudopygidial area entirely brown, vaguely distinct from rest of T5, apically with noticeably coarser, sparser setae; S5 very slightly downcurved apically. Male: Pygidial plate keyhole shaped, with distinct transverse basal ridge and apical downturned plate; S2–S3 with apical bands of white setae, S3 with white setae slightly surpassing apical margin; S4–S5 with apical fringes of setae pale golden brown on S4, brown on S5. Comments.—This species is known by its small size, relatively uniform setae of the pseudopygidial area, and the lack of a median notch in the posterior margin of the head. Distribution.—CUBA: Santiago de Cuba.

50 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Seasonal Records.—December (day unspecified). According to Genaro (1999), a specimen of this species was collected in October. Specimens examined.—1 &, 1 % (GAINESVILLE, LAWRENCE).

TRIEPEOLUS DACOTENSIS (Stevens) (Figs. 45, 46, 249)

Epeolus dacotensis Stevens 1919: 210 [Holotype: U. S. National Museum of Natural History No. 23160; &, Williston (Williams Co.), North Dakota; 8 August 1915; at clay bank]. Triepeolus dacotensis; Bohart 1970: Figs. 20, 22, 24, 27 [photographs of egg membrane, first instar, and mature larvae]; Torchio 1986: 588–596, Figs. 1–8 [biological data, description of embryogenesis and egg eclosion, illustrations of egg, embryo]; Torchio and Burdick 1988: 632 [comparison of egg with that of E. compactus]; Rozen 1989b: 15, 16, Figs. 26–28 [description, illustrations of first instar].

Description.—Length ca. 10–12 mm; ITW 2.3–2.7 mm. Integument black with dark red medially on mandible; dorsum of mesosoma and metasoma with bands of pale yellow to white setae. Clypeus with weak dorsal midline and faint larger punctures. Head and mesosoma with conspicuous erect, simple setae. Paramedian band diffuse, joined laterally to diffuse, pale, erect to suberect setae on anterior of mesoscutum. Scutellum weakly bigibbous; axillar spine rounded, not attaining midpoint of scutellum. Mesepisternum with dense, dark brown, erect, simple setae; dorsally with patch of white, branched setae beneath pronotal lobe; pronotal lobe also covered with white, branched setae. T1 almost entirely covered by white setae except for black semicircle mediobasally; rest of terga with apical bands uninterrupted; T2 with LLB absent. Venter of mesosoma and metasoma entirely black (both sexes). Female: Pseudopygidial area subrectangular, with apical, transverse rows of stout, relatively sparse setae, basally with fine black setae, poorly differentiated from rest of T5; S5 straight in profile. Male: Pygidial plate very longitudinally rectangular (appearing square when T5 overlaps base), with apical and lateral margins upturned into lamellate rim, apparently lacking transverse basal ridge; S3 with slight medial extension of dark, apical setae (in one specimen); S4–S5 with apical fringes of dark brown setae. Comments.—This is a distinctive species, similar only to T. mojavensis, from which it can be separated by the relative amounts and pattern of pale setae on the metasomal terga. Distribution.—CANADA: Alberta; USA: North Dakota, Utah. Host Records.—Anthophora neomexicana Cockerell? [=Anthophora (Melea) bomboides Kirby] (Stevens, 1919, unspecified observations of nests), Anthophora (Melea) occidentalis Cresson (Stevens, 1919, unspecified observations of nests; Bohart, 1970, egg and larvae in cells; Torchio, 1986, eggs and immatures from nest; Rozen, 1989b, larvae from nest). Floral Records.—Brauneria pallida [= Echinacea pallida (Nutt.) Nutt.], Helianthus petiolaris Nutt., Lactuca tatarica (L.) C.A. Mey. var. pulchella (Pursh) Breitung, Senecio sp. Seasonal Records.—15 April to 14 July. Specimens examined.—11 &, 4 % (AUSTIN, BOULDER, LAWRENCE, NEW YORK, WASHINGTON D.C.).

TRIEPEOLUS DENVERENSIS Cockerell (Figs. 47–50)

Triepeolus denverensis Cockerell 1910a: 91 [Holotype: U. S. National Museum of Natural History No. 100022; %, Den- ver, Colorado; 11 August 1908 (Peritoma serrulatum)].

Description.—Length ca. 8.5–14 mm; ITW 1.8–3.1 mm. Integument black, with red on legs excluding basal portions of coxae and spurs, and sometimes parts of femora (sometimes with red on part of mandible, labrum,

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 51 scape, pedicel, F1, and tegula); dorsum of mesosoma and metasoma with bands of pale yellow setae (some female specimens from Utah lacking pale banding). Clypeus lacking or with weak midline, lacking larger punctures, sometimes covered with sparse pale setae (dense in males). Paramedian band absent (black mor- phs), or present and clearly separated from other pale setae on mesoscutum, or contiguous with diffuse, pale setae on anterior margin of mesoscutum. Scutellum strongly bigibbous; axillar spine reaching or slightly surpassing midline of scutellum, apical point slightly incurved. Mesepisternum with sparse, erect, simple setae; with patch of pale, dense, branched setae beneath scrobal groove and pronotal lobe; ventrally with punctures small and nearly contiguous, mostly asetose (females) or covered with black, branched setae (males); black forms of female with mesepisternum entirely covered with black setae. T1 discal patch transversely rectangular to subovate; T2 with LLB absent or forming 90 degree angel with ATB. Female: Pseudopygidial area longitudinally elongate, strongly triangular, mostly silvery shining, but with small patch of coarse, darker setae subapically, this area of coarse, darker setae shorter than basal region of silvery setae; S5 not or very slightly downcurved, medially longitudinally convex. Male: Pygidial plate with distinct transverse basal ridge and apical downturned plate; S2–S3 with apical bands of white setae, S3 with setae slightly surpassing apical margin; S4–S5 with apical fringes of dark brown setae. Comments.—Triepeolus denverensis females are readily recognized by the triangular pseudopygidial plate, which is mostly silvery except for the small, subapical patch of dark, coarse setae. This pseudopygidial area is somewhat similar to that of T. isohedrus, and especially T. punctoclypeus. Triepeolus denverensis may be distinguished from T. isohedrus by the paramedian bands of pale setae on the mesoscutum (forming distinct anchor-shaped pattern in T. isohedrus), the LLB on T2 (lacking or forming 90 degree angle in T. denverensis, forming acute angle in T. isohedrus), and by the predominantly silver pseudopygidial area in T. denverensis, among other characters. Triepeolus denverensis may be differentiated from T. punctoclypeus by the distinctly coarser texture of the dark, submedian patch of setae on the pseudopygidial area in T. denverensis, by the denser punctation of the mesepisternum in T. denverensis, and by the distinct larger punctures of the clypeus in T. punctoclypeus. Some females of this species from Utah are noteworthy for the absence (or par- tial absence) of bands of pale setae, sometimes resulting in an entirely black appearance. Distribution.—USA: Arizona (Coconino Co., Navajo Co.), Colorado, Montana, New Mexico (McKin- ley Co.), North Dakota, Utah. Host Records.—Melissodes [=Svastra (Epimelissodes)] obliqua (Say)? and Melissodes (Eumelissodes) agilis Cresson? (Cockerell 1910a, collected at same flowers). Floral Records.—Chrysothamnus sp., Grindelia sp., Peritoma serrulatum (=Cleome serrulata Pursh), Senecio longilobus (=Senecio flaccidus Less. var. flaccidus). Seasonal Records.—17 August to 23 September. Specimens examined.—29 &, 2 % (BOULDER, DAVIS, LOGAN, NEW YORK, WASHINGTON D.C.).

TRIEPEOLUS DIFFUSUS new species (Figs. 51, 52)

Diagnosis.—This species can be distinguished by the combination of the sparse, pale setae on much of the mesoscutum and scutellum in many specimens; the clearly-present silvery setae basally on the pseudopygidial area, which smoothly transition to coarser, sparser setae apically; the acute angle formed by the LLB and ATB on T2; and the lack of erect, simple setae on the mesepisternum. Triepeolus diffusus strongly resembles both T. circumculus and T. vernus, from which it usually can be distinguished by the paramedian bands, which are separated from, or only faintly contiguous with, other pale setae in T. circumculus and T. vernus, while in T. diffusus the paramedian bands are at least laterally contiguous with pale setae on the anterior margin of the mesoscutum (and sometimes entirely surrounded by pale setae on mesoscutum). In addition, the scutellum of

52 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER T. vernus is weakly bigibbous, and the species is apparently found only in Texas during the months of March and April whereas T. diffusus is more widespread and appears later in the year. Triepeolus diffusus also can be differentiated from T. circumculus by the pseudopygidial area, in which the boundary between the basal and apical regions is strongly demarcated in T. circumculus, while in T. diffusus the two gradually intergrade. In addition, T. diffusus resembles T. dilutus, but in T. dilutus the T1 discal patch is more distinctly rectangular (almost forming plus-shaped sign). Triepeolus eldoradensis and T. fraserae also resemble T. diffusus, but these species have erect, simple setae on the mesepisternum. Description of holotype female.—Length ca. 10 mm (6–10 mm); ITW 1.9 mm (1.3–1.9 mm). Integu- ment black, grading to reddish brown on metasoma, with dark reddish orange on basal half of mandible, labrum (or part of labrum), apical margin of clypeus, pronotal lobe, and tegula; orange on scape, pedicel, F1, and legs excluding basal halves of coxae and spurs (sometimes dark brown on scape, pedicle, front leg, and axillar spine); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus shining, lacking (or with faint) midline, with faint larger punctures (densely covered with setae in male). Paramedian band poorly distinguished from diffuse setae on mesoscutum (or distinct but contiguous with diffuse lateral setae in specimens from southern locales). Scutellum strongly (to moderately) bigibbous, sparsely covered with pale setae (in most specimens); axillar spine triangular, not (or barely) attaining midpoint of scutellum. Mesepis- ternum apparently lacking (or with very sparse, short) erect, simple setae; dorsally with dense, pale yellow, branched setae, grading to sparser, pale yellow, branched setae ventrally; integument beneath very densely, minutely punctate (or entirely covered in dense, branched setae, mostly white except for small, brown, branched, ventroposterior spot in male). T1 discal patch subtriangular (to narrowly, transversely ovate); T2 with LLB forming strongly acute angle with ATB. Pseudopygidial area subquadrate (to longitudinally ovate), with silvery basal crescent of fine setae grading to coarser, golden apical setae; S5 weakly downcurved apically (to straight in profile). (Male: Pygidial plate somewhat keyhole shaped, only weakly emarginate laterally; S2–S3 with white apical bands of setae; S4–S5 with pale brown apical fringes, white laterally on S4, darker and shorter on S5.) Comments.—A specimen of T. diffusus was labeled “PCAM 21” by D. Yanega. Distribution.—MéXICO: Durango, Sonora; USA: Arizona, California (Riverside Co.), Colorado, Idaho, Nevada, New Mexico, Texas, Utah. Floral Records.—Asclepias sp., Bahia absinthifolia var. dealbata (Gray) Gray, Crepis sp., Dyssodia sp., Euphorbia sp., Helianthus petiolaris Nutt., Melilotus alba [= Melilotus officinalis (L.) Lam.], Pectis papposa Harvey & Gray, Sphaeralcea sp., Tamarix gallica L. Seasonal Records.—6 June to 6 August (more setose specimens, from Colorado, Idaho, Nevada, and Utah), or 13 August to 14 October (less setose specimens, from remaining locales). Holotype.—“UT [Utah], Millard Co., Fillmore, 12 mi NW, VI [June]-20-72 [1972], F. Parker, D. Vincent // on Tamarix gallica // Native Bee Survey USDA, Logan, Utah BBSL 582487 // M. G. Rightmyer Triepeolus Database Specimen No. 5602 // HOLOTYPE & Triepeolus diffusus Rightmyer 2007” (LOGAN). Paratypes.—USA: UTAH: Millard Co., Delta, 19 June 1950, G. F. Knowlton (1%, LOGAN); 24 June 1949, G. E. Bohart (4&, LOGAN); 25 June 1949, G. F. Knowlton (3&, LOGAN); 28 June 1950, C. D. Michener, Asclepias (2&, LAWRENCE); 30 June 1947, G. E. Bohart (1&, LOGAN); 15 July 1970, G. E. Bohart (1&, LOGAN); 16 July 1949, G. F. Knowlton (1&, LOGAN); 12 mi NW Fillmore, 20 June 1972, F. Parker, D. Vin- cent, Tamarix gallica (17&, LOGAN); Tamarix sp. (5&, LOGAN); Flowell, 22 June 1961, G. E. Bohart (4&, LOGAN); Fort Deseret, 17 June 1953, G. E. Bohart, E. A. Cross (1&, LOGAN); Salt Lake Co., 6600 W & 2000 N St., 8 June 1956, J. L. Eastin (1&, LOGAN). Additional specimens.—MÉXICO: DURANGO: 62 km NE Ceballos, Est. Biol. Mapimí, 22 August 1991, J. L. Neff (1&, LAWRENCE); Reserva Biósfera Mapimí near Ceballos, 22 August 1991, J. G. Rozen (1&, LAWRENCE); SONORA: Navajo, 27 September 1966, G. E. & A. S. Bohart (1&, LOGAN); USA: ARIZONA: Cochise Co., 1 mi E Douglas, 16 August 1962, M. Statham (2&, NEW YORK); 18 August 1962, M. Statham

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 53 (1&, NEW YORK); 19 August 1962, M. A. Cazier (1&, DAVIS); 20 August 1968, Rozen, Favreau (2&, NEW YORK); 1 mi N Douglas, 16 August 1962, E. G. Linsley, Bahia absinthifolia var. dealbata (3&, DAVIS); 25 mi NE Douglas, 26 August 1969, W. J. Hanson, T. Hsiao (2&, LOGAN); 3 mi E Portal, 13 August 1965, M. E. Irwin (1&, RIVERSIDE); San Simon, 13 August 1981, R. M. Bohart (1&, DAVIS); 5 mi S San Simon, 23 August 1996, J. G. & B. L. Rozen, A. Pence (1&, NEW YORK); CALIFORNIA: Riverside Co., 1 mi E Ford Dry Lake Road at Hwy I-10, 14 October 2000, G. R. Ballmer, Pectis papposa (4&, RIVERSIDE); COLORADO: (Delta Co.), Delta: 25 June 1938, U. Lanham (1&, BOULDER); 26 June 1938, R. Bauer (2&, BOULDER); U. Lanham (1&, BOULDER); Moffat Co., Artesia, 23 July 1950, C. D. Michener, Helianthus petiolaris (1&, LAWRENCE); IDAHO: Bonneville Co., 23 mi W Idaho Falls, 17 July 1975, G. E. Bohart (1&, LOGAN); 21 July 1975, G. E. Bohart (1&, LOGAN); Cassia Co.: Burley, 6 July 1931, J. Nottingham (1&, LAWRENCE); Raft R. Narrows, 8 July 1978, G. F. Knowlton (2&, LOGAN); Franklin Co., Preston, 10 June 1968, G. E. Bohart (1&, LOGAN); Oneida Co., Curlew Reservoir, 30 June 1969, G. F. Knowlton (1&, LOGAN); Stone Reservoir, 18 June 1969, G. F. Knowlton (1&, LOGAN); NEVADA: Lincoln Co., Panaca, 21 July 1958, R. C. Bechtel, Melilotus alba, (1&, GAINESVILLE); NEW MEXICO: (Hidalgo Co.), 4 mi S Animas, 24 August 1974, Rozen, Favreau (1&, NEW YORK); 5 mi W Cotton City, August 1959, R. M. Bohart (1&, DAVIS); 9 mi N Cot- ton City, 24 August 1989, R. M. Bohart (1&, DAVIS); Rodeo, 9 September 1973, M. Favreau (3&, NEW YORK); TEXAS: (Hidalgo Co.), Hidalgo, 18 July 1954, Univ. Kans. Expedition (1&, LAWRENCE); (Webb Co.), Laredo, July (1&, WASHINGTON D.C.); UTAH: Box Elder Co., Hardup, 18 June 1969, G. F. Knowlton (1&, LOGAN); Kelton Pass, 9 June 1969, G. F. Knowlton (1%, LOGAN); 5 mi W Snowville, 6 June 1969, G. F. Knowlton (1&, LOGAN); Snowville, 16 June 1969, G. F. Knowlton (1%, LOGAN); (Davis Co.), Davis, 23 June 1943, G. F. Knowlton, P. F. Telford (1&, LOGAN); Emery Co., 15 mi NW Woodside, 24 June 1963, A. S. Menke (1&, DAVIS); Tooele Co., Vernon, 19 June 1939, G. F. Knowlton, F. C. Harmston (1&, LOGAN); Wasatch Mts., 18 June 1952, E. I. Schlinger (1&, LOGAN); Uintah Co., Bonanza: 27 June 1978, G. Bohart, Helianthus (1&, BOULDER); 11 July 1974, G. E. Bohart, Melilotus alba (1&, LOGAN); 17 mi S Bonanza, 25 July 1975, Menke, Pulawski (1&, WASHINGTON D.C.); (Weber Co.), Ogden, 6 August 1958, G. E. Bohart (1&, LOGAN); (unknown county): 12 mi E Jenson, 23 June 1939, Timberlake, Sphaeralcea sp. (1&, WASHING- TON D.C.). Etymology.—This name is in reference to the paramedian bands of pale setae on the mesoscutum, which are connected laterally to pale setae on the apical margin of the mesoscutum. From the Latin, “diffusus” (spread out, extended, dispersed).

TRIEPEOLUS DILUTUS new species (Figs. 53, 54)

Diagnosis.—This species can be distinguished by the combination of the relatively short clypeus, the lack of erect, simple setae on the mesepisternum, the red legs, strongly rectangular T1 discal patch, the medially continuous ATBs on T3 and T4, the ovate pseudopygidial area with its well-differentiated basal region of silvery setae, and the strongly-downturned female S5. Triepeolus dilutus resembles T. diffusus, but in T. dilutus the T1 discal patch is more distinctly rectangular (almost forming plus-shaped sign). Triepeolus dilutus also resembles T. warriti, but can be differentiated from that species by the strongly downturned S5 (straighter in profile in T. warriti) and by the relatively strongly differentiated apical and basal setae on the pseudopygidial area (very poorly differentiated in T. warriti). Description of holotype female.—Length ca. 7.5 mm (7–8 mm); ITW 1.8 mm (1.5–1.8 mm). Integu- ment black, with dark reddish-orange on basal half of mandible, labrum, apical margin of (or entire) clypeus; orange on scape, pedicel, F1, pronotal lobe (in most specimens), tegula, and legs excluding basal halves of coxae and spurs; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint

54 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER midline and larger punctures. Paramedian band continuous with lateral setae (or sometimes clearly separated from other pale setae on mesoscutum). Scutellum weakly bigibbous; axillar spine triangular, not attaining midpoint of scutellum. Mesepisternum lacking erect, simple setae; dorsally with dense, pale yellow, branched setae, grading ventrally along margins to sparser setae; medially asetose (or with sparse, brown, branched setae); integument shining, punctures small and fairly regularly spaced ca. 0.5 puncture diameter apart. T1 discal patch widely subovate (to rectangular), weakly suggesting plus-shaped sign; T2 with LLB mostly on lateral surface of T2, forming weakly acute angle with ATB. Pseudopygidial area small, subcircular, with basal silvery setae somewhat pilose, with apical setae golden, relatively long; S5 downcurved. (Male: Unknown.) Comments.—This species was labeled “PCAM 22” by D. Yanega. Distribution.—MéXICO: Chihuahua, Durango; USA: Texas. Floral Records.—Coreopsis sp., Dyssodia sp., Haplopappus gracilis [= Machaeranthera gracilis (Nutt.) Shinners]. Seasonal Records.—9 April to 28 September. Holotype.—“Brownwood Tex. [Texas] VI [June].12-1924 // R. M. Fouts Collector // M. G. Rightmyer Triepeolus Database Specimen No. 6943 // HOLOTYPE & Triepeolus dilutus Rightmyer 2007” (WASHING- TON D.C.). Paratypes.—MÉXICO: CHIHUAHUA: 16 km N Jiménez, 16 August 1991, R. L. Minckley, Dyssodia (2&, LAWRENCE); DURANGO: Mapimí Reserva Biósfera, Rancho La Flor, 26°34’8”N 103°57’37”W, 28 September 1994, L. Godinez, 1205 m. (1&, MEXICO CITY); Reserva Biósfera Mapimí near Ceballos, 22 August 1991, J. G. Rozen (1&, LAWRENCE); 62 km NE Ceballos, Est. Biol. Mapimí, 22 August 1991, J. L. Neff (1&, LAWRENCE); USA: TEXAS: Brewster Co., Big Bend National Park, Tenapin Tanks, 15 July 1950, R. F. Smith (1&, NEW YORK); Brooks Co., 10 mi S Falfurrias, 9 April 1954, R. E. Beer et al., Coreopsis (1&, LAWRENCE); Brown Co., Brownwood, 12 June 1924, R. M. Fouts (1&, WASHINGTON D.C.); 10–15 June 1924, R. M. Fouts (1&, LOS ANGELES); Dimmit Co., Chaparral Wildlife Management Area, 7–8 June 1992, A. W. Hook (1&, AUSTIN); 13–15 June 1992, A. Hook, J. Neff (1&, AUSTIN); 12 mi E Catarina, Chaparral Wildlife Management Area, 13 June 1992, J. L. Neff, A. Hook (1&, AUSTIN); Jeff Davis Co., upper Limpia Canyon, Davis Mountains, 31 August 1986, R. R. Snelling, 5600 ft., Haplopappus gracilis (1&, LOS ANGE- LES); Kimble Co., 18–19 mi WNW Doss, 19 May 2001, A. Hook, J. Neff (1&, AUSTIN); Stonewall Co., 16 April 1953, R. H. Beamer (1&, LAWRENCE); Travis Co., Austin, 1 June 1985, J. Neff (1&, AUSTIN); Val Verde Co., Devils River, Dolan Falls, 26–29 May 1994, A. Hook, O. Hernandez (4&, AUSTIN); 28 May 1994, C. R. Nelson (1&, AUSTIN). Etymology.—This name is in reference to the more weakly differentiated setal types on the pseudopygidial area, in comparison to those of T. warriti. From the Latin, “dilutus” (mixed, weak, thin).

TRIEPEOLUS DISTINCTUS (Cresson) (Figs. 55–57, 244, 250, 251)

Epeolus distinctus Cresson 1878: 84 [Lectotype: Academy of Natural Sciences No. 2236; &, Georgia]; Cresson 1916: 117 [lectotype designation]. Epeolus bardus Cresson 1878: 84–85 [Lectotype: Academy of Natural Sciences No. 2237; &, Texas]; Cresson 1916: 113 [lectotype designation]. new synonymy Triepeolus bardus; Cockerell 1903: 331. Triepeolus mesillae var. a. Cockerell 1903: 331 [&, Las Cruces (Dona Ana Co.), New Mexico; September 22; Verbesina encelioides]. (not seen) Triepeolus mesillae Cockerell 1904: 36–37 [Holotype: U. S. National Museum of Natural History No. 9704; &, Mesilla, (Dona Ana Co., New Mexico); 24 September]; Rozen 1966: 16, 17, Figs. 19–23 [description, illustrations of postdefecating larva]. new synonymy

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 55 Triepeolus pimarum Cockerell 1904: 36–37 [Holotype: U. S. National Museum of Natural History No. 9703; &, Alham- bra, (Maricopa Co.), Arizona; Verbesina encelioides]. new synonymy Triepeolus distinctus; Mitchell 1962: 467, Fig. 112 [redescription, floral records, illustrations of scutellum, axillae]; Wuellner and Hixon 1999: 145–147 [behavior before leaving host nest, chemical components of Dufour’s glands, venom glands, and glandular pouches]. Triepeolus distincta; Minckley et al. 1994: 1415.

Description.—Length ca. 10–12.5 mm; ITW 2.1–2.9 mm. Integument black, with red to orange usually entirely or partly on the following: mandible, labrum, clypeus (especially apically), antenna (especially basally), pronotal lobe, tegula, mesoscutum, scutellum, axilla, mesepisternum, legs, pygidial plate of both sexes; dorsum of mesosoma and metasoma with bands of yellow setae. Clypeus lacking or with weak midline; covered with golden setae. Head somewhat globular, with preoccipital carina on posterior margin of head behind vertex, as well as on gena. Paramedian band narrow, clearly separated from other pale setae on mesoscutum. Scutellum moderately bigibbous; axillar spine well surpassing midpoint of scutellum, incurved, pointed. Mesepisternum with sparse, short, erect, simple setae; irregularly punctate, with punctures relatively large, separated up to one puncture diameter in some places, with integument between punctures raised, shining. T1 discal patch small, diamond-shaped or rectangular; T2 with LLB absent or rarely weakly present, forming 90 degree angle with ATB. Female: Pseudopygidial area rectangular, with apical margin straight or weakly concave; setae uniformly dense, coarse, short, and golden-shining; apical margin usually silvery-shining; S5 straight in profile, strongly convex along longitudinal midline, with bristle-like setae present on apical margin; S2–S4 lacking apical bands of white setae or with sparse pale setae on apical margins. Male: Pygid- ial plate wide, with slightly upturned (lamellate) margins; medioapically often notched; lacking distinct basal transverse ridge; S2–S3 with white to pale golden setae on apical margins (often slightly extended sublaterally on S3); S4–S5 with apical fringes of golden brown setae (often white basally on apical fringe of S4). Comments.—This species is known by the unique pseudopygidial area (see Fig. 56), pattern of pale setae on T1, and the long axillar spines. Distribution.—USA: Arizona, Colorado, Georgia, Illinois, Iowa, Kansas, Minnesota, Nebraska, New Mexico, Texas, Utah. Host Records.—Dieunomia (Dieunomia) heteropoda (Say) (36 specimens from nesting sites, in Cochise Co., Arizona and Bastrop Co., Texas; Wcislo, 1993, adults observed entering nests, prepupae and pupa recov- ered from nests), Dieunomia (Epinomia) triangulifera (Vachal) (1 specimen entering nest in Lawrence, Kan- sas; 3 specimens from nesting site in Lawrence, Kansas; Rozen 1966, larvae presumably taken from nest; Wcislo et al., 1994, adults caught emerging from nests; Wuellner and Hixon, 1999, adults entering nests). Floral Records.—Euphorbia dentata Michx., Helianthus annuus L., H. petiolaris Nutt., Heterotheca grandiflora Nutt., He. latifolia [= He. subaxillaris (Lam.) Britt. & Rusby], Monarda punctata L., Solidago speciosa Nutt., Verbena stricta Vent., Verbesina encelioides (Cav.) Benth. & Hook. f. ex Gray, V. helianthoides Michx. Seasonal Records.—10 May to 22 October. Specimens examined.—271 &, 195 % (AUSTIN, BERKELEY, BOULDER, ITHACA, LAWRENCE, LOS ANGE- LES, NEW YORK, URBANA, TERRE HAUTE, WASHINGTON D.C.).

TRIEPEOLUS DIVERSIPES Cockerell (Figs. 59, 60)

Triepeolus diversipes Cockerell (in Cockerell and Sandhouse 1924): 314 [Holotype: California Academy of Sciences No. 1608; %, San Pedro (Los Angeles Co.), California; 25 October 1909].

Description.—Length ca. 10–13 mm; ITW 2.2–3.2 mm. Integument black, with red on apical half of mandible; orange on F1 and middle and hind legs excluding basal portions of coxae and spurs (sometimes also on

56 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER tegula and front leg); dorsum of mesosoma and metasoma with bands of pale yellow to yellow setae. Clypeus with faint midline and larger punctures; covered with dense white setae in males. Paramedian band clearly separated from other pale setae on mesoscutum or contiguous with lateral setae (anterior third to entire mesoscutum sometimes covered with diffuse pale yellow setae). Scutellum strongly to moderately bigibbous; axillar spine triangular, reaching scutellum midpoint, apically somewhat curved inward. Mesepisternum apparently lacking or with extremely sparse, short, erect, simple setae; dorsal third to fourth (except hypoepimeron) covered with dense, pale yellow, branched setae, sometimes with pale setae extending posteroventrally (females), or entire mesepisternum covered with dense, branched, white setae (males); with punctures small, nearly contiguous; integument between somewhat raised, rough. T1 discal patch transversely ovate, rectangular, or triangular; T2 with LLB forming acute to weakly angle with ATB. Female: Pseudopy- gidial area triangular, mostly covered with very pilose, coarse, golden setae, but basally with fine, dense, golden setae; S5 straight in profile. Male: Pygidial plate keyhole shaped, with distinct transverse basal ridge and apical downturned plate; S2–S3 with apical bands of white setae; S4–S5 with apical fringes of brown to white setae (if brown, then S4 with white setae apicolaterally). Comments.—The pseudopygidial area of this species somewhat resembles that of T. parvidiversipes; however the two species can easily be distinguished by the mesepisternum (with much longer erect, simple setae and much more widely-spaced punctures in T. parvidiversipes), in addition to the much smaller size of T. parvidiversipes. A specimen of this species was labeled with a manuscript name indicating a “false pygidium,” while three other specimens were given a manuscript name referring to the ornate nature of the pseudopygidial area. Distribution.—USA: California (San Bernardino Co., Inyo Co.), Colorado, New Mexico, Oregon (Lake Co.), Utah. Floral Records.—Chrysothamnus sp., “Helianthoid composite”. Seasonal Records.—24 August to 29 October. Specimens examined.—19 &, 2 % (BOULDER, BERKELEY, DAVIS, LOGAN, SAN FRANCISCO, WASHING- TON D.C.).

TRIEPEOLUS DONATUS (Smith) (Figs. 58, 61, 62, 252)

Epeolus donatus Smith 1854: 256 [Holotype: The Natural History Museum, London No. 17B.519; &, (Mount Pleasant, Jefferson Co., Ohio)]. Triepeolus donatus; Robertson 1901: 231; Mitchell 1962: 467–468 [redescription]. Triepeolus cirsianus Mitchell 1962: 463–464 [Holotype: Purdue Entomological Research Collections; %, Warren Co., Indiana, 11 August 1953; thistle]. new synonymy Triepeolus crisianus; Commonwealth Institute of Entomology 1962: 312 [lapsus calami].

Description.—Length ca. 10–13 mm; ITW 2.5–2.7 (rarely as small as 2.0) mm. Integument black (sometimes with purplish tint), with red on distal half of mandible (specimens from Iowa, Minnesota, and North Dakota with red legs); dorsum of mesosoma and metasoma with bands of pale gray to white setae. Face elongate; clypeus with strong to moderate midline present and weak larger punctures, asetose or covered with white setae. Paramedian band clearly separated from other pale setae on mesoscutum (females and some males), or connected laterally to diffuse white setae on anterior margin on mesoscutum (most males). Scutel- lum moderately bigibbous; axillar spine triangular, reaching midpoint of scutellum. Mesepisternum with relatively sparse, long, erect, simple setae; punctures small, nearly contiguous to separated by 1–2 puncture widths in some places, especially ventrally; integument between flat or slightly raised (punctures generally denser and integument between sometimes slightly tuberculate in males, especially those from Midwestern states). T1 discal patch transversely ovate (sometimes subrectangular), with ATB interrupted medially; T2

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 57 with LLB reduced or forming very weakly acute, nearly 90 degree, angle with ATB (mostly on lateral surface of T2). Female: Pseudopygidial area subovate to subquadrate, with distinct basal shining crescent; S3–S4 with apical bands of pale setae, sometimes restricted laterally (Southeastern specimens usually with pale bands only S4, these bands sometimes reduced to a few setae on apicolateral margins); S5 straight in profile. Male: Pygidial plate of moderate size, with strong to weak basal transverse ridge; S2–S3 with apical bands of white setae; S4–S5 with apical fringes of setae pale brown intermixed with white (S4), or brown medially and white laterally (S5). Comments.—This species is similar to T. texanus in size, the elongate face, and erect, simple setae on mesepisternum; however, in T. donatus the bands of setae are white (rather than pale yellow), the T1 discal patch is usually more ovate than rectangular, and the pseudopygidial area’s basal shining crescent is less strongly differentiated from the more apical, coarser setae. Triepeolus donatus is also similar to T. georgicus and T. atripes, especially in size and coloration, but unlike the latter two species, T. donatus has an elongate face and erect, simple setae on the mesepisternum. Triepeolus rugosus is similar in coloration, the elongate face, and erect, simple setae, but is generally smaller and has an irregularly punctate mesepisternum, with the integument between the punctures distinctly raised and somewhat tuberculate. Distribution.—USA: Connecticut, Delaware, Florida, Illinois, Indiana, Iowa, Maine, Maryland, Massa- chusetts, Michigan, Minnesota, Missouri, New Hampshire, New Jersey, New York, North Dakota, Ohio, Pennsylvania, Vermont, West Virginia, Wisconsin. Host Records.—Eutechnia (sic) (=Melitoma) taurea (Say)? (Robertson 1901:231, adult entering nest); Melissodes (Heliomelissodes) desponsa Smith (John S. Ascher, in lit. 2004, unpublished data from observations and specimens in New York state). The Melitoma host record is for T. donatus entering the nest of Mel- itoma taurea, observed by W. H. Ashmead, who incorrectly concluded that T. donatus was the builder of the nest (Robertson, 1899). Floral Records.—Carduus undulatus [= Cirsium undulatum (Nutt.) Spreng. var. undulatum], C. altissi- mus [= Cirsium altissimum (L.) Hill], Centaurea jacea L., C. vulgare (Savi) Ten., Inula helenium L., Monarda fistulosa L., Rudbeckia laciniata L., Silphium perfoliatum L. Seasonal Records.—26 July to 8 October. Specimens examined.—33 &, 40 % (ANN ARBOR, BERKELEY, BOULDER, CORVALLIS, DAVIS, ITHACA, LAWRENCE, LOGAN, LONDON, LOS ANGELES, NEW YORK, SAN FRANCISCO, TUCSON, URBANA, WASHINGTON D.C., WEST LAFAYETTE).

TRIEPEOLUS EDWARDI new species (Figs. 63–65, 253)

Diagnosis.—This species primarily can be distinguished by the unusual form of the pseudopygidial area, which is almost entirely dark brown. Coupled with the fact that the T5 usually lacks white setae laterally on the tergum, the pseudopygidial area is mostly discernable by the change in angle in T5 created by the flat plane of the pseudopygidial area. In dorsal aspect, this species strongly resembles T. antiochensis, but the two species can be differentiated by the female T5 (the pseudopygidial area of T. antiochensis has a distinctly differentiated basal region of silvery setae, as well as a large patch of white setae lateral to the pseudopygidial area). If the T5 does have white setae present laterally, then T. edwardi can be recognized by the erect, simple setae on the mesepisternum, the 90 degree angle formed by the LLB and ATB of T2, the relatively short clypeus, and the rectangular to reduced T1 discal patch. Description of holotype female.—Length ca. 11 mm (10.5–11.5 mm); ITW 2.3 mm (2.1–2.4 mm). Integument black, with red on apical half of mandible, and reddish orange on outer F1 (also sometimes on legs excluding coxae and spurs); dorsum of mesosoma and metasoma with bands of very pale yellow setae.

58 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Clypeus lacking midline, with distinct larger punctures; face entirely dark except with small areas of white setae near anterior tentorial pits and antennal bases (or entirely covered with dense, white setae in males). Paramedian band relatively long, mostly separated from other pale setae on mesoscutum but with few, scattered setae medially (entirely isolated in some females; or nearly indistinct from diffuse, pale setae on anterior to entire mesoscutum in males). Scutellum weakly (to moderately) bigibbous; axillar spine triangular, not quite reaching (sometimes reaching) midpoint of scutellum. Mesepisternum with erect, simple setae; dorsally with small patch appressed, pale yellow, branched setae between pronotal lobe and hypoepimeron (or entire upper third or more covered with appressed, white, branched setae in males); ventrally with sparse, black, branched setae; punctures rough, nearly contiguous to separated by nearly a puncture diameter in some areas, with integument between raised and somewhat shining. T1 discal patch partially covered with sparser, pale setae to form very small, circular area (or slightly larger triangular area, or with T1 discal patch appearing as short, but relatively wide, rectangle); T2 with LLB reduced, forming 90 degree angle with ATB; ATBs of all terga robust, not medially interrupted. Pseudopygidial area poorly differentiated from rest of T5, represented by a flattened, dark, semicircular plane, with very vague basal crescent; T5 with lateral pale setae reduced (or absent); mesosoma and metasomal venter entirely black; S5 very faintly downcurved. (Male: Pygidial plate keyhole shaped, with poorly differentiated transverse basal ridge and weakly downturned apical plate; S2–S3 with apical bands of white setae restricted to lateral margins; S4–S5 with apical fringes of black setae.) Comments.—Various specimens of this species bear Timberlake manuscript names meaning “beautiful” and “from Siskiyou.” Distribution.—USA: California (Lassen Co., Modoc Co., Siskiyou Co.), Oregon (Klamath Co.). Floral Records.—Chrysothamnus nauseosus [= Ericameria nauseosa (Pallas ex Pursh) Nesom & Baird]. Seasonal Records.—21 August to 22 September. Holotype.—“Gazelle Cal [California], Siskiyou Co., IX [September] .22.1951 // Jack Hall, E. I. Schlinger Collectors //Native Bee Survey USDA Logan, Utah BBSL522457 // M. G. Rightmyer Triepeolus Database Specimen No. 6967 // HOLOTYPE & Triepeolus edwardi Rightmyer 2007” (LOGAN). Paratypes.—USA: CALIFORNIA: Lassen Co., 2 mi E Herlong, 12 September 1983, J. A. Powell (1&, BERKELEY); Standish, 15 September 1983, J. Chemsak, Chrysothamnus sp. (1&, BERKELEY); Modoc Co., 1 mi N Cedarville, 14 November 1969, E. E. Grissell, R. F. Denno (1&, DAVIS); Siskiyou Co., Gazelle, 11 Sep- tember 1950, A. T. McClay (2&, 1%, LOGAN); Lava Beds National Monument, 22 August 1954, G. Ferguson (1%, LOGAN); Lower Klamath Lake, 21 August 1954, G. Ferguson (1%, LOGAN), 3 September 1959, J. Schuh (1&, CORVALLIS); Orr Lake, 3 September 1963, J. Schuh, Chrysothamnus sp. (1&, GAINESVILLE); Tule Lake, 22 August 1954, G. Ferguson (1%, LOGAN); OREGON: Klamath Co., 7 September 1954, J. Schuh (1&, LOGAN), 3 mi W Merrill, 27 August 1970, Penrose, Westcott, Chrysothamnus nauseosus (1&, LOGAN). Etymology.—This species is named in honor of my uncle, Edward G. Marquette, who gave me a microscope when I was young, introducing me to whole worlds I had never before noticed.

TRIEPEOLUS ELDORADENSIS (COCKERELL) (Figs. 66, 67, 254)

Epeolus eldoradensis Cockerell 1910b: 245–246 [Holotype: U. S. National Museum of Natural History No. 100021; %, Eldora (Boulder Co.), Colorado; 18–19 August; Grindelia (subalpina)]. Epeolus eldoradensis var. a Cockerell 1910b: 246. Triepeolus eldoradensis; Hurd 1979: 2092.

Description.—Length ca. 7–10 mm; ITW 1.5–2.3 mm. Integument black, usually with red on apical half of mandible and margins of labrum; orange on F1, tegula, and legs excluding basal portions of coxae and spurs (sometimes on scape and pedicel); dorsum of mesosoma and metasoma with bands of pale to very pale yellow

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 59 setae. Clypeus lacking or with very faint midline and larger punctures (sometimes not visible due to white setae). Paramedian band poorly distinguished from diffuse pale setae covering mesoscutum. Scutellum weakly bigibbous; axillar spine almost reaching midpoint of scutellum. Mesepisternum with dense, erect, simple setae (sometimes rather short); entirely covered with appressed, pale yellow to white, branched setae (slightly less dense ventrally in females). T1 discal patch widely rectangular to ovate, often obscured by sparser pale setae; T2 with LLB forming 90 degree angle with ATB. Female: Pseudopygidial area semicircular to subquadrate, with distinct, relatively long, basal region of silvery setae and larger apical region of coarse, darker setae; S5 very slightly downcurved. Male: Pygidial plate keyhole shaped, with distinct transverse basal ridge and apical downturned plate; S2–S3 with apical bands of white setae, S3 with white setae slightly surpassing apical margin; S4–S5 with apical fringes of setae white on S4, golden brown on S5. Comments.—This species is similar to T. argyreus and T. paenepectoralis; see Table 1 for differentiating characters among these species. Triepeolus eldoradensis also resembles T. diffusus, but T. eldoradensis has erect, simple setae on the mesepisternum, the LLB on T2 forms a 90 degree angle with the ATB, and the pseudopygidial area has a distinct basal shining crescent. Distribution.—USA: Colorado, Idaho, Utah, Wyoming. Floral Records.—Aster sp. (= Symphyotrichum), Grindelia squarrosa (Pursh) Dunal, G. subalpina Greene. Seasonal Records.—20 July to 17 October. Specimens examined.—28 &, 1 % (LAWRENCE, LOGAN, URBANA, WASHINGTON D.C.).

TRIEPEOLUS ENGELI new species (Figs. 68, 69)

Diagnosis.—This species is readily recognized by the female pseudopygidial area, due to the unique medioapical tuft of long, golden setae. The scutellum of the paratype approaches that of T. bimorulus, but the axilla is not fringed with pale setae in T. engeli as it is in that species. Description of holotype female.—Length ca. 10 mm; ITW 2.0 mm. Integument black, with dark reddish orange on most of mandible (basal half on paratype), labrum, apical half of clypeus, scape, pedicel, F1, and legs excluding coxae and spurs; pale brown on tegula; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint midline, stronger dorsally, and larger punctures. Paramedian band nearly laterally contiguous with pale setae curving medially towards paramedian band on anterior of mesoscutum. Scutellum weakly bigibbous; axillar spine triangular, not attaining midpoint of scutellum. Mesepisternum lacking erect, simple setae; with dense, pale yellow, branched setae on all margins, enclosing circular patch of short, brown, branched setae medioventrally; integument with punctures nearly contiguous to separated by 0.5 puncture diameter. T1 discal patch transversely ovate, with BTB notched medially and ATB medially interrupted; T2 with LLB forming acute angle with ATB. Pseudopygidial area transversely ovate, covered with fine, brown setae, except medioapically with tuft of long, golden setae, these setae slightly upturned and slightly shorter medially; S5 straight in profile. (Male: Unknown.) Comments.—The holotype specimen has greatly reduced mandibles, while the paratype specimen has normal length mandibles. In all other observable respects the two specimens are very similar. Distribution.—USA: Texas (Maverick Co., Lee Co.). Floral Records.—Dalea lasiathera Gray. Seasonal Records.—11 April to 1 June. Holotype.—“9 mi SW Eagle Pass, Tex. [Texas], IV [April]-11-1959 // Michener, Rozens, Beamers, Stephen // Taken on Dalea lasianthera // M. G. Rightmyer Triepeolus Database Specimen No. 1306 // HOLO- TYPE & Triepeolus engeli Rightmyer 2007” (LAWRENCE).

60 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Paratype.—USA: TEXAS: (Lee Co.), Fedor, 1 June 1896, Birkmann (1&, BERLIN). Etymology.—This species is named in honor of Michael S. Engel, for his inspiring and generous encour- agement in my transition from scientific illustrator to melittologist.

TRIEPEOLUS EPEOLURUS Rightmyer (Figs. 70, 71)

Triepeolus epeolurus Rightmyer 2004a: 25–28 [Holotype: University of Kansas Natural History Museum and Biodiver- sity Research Center No. 9162; &, Los Sabinos, Michoacán, México; 1190 m; 29 October 1987].

Description.—Length ca. 8.5–10.5 mm; ITW 1.6–2.1 mm. Integument black to brown, with red on apical half of mandible, and sometimes on scape, pedicel, and F1; dorsum of mesosoma and metasoma with bands of yellow to pale yellow setae (males with bands of setae on metasomal terga grading from yellow to white posteriorly). Labrum with apical tubercles often produced into scoop-like structure. Clypeus lacking or with weak dorsal midline; lacking larger punctures, sometimes covered with sparse, white setae (usually denser in males). Paramedian band relatively short, clearly separated from other pale setae on mesoscutum. Scutellum weakly to moderately bigibbous; axillar spine triangular, reaching or not attaining midpoint of scutellum. Mesepisternum lacking erect, simple setae; with regions of dense, white, branched setae beneath scrobal groove and below pronotal lobe (sometimes sparse on hypoepimeron); ventral half of lateral surface of mesepisternum with punctures nearly contiguous, sometimes covered with sparse, brown, branched setae (usually denser and paler in males). T1 discal patch triangular to ovate; T2 lacking LLB. Metasomal terga with apical bands interrupted medially (at least T1–T3). Female: Pseudopygidial area with distinct transverse, basal, region of silvery setae, apically with dark, long, stout setae; S5 straight in profile. Male: Pygid- ial plate keyhole shaped, with distinct transverse basal ridge and downturned apical plate; S2–S3 with white apical bands of setae, S3 with white setae extending slightly past apical margin; S4–S5 with relatively weak apical fringes of setae, brown and white on S4, brown on S5. Comments.—Females of this species are easily recognized by the unique pseudopygidial area, which has a subapical transverse band of silvery, fine, appressed setae contrasted with an apical band of long, stout, suberect setae. Distribution.—COSTA RICA: Alajuela, Guanacaste; MéXICO: Jalisco, Michoacán, Oaxaca. Seasonal Records.—22 September to 5 December. Specimens examined.—24 &, 9 % (BERKELEY, CHAMELA, GAINESVILLE, LAWRENCE, LOGAN).

TRIEPEOLUS EXILICURVUS new species (Figs. 72, 73)

Diagnosis.—This species primarily can be distinguished by the combination of the weakly downturned S5 and the pseudopygidial area, which has a longitudinally elongate basal area of silvery setae that extends apicolaterally to nearly enclose a subapical circular region of dark, coarse, relatively sparse setae. In addition, T. exilicurvus is known by the relatively short clypeus, the brown scape, the paramedian band, which is laterally contiguous with pale setae subapically on mesoscutum, the absence of erect, simple setae on the mesepisternum, and the medially-interrupted ATBs on T1 and T2 (those of T1 interrupted by nearly 1 OD). Triepeolus exilicurvus strongly resembles T. argus and T. circumculus due to the form of the pseudopygidial area. It is differentiated from T. circumculus by its much larger size and by the more elongated basal region of silvery setae on the pseudopygidial area in T. exilicurvus. It is differentiated from T. argus by the weakly downcurved S5 and by the pseudopygidial area, in which the apicomedial, circular region of coarse setae is

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 61 nearly fully enclosed by silvery setae in T. exilicurvus. The form of the pseudopygidial area found in these species also resembles those found in T. micropygius (from which T. exilicurvus can be differentiated by the only weakly downcurved S5), and T. atripes (from which T. exilicurvus can be differentiated by the bands of pale yellow setae on the dorsal metasoma and red legs). Description of holotype female.—Length ca. 11.5 mm; ITW 2.6 mm. Integument black, with red on apical half of mandible; orange on most of protibia and distal on foreleg, and middle and hind legs distal to trochanters excluding spurs; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with distinct midline and larger punctures. Paramedian band laterally contiguous with pale setae subapically on mesoscutum. Scutellum strongly bigibbous; axillar spine triangular, not attaining midpoint of scutellum. Mesepisternum lacking erect, simple setae; dorsal half covered with dense, pale, branched setae, sparser on hypoepimeron; ventrally with pale, branched setae on margins; medioventrally mostly asetose, with punctures nearly separated by 0.5 puncture diameter or less; integument between punctures relatively flat. T1 discal patch trapezoidal; T2 with LLB forming acute angle with ATB. Pseudopygidial area longitudinally ovate, with dense, fine, silvery setae on basal half and on lateral margins, nearly enclosing distinct, apicomedial, circular region of sparse, coarse setae; S5 weakly downcurved apically. (Male: Unknown.) Distribution.—MéXICO: Coahuila. Seasonal Records.—20 October. Holotype.—“MÉXICO Coahuila, 39 km S Agua Nueva, 20-X [October]-1994, Alt. 1770 m, 24°53’21”N 101°04’63”O, Col. E. Ramirez // M. G. Rightmyer Triepeolus Database Specimen No. 1302 // HOLOTYPE & Triepeolus exilicurvus Rightmyer 2007” (CHAMELA). Etymology.—This name is in reference to the weakly downcurved S5 of the female. From the Latin, “exilis” (weak) and “curvus” (bent).

TRIEPEOLUS FLAVIGRADUS new species (Figs. 74, 75)

Diagnosis.—This species is known by the recessed appearance of the flat clypeus, due to the long, erect setae that surround it. Triepeolus flavigradus shares this facial appearance with T. robustus, but can be differentiated from that species by the coloration of the transverse bands of pale setae on the metasomal terga (grading posteriorly from yellow to white in T. flavigradus; uniformly pale yellow in T. robustus), as well as by the general shape of the T1 discal patch (quadrate to subtriangular in T. flavigradus; strongly rectangular in T. robustus) and the LLB of T2 (forming a right angle with the ATB in T. robustus, usually absent in T. flavigradus). This species also strongly resembles T. nayaritensis, but in T. flavigradus the paramedian band is absent or poorly distinguished from other pale setae on the mesoscutum (present and isolated in T. nayaritensis) and the clypeus appears recessed (face lacking such erect setae surrounding the clypeus in T. nayaritensis). Description of holotype female.—Length ca. 13 mm (10–15.5 mm); ITW 2.7 mm (2.1–2.8 mm). Integ- ument black to brown, with red medially on mandible; brownish-orange on outer F1; dorsal metasoma with bands of setae pale yellow on T1, grading to white posteriorly on terga. Clypeus shining, with very faint midline (not visible in some specimens), with distinct larger punctures (or covered with appressed white setae in male), very flat, appearing recessed due to erect setae on upper face. Mesoscutum shining; paramedian band interspersed with diffuse white setae on anterior mesoscutum (or absent). Scutellum very weakly bigibbous, weakly extended posteriorly; axillar spine triangular, rounded apically, not attaining midpoint of scutellum. Mesepisternum with long, erect, simple setae; with small patch dense, white, appressed, branched setae below scrobal groove and along posterior margin of mesepisternum to top of mesocoxa (or more densely covered with appressed, white, branched setae, usually sparser medially to give impression of rounded, asetose or very sparsely, darkly setose, black region in male); punctures very small, separated by 2 or 3 puncture widths (or a

62 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER puncture width or less in male); integument shining. T1 quadrate, nearly subtriangular (usually quadrate, rarely subovate or subtriangular); ATB and BTB interrupted medially, basal interruption continued along anterior surface of T1 forming rectangular dark area; T2 with LLB absent (or mostly on lateral surface of T2, forming very weakly acute angle with ATB). Pseudopygidial area similar to that of T. concavus, but area wider and setae less erect; S5 strongly downcurved, apical margin surpassing apical margin of T5. (Male: Pygidial plate relatively wide, with transverse basal ridge often hidden beneath long setae and relatively weak downturned apical plate; S2 with apical band of white setae; S3 with white apical setae surpassing apical margin of sternum, slightly curled apically; S4–S5 with white to golden brown apical fringes.) Distribution.—COSTA RICA: Guanacaste, Puntarenas; MéXICO: Chiapas, Jalisco, Michoacán, Nayarit, Oaxaca, Puebla, Veracruz. Floral Records.—Baltimora recta L., Eysenhardtia polystachya [= Eysenhardtia orthocarpa (Gray) S. Wats.], Sida sp., Asteraceae. Seasonal Records.—16 June to 2 December. Holotype.—“17 km W Tehuantepec, Oaxaca, México, 8 Sept [September] 1965// D. H. Janzen Collector // M. G. Rightmyer Triepeolus Database No. 6677 // HOLOTYPE & Triepeolus flavigradus Rightmyer 2007” (WASHINGTON D.C.). Paratypes.—MÉXICO: OAXACA: 55 mi NW Tehuantepec, 31 August 1957, H. A. Scullen, 3050 ft. (1&, CORVALLIS); 3 mi W El Cameron, 6 August 1963, F. D. Parker, L. A. Stange (1&, RIVERSIDE); Nopala, 50 km N Puerto Escondido, 2 September 1990, L. M. Godinez, 700 m, Sida sp. (1%, LAWRENCE); 30.6 km S Pinotepa Nacional, 23 October 1989, F. A. Noguera, A. Rodriguez (1%, CHAMELA); 10 mi SE Tapanatepec, 8 August 1963, F.. D. Parker, L. A. Stange (1%, RIVERSIDE); PUEBLA: 3 km SE Zapotitlán, near Los Reyes, Metzontla, 10 September 1996, R. Brooks, 1680 m, tall yellow comp. (Asteraceae) (1&, LAWRENCE); 14 km SE Acatlán, 19 October 1989, A. Rodriguez, F. A. Noguera (1%, CHAMELA); 20 km NW Acatlán, 1 November 1991, T. Griswold, 1200 m (1&, LAWRENCE). Additional specimens.—COSTA RICA: GUANACASTE: 4 mi NW Cañas, La Pacifica, 23 July 1972, P. A. Opler, Baltimora recta (1&, BERKELEY); 4 km N Cañas, 15 July 1965, D. H. Janzen (4%, LAWRENCE); 14 km S Cañas, 23 July–6 August 1991, F. D. Parker (1%, LOGAN); 4 mi E Cañas, 1 July 1974 (4%, DAVIS); 3 km NW Nacaome, P. N. Barra Honda, 3–25 Aug 1992, M. Reyes, 100 m (1&, HEREDIA); 5–27 July 1992 (2&, HEREDIA); 8 km SW Cuajiniquil, Est. Murciélago, F. Quesada, 16 June–4 July 1993, 100 m (1&, 1%, HEREDIA); C. Cano (1&, 6%, HEREDIA); Estación Experimental Horizontes, June 1993, P. Rios, 100–150 m. (1%, HEREDIA); Sugar Beach, 3 km N Potrero, 9–11 November 1991, A. S. Menke (3%, WASHINGTON D.C.); F. D. Parker (1%, LOGAN); MÉXICO: JALISCO: 12 km S El Tuito, 6 November 1987, T. Griswold (2&, LOGAN); 8 km N El Tuito, 2 December 1988, J. Chemsak (1&, BERKELEY); Chamela, 7 November 1986, R. Ayala (1&, CHAMELA); Chamela (Est. Biológica), 17 October 1988, Buickerood, E. S. Ross, 175 m (1&, SAN FRANCISCO); 15 October 1988 (1%, SAN FRANCISCO); 14 October 1988 (1%, SAN FRANCISCO); San Juan Lagos, 27 July 1951, H. E. Evans, flowers of Eysenhardtia polystachya (Ort.) (1%, LAWRENCE); MICHO- ACÁN: Los Sabinos, 28 km S Ario de Rosales, 29 October 1987, R. Ayala, 1190 m (1&, CHAMELA); T. Gris- wold (1&, 1%, LOGAN); 20 km NE La Huacana, 3 November 1992, A. Rodriguez, F. A. Noguera (1&, CHAMELA); NAYARIT: 29 km N Peñitas, mi. 75157.7, 28 September 1976, C. D. George, R. R. Snelling, 90 m (2&, 7%, WASHINGTON D.C.); VERACRUZ: Cotaxtla, Cotaxtla Exp. Sta., 15 August 1962, D. H. Janzen (2&, BERKELEY); 5 mi NE Tinajas, 18 August 1963, F. D. Parker, L. A. Stange Co. (2&, 1%, DAVIS). Etymology.—This name is in reference to the coloration of the dorsal metasomal banding, which grades from yellow to white posteriorly. From the Latin, “flavus” (yellow) and “gradus” (degree, pitch).

Triepeolus cressoni var. fraserae Cockerell 1904: 39 [Holotype: U. S. National Museum of Natural History No. 9708; %, Beulah (San Miguel Co.), New Mexico; 29 June; Frasera]. Triepeolus loganensis Cockerell 1925a: 624 [Holotype: U. S. National Museum of Natural History No. 100032; %, Logan Co., Colorado; 23 August 1923]. new synonymy Triepeolus sandhousae Cockerell 1925a: 624–625 [Holotype: U. S. National Museum of Natural History No. 100036; %, Logan Co., Colorado; 23 August 1923]. new synonymy Triepeolus fraseri; Hurd 1979: 2093.

Description.—Length ca. 7–12 mm; ITW 1.2–2.2 mm. Integument black, with red on most of mandible; orange on F1 and legs excluding basal portions of coxae and spurs (often with brown patches on at least foreleg), sometimes on part or entire labrum, scape, pedicel, and tegula; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with distinct (sometimes weak) midline and weak larger punctures, integument shining or less frequently covered with sparse pale yellow setae (often entirely covered by dense white setae in males). Paramedian band clearly separated from other pale setae or laterally contiguous with pale setae near anterior margin of mesoscutum. Scutellum moderately bigibbous, sometimes with longitudinal line of pale yellow setae between biconvexities; axillar spine triangular, reaching midpoint of scutellum, often slightly incurved apically. Mesepisternum with sparse, erect, simple setae; dorsal half with dense, pale yellow, branched, appressed setae; medioventrally and hypoepimeron with pale yellow, branched setae sparser or with sparse brown, branched setae; punctures small, nearly contiguous to separated by up to two puncture diameters in some places, integument between punctures shining, somewhat elevated (females), or entire mesepisternum covered with dense, pale yellow, branched setae (sparser on hypoepimeron; males). T1 discal patch transversely ovate; T2 with LLB forming acute angle with ATB. Female: Pseudopygidial area semicircular, with distinct basal crescent of silvery setae; S5 weakly to moderately downcurved. Male: Pygidial plate keyhole shaped, with distinct basal transverse ridge and apical downturned plate; S2–S3 with strong, broad white apical bands strongly contrasting with dense brown setae basally (S3 with white setae slightly surpassing apical margin); S4–S5 with apical fringes of brown or brown and white setae. Comments.—This species is similar to T. diffusus, T. vernus, and T. dilutus, but can be separated from those species by the erect, simple setae on the mesepisternum of T. fraserae. It is also similar to T. micropygius; however, S5 and T5 of T. micropygius are conspicuously narrowed distally, S5 is much more strongly downcurved, the pseudopygidial area is even more distinctly rounded, and the erect, simple setae of the mesepisternum are much shorter in T. micropygius (0.25 OD vs. greater than 0.5 OD in T. fraserae). Distribution.—USA: Arizona, California (Riverside Co.), Colorado, Montana, New Mexico, North Dakota, Utah, Wyoming. Floral Records.—Frasera sp., Grindelia squarrosa (Pursh) Dunal, Gutierrezia sp., Helianthus annuus L., Isocoma tenuisecta Greene. Seasonal Records.—21 June to 2 November. Specimens examined.—55 &, 6 % (BOULDER, DAVIS, LAWRENCE, LOGAN, LOS ANGELES, NEW YORK, RIVERSIDE, TEMPE, WASHINGTON D.C.).

TRIEPEOLUS FULGIDUS new species (Figs. 78, 79)

Diagnosis.—This species can be distinguished by the pseudopygidial area, in which the setae are nearly uniformly pale brown or golden, and the apical margin is straight. This type of pseudopygidial area is similar to that of T. joliae, but the T1 discal patch of T. fulgidus lacks the median circular region of dark setae that is

64 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER found in T. joliae. Triepeolus fulgidus is similar to T. phaeopygus, but in T. fulgidus the paramedian band is isolated from other pale setae on the mesoscutum, and the setae of the pseudopygidial area are more appressed than those of T. phaeopygus. Triepeolus fulgidus is also similar to T. medusa in the sparse, erect setae of the mesepisternum (these setae are difficult to see in many specimens of T. fulgidus, and denser in T. medusa), the setae and punctures of the clypeus and mesepisternum, and the isolated paramedian bands, but the setae of the pseudopygidial area are more uniformly pale golden in T. fulgidus than in T. medusa. Description of holotype female.—Length ca. 10.5 mm (10.5–11.5 mm); ITW 1.9 mm (1.9–2.2 mm). Integument black, with brownish orange on basal half of mandible, labrum, and tegula (also on apical margin of clypeus, scape, pedicel, and medially on pronotal lobe in paratype); orange on F1 and legs excluding basal halves of coxae and spurs; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus distinctly shining, with weak midline and distinct larger punctures. Paramedian band clearly separated from other pale setae on mesoscutum. Scutellum weakly bigibbous; axillar spine triangular, reaching midpoint of scutellum. Mesepisternum with sparse, erect, simple setae (very short, and therefore possibly overlooked); dorsal third covered with dense, pale, branched setae, absent on hypoepimeron, extending down anterior surface (setae on anterior surface brown in paratype); ventrally mostly asetose (or with sparse, brown, branched setae in paratype); punctures nearly contiguous to separated by 0.5 puncture diameter; ventral mesosoma covered with white (or brown in paratype), branched setae. T1 discal patch transversely ovate; T2 with LLB forming acute angle with ATB. Pseudopygidial area subquadrate, with setae nearly uniform and golden, but setae on basal margin of pseudopygidial area very slightly finer and denser than rest of setae; S2–S4 with distinct lateral or continuous bands of white setae (also vaguely present on apical margin of S1); S5 straight in profile. (Male: Unknown.) Distribution.—USA: Arizona (Navajo Co.), New Mexico (Sandoval Co.). Seasonal Records.—8 August to 13 August. Holotype.—“ARIZ [Arizona] Navaho Co., Show-Low, VIII [August]-8-71 [1971], Bill Apperson // Native Bee Survey USDA, Logan, Utah, BBSL 581422 // M. G. Rightmyer Triepeolus Database Specimen No. 1297 // HOLOTYPE & Triepeolus fulgidus Rightmyer 2007” (LOGAN). Paratype.—USA: NEW MEXICO: (Bernalillo Co. or Sandoval Co.), Sandia Mountains, 13 August 1935, 5000 ft. (1&, DAVIS). Etymology.—This name is in reference to the pale golden setae of the pseudopygidial area. From the Latin, “fulgidus” (shining, gleaming).

TRIEPEOLUS GEORGICUS Mitchell (Figs. 80, 81)

Triepeolus georgicus Mitchell 1962: 469–470, Fig.112 [Holotype: U. S. National Museum of Natural History No. 400192; &, Fort Gordon, Richmond Co., Georgia; 24 September 1958]. Triepeolus floridanus Mitchell 1962: 468–469 [Holotype: Florida State Collection of Arthropods; %, Gainesville (Ala- chua Co.), Florida; 28 October 1956]. new synonymy

Description.—Length ca. 11 mm; ITW 2.2–2.3 mm. Integument black, with red on distal half of mandible and sometimes on F1; dorsum of mesosoma and metasoma with bands of pale gray to white setae. Tegula more or less transparent on apical margin. Clypeus with weak midline, lacking or with weak larger punctures, nearly or entirely asetose. Paramedian band connected laterally to diffuse white setae on anterior margin of mesoscutum, or barely distinct in some females. Scutellum moderately to strongly bigibbous; axillar spine reaching or slightly surpassing scutellar midpoint, weakly incurved apically. Mesepisternum lacking erect, simple setae; punctures spaced up to two puncture diameters apart, integument between flat; with dense, white, branched setae mostly restricted to dorsal half or fourth. T1 discal patch transversely ovate; T2 with

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 65 LLB of pale setae forming weakly acute angle with ATB (mostly on lateral margin of T2). Female: Pseudopygidial area ovate, with distinct basal shining setae; S2–S4 with white setae on apical, or only apicolateral, margins (sometimes faint on S2); S5 weakly downcurved apically. Male: Pygidial plate moderately wide, weakly keyhole shaped, with distinct basal transverse ridge; S2–S3 with white setae apically; S4–S5 with brown apical fringes (sometimes with small patch white setae on apicolateral margin of S4). Comments.—This species is very similar to T. donatus, but T. georgicus lacks erect, simple setae on mesepisternum and the clypeus is not elongate. Triepeolus georgicus is also similar to T. atripes, except in T. georgicus the clypeus is more convex in profile, with a modest midline (as opposed to a relatively flattened clypeus with a strong midline), the integument of the ventral half of the lateral surface of the mesepisternum is relatively flat, with punctures separated by up to a puncture diameter (as opposed to almost contiguous, with the integument between the punctures raised, slightly tuberculate in appearance), and in females, the paramedian bands reach the anterior margin of the mesoscutum, the pseudopygidial area has a basal crescent that is shorter than that of T. atripes, and S5 is slightly less downcurved. Triepeolus georgicus is less likely to be confused with T. cressonii, but can be distinguished from that species by the pale gray to white band coloration and black integument except for the mandibles and F1 (as opposed to pale yellow band coloration, usually with at least some areas of red integument on the legs and face in T. cressonii). As in T. obliteratus, the number of submarginal cells in this species appears to be labile: there is a tiny “fourth” submarginal cell in one wing of the holotype of T. floridanus. Distribution.—USA: Florida, Georgia, Mississippi. Seasonal Records.—24 September to 28 October. Specimens examined.—6 &, 3 % (GAINESVILLE, STARKVILLE, WASHINGTON D.C.).

TRIEPEOLUS GRANDIS (Friese) (Figs. 82, 83, 241–243, 255)

Epeolus grandis Friese 1917 [1916]: 338 [Holotype: Zoologisches Museum, Humboldt-Universität; %, Jacubaya (=Tacu- baya?, see Comments below), México]. Triepeolus species A; Rozen 1966: 12–14, Figs. 8–14 [description, illustrations of postdefecating larva]. Triepeolus species b; Rozen 1984: 7, 11, 18 Figs. 6, 23 [potential defense by host, biology, illustrations of postdefecating larva and first instar on host provisions]. Triepeolus grandi; Ayala 1988: 404. [lapsus calami]. Triepeolus grandis; Rozen 1989a: 2–10, Figs. 1–6, 10–17 [redescription, biology, illustrations of adult, pupa, egg]; Rozen 1989b: 1–14, Figs. 1–18 [descriptions, illustrations of first and second instars].

Description.—Length ca. 11–15 mm; ITW 2.9–3.6 mm. Integument black, with dark reddish brown to orange on basal mandible and F1, usually on labrum, scape, and pedicel, sometimes on apical margin of clypeus, rarely on pronotal lobe, almost always orange on tegula, usually on legs (excluding basal coxae and spurs, sometimes with black areas on tibiae, profemur, and trochanters; or legs rarely entirely black); dorsum of mesosoma and metasoma with bands of pale yellow to yellow setae. Clypeus with weak to strong midline and larger punctures, entirely covered with white setae in males. Paramedian band usually clearly separated from other pale setae on mesoscutum, robust (rarely barely contiguous with lateral setae). Scutellum strongly to moderately bigibbous; axillar spine triangular, reaching or well surpassing midpoint of scutellum, apical spine somewhat incurved in those specimens with longer spine. Mesepisternum with erect, simple setae; dorsal half with dense, pale yellow, branched setae (absent or sparser on hypoepimeron), sometimes extending along anterior and posterior margins; ventrally with sparser, black, branched setae; punctures small, nearly contiguous to separated by a puncture diameter, with integument between punctures raised, tuberculate (females); or mesepisternum covered with dense, white, branched setae, except for small medioventral patch of dense, brown, branched setae (males). T1 discal patch trapezoidal, sometimes subtriangular or triangular;

66 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER T2 with LLB forming acute to weakly acute angle with ATB. Female: Pseudopygidial area ovate, with uniformly fine, darkly shining setae; S5 straight in profile. Male: Pygidial plate relatively wide, with distinct transverse basal ridge and apical downturned plate; S2 with apical band of white setae; S3–S5 with apical fringes of setae, white and shorter on S3, golden on S4–S5. Comments.—Triepeolus grandis is very similar to T. zacatecus; presumably the latter species is the black-legged form of T. grandis from the west coast of México referred to by Rozen (1989a: 8). In fact, T. grandis is only separable from T. zacatecus (usually) by the orange legs, paler yellow bands of setae, narrower bands of yellow setae on T1 and T2, and shorter axillar spines. Triepeolus grandis is superficially similar to T. remigatus, but T. grandis has erect, simple setae on the mesepisternum, the paramedian bands are usually clearly separated from other pale setae on the mesoscutum, and the T1 discal patch is trapezoidal to subtriangular. The original description of Epeolus grandis is based on one male specimen; thus it is the holotype specimen by default. Ayala (1999: 24) states that Jacubaya is possibly a locality in Veracruz; however, I cannot find “Jacubaya” in any atlas available to me. Alternatively, Tacubaya is a possible correct spelling of the type locality. There is a locality called Tacubaya in each of the following Mexican states: Chihuahua, Guanajuato, Oaxaca, and Yucatán, as well as in the Distrito Federal. A specimen of this species was labeled “PCAM 2” by D. Yanega; another specimen was given a manuscript name by Paul Hurd honoring Mont A. Cazier. Triepeolus grandis is found in abundance in the Southwestern United States. This species is parasitic on the predominately crepuscular or nocturnal subfamily Diphaglossinae; one of its known host species, Ptilo- glossa arizonensis, was recorded flying as early as 5 am in Portal, Arizona by Rozen (1984). Distribution.—MéXICO: Baja California, Chihuahua, Distrito Federal, Durango, Jalisco, Querétaro, Sonora, Zacatecas; USA: Arizona, Kansas, New Mexico, Oklahoma, Texas. Host Records.—Caupolicana (Caupolicana) yarrowi (Cresson) (James H, Cane, in lit., 2005, visiting nest); Ptiloglossa arizonensis Timberlake (Rozen, 1984, eggs and larvae from cells; Rozen, 1989b larvae from cells), Ptiloglossa jonesi Timberlake (Rozen, 1966, larvae from cells; Rozen, 1984, larvae taken from cells; Rozen, 1989b, larva from cell). Floral Records.—Aloysia gratissima (Gillies & Hook.) Troncoso, A. wrightii (Gray) A. Heller ex Abrams, Asclepias subverticillata (A. Gray) Vail, Baccharis sp., Bahia absinthifolia var. dealbata (Gray) Gray, Baileya multiradiata Harv., Chilopsis linearis (Cav.) Sweet, Eriogonum deflexum Torr., Gaillardia pulchella Foug., Gutierrezia sp., Helianthus petiolaris Nutt., Heliotropium sp., Heterotheca subaxillaris (Lam.) Britt. & Rusby, Hoffmanseggia jamesii [= Pomaria jamesii (Torr. & Gray) Walp.], Kallstroemia grandiflora Torr. ex Gray, Larrea tridentata (Sessé & Moc. ex DC.) Coville, Marrubium vulgare L., Melilotus officinalis (L.) Lam., Verbesina encelioides (Cav.) Benth. & Hook. f. ex Gray. Seasonal Records.—3 March to 14 October. Specimens examined.—195 &, 3 % (AUSTIN, BOULDER, CHAMELA, CORVALLIS, DAVIS, GAINESVILLE, ITHACA, LAWRENCE, LOGAN, LOS ANGELES, MEXICO CITY, NEW YORK, RIVERSIDE, SAN FRANCISCO, TEMPE, WASHINGTON D.C.).

TRIEPEOLUS GRISWOLDI new species (Figs. 84, 85)

Diagnosis.—This species can be separated from all others in the genus by the combination of the dense, golden, erect setae covering the vertex and mesoscutum; the strong, outcurved preoccipital carina on the gena; the paramedian band, which is joined laterally to pale yellow setae on anterior third to fourth of the mesoscutum; the mesepisternum, which has plentiful long, erect, simple setae mixed with pale, appressed, plumose setae; and the very wide, rectangular T1 discal patch (nearly reaching lateral margin of tergum in dorsal view).

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 67 Description of holotype female.—Length ca. 10.5 mm (10–10.5 mm); ITW 2.6 mm (2.2–2.6 mm). Integument black, with red on basal half of mandible, margins of labrum, and apically on clypeus; orange on F1, parts of scape, pedicel, pronotal lobe (in two of the three specimens), and legs excluding basal halves coxae and spurs; brownish orange on tegula; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint midline and larger punctures (sometimes covered by setae). Paramedian band somewhat diffuse, joined laterally to pale setae on anterior of mesoscutum. Scutellum moderately (to weakly) bigibbous; axillar spine triangular, slightly flattened, reaching midpoint of scutellum. Mesepisternum with long, erect, simple setae; mostly covered by pale, branched, yellow subappressed setae but with shining, somewhat tuberculate, black integument visible beneath setae ventrally. T1 discal patch widely, distinctly rectangular; T2 with LLB setae forming 90 degree angle with ATB. Pseudopygidial area semicircular to subquadrate, with vaguely differentiated basal crescent of golden shining setae; S5 slightly downcurved apically. (Male: Unknown.) Distribution.—USA: New Mexico (McKinley Co.), Utah (Garfield Co.). Floral Records.—Chrysothamnus sp., Senecio longilobus (= Senecio flaccidus Less. var. flaccidus). Seasonal Records.—29 August to 21 September. Holotype.—“USA Utah, Garfield Co. Calf Creek, 29 Aug [August] 1985 // T. L. Griswold Collector // on Chrysothamnus // Native Bee Survey USDA, Logan, Utah BBSL 224700 // M. G. Rightmyer Triepeolus Data- base Specimen No. 3564 // HOLOTYPE & Triepeolus griswoldi Rightmyer 2007” (LOGAN). Paratypes.—USA: TEXAS (sic, should be NEW MEXICO): McKinley Co., Tohatchi, 21 August 1965, G. E. Bohart, Senecio longi. (1&, LOGAN); UTAH: Garfield Co., Calf Creek, 29 August 1985 (1&, LOGAN). Etymology.—This species is named in honor of Terry L. Griswold, for his gracious hospitality and support during my studies of Triepeolus.

TRIEPEOLUS HELIANTHI (Robertson) (Figs. 86, 87)

Epeolus helianthi Robertson 1897: 344 [Lectotype: Illinois Natural History Survey No. 9496; &, Carlinville, Macoupin Co., Illinois; 18 September 1890; Helianthus grosseserratus]; Webb 1980: 108 [lectotype designation (by W. E. LaBerge)] Triepeolus helianthi; Robertson 1901: 231; Graenicher 1905: 164–166, Fig. 7 [description, illustrations of larva]; Parker et al. 1981: 48, 51, Figs. 9–14 [description, photographs of prepupae and egg chorion, behavior of adults]. Triepeolus helianthi var. arizonensis Cockerell 1904: 39 [Holotype: U. S. National Museum of Natural History No. 9707; %, Phoenix, Arizona; 9 October; Helianthus annuus]. new synonymy Triepeolus coquilletti Cockerell 1905c: 106 [Holotype: U. S. National Museum of Natural History No. 9908; &, San Ber- nardino Co., California; October]. new synonymy Triepeolus helianthi helianthi; Cockerell 1919b: 300 Triepeolus helianthi pacificus Cockerell 1919b: 300 [Holotype: U. S. National Museum of Natural History No. 100031; %, Peaceful Valley (Boulder Co.), Colorado; 26 August]. new synonymy Triepeolus maculiventris Cockerell 1921: 11–12 [Holotype: American Museum of Natural History No. 25095; &, (Navajo Canyon), Mesa Verde (National Park), Colorado; about 37°11’N, 108°30’W; 6600 ft.; (5) July 1919 (Helianthus petiolaris)]. new synonymy Triepeolus lineatulus Cockerell and Sandhouse 1924: 306–307 [Holotype: California Academy of Sciences No. 1598; &, Stockton (San Joaquin Co.), California; 20 August 1919]. new synonymy

Description.—Length ca. 8.5–12 mm; ITW 1.7–2.7 mm. Integument black, with the following sometimes at least partly reddish: mandibles, labrum, clypeus, scape, pedicel, F1, pronotal lobe, tegula, legs; dorsum of mesosoma and metasoma with bands of yellow to pale yellow setae. Clypeus slightly elongate, asetose (in females) with strong to moderate midline (male clypeus basally covered with white setae). Paramedian band clearly separated from other pale setae on mesoscutum, or rarely laterally contiguous with diffuse pale setae

68 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER on anterior margin of mesoscutum in some males. Scutellum moderately to strongly bigibbous; axillar spine triangular, reaching or exceeding scutellar midpoint, often apically pointed and slightly incurved. Mesepister- num lacking erect, simple setae (sometimes appearing to have very short, sparse, erect, simple setae in males), females with distinct, “L” shaped dorsal region of pale, branched setae (pale setae absent on hypoepimeron), lower pleuron black, asetose with very dense, small punctures (separated by up to one puncture diameter in some places); male mesepisternum more generally covered with white, branched setae, often with brown patch medioventrally. T1 with very wide, parallel-sided (i.e., BTB and ATB parallel), laterally rounded, ovate discal patch. T2 with LLB forming weakly acute angle with ATB. Female: Pseudopygidial area subovate, with very vague, almost indistinguishable basal crescent (formed by increase in density, but not texture, of setae), appearing brownish (rather than silvery) throughout; S5 very slightly downcurved. Mesosomal and metasomal venter black except for slight lateral patches of white setae on S3–S4 (sometimes also on S2). Male: Pygidial plate strongly keyhole shaped, with distinct basal transverse ridge; S2–S3 with white apical bands, often restricted laterally, sometimes with white setae apicolaterally on S4; S4–S5 with apical fringes of brown setae. Comments.—This species is very similar to T. cressonii. Females can be separated by the pseudopygidial area, which is a more elongate oval, with a distinct basal crescent of shining setae in T. cressonii, while in T. helianthi the pseudopygidial area is more circular, and the setae of the entire area are nearly uniform in reflectance; the mesepisternum in T. cressonii has dense pale setae more or less restricted to beneath the scrobal groove (sometimes these pale setae are more extensive), while in T. helianthi there are more such pale setae dorsally on the mesepisternum, forming an “L”-shaped pattern; and in T. cressonii there are pale setae laterally on S2–S4, while in T. helianthi the pale setae are sometimes only on S3–S4. The males are extremely similar, but may be separated based on the following characters: the clypeus tends to be entirely covered with white setae in T. cressonii, while in T. helianthi the clypeus is usually apically asetose; the paramedian bands are usually laterally contiguous with pale setae on the anterior margin of the mesoscutum in T. cressonii, while in T. helianthi the paramedian bands are usually distinct; and S2–S3 usually have white apical bands entire in T. cressonii, while in T. helianthi the white apical bands often medially interrupted. Distribution.—CANADA: Alberta; MéXICO: Chihuahua, Coahuila, Nuevo León; USA: Arizona, Cali- fornia (ranging from Modoc Co. to Orange Co. and San Bernardino Co.), Colorado, Connecticut, Illinois, Indiana, Kansas, Maryland, Minnesota, Mississippi, Missouri, Montana, Nebraska, Nevada, New Jersey, New Mexico, New York, North Carolina, North Dakota, Oregon, South Dakota, Texas, Utah, Virginia, Washington, Wisconsin, Wyoming. Host Records.—Dieunomia (Dieunomia) heteropoda (Say)? (1 specimen from nest site), Nomia (Acuno- mia) melanderi Cockerell? (1 specimen from nest site in Utah); Melissodes (Eumelissodes) agilis Cresson (Parker et al. 1981, adult entering nest, prepupae from cells), Melissodes (Callimelissodes) composita Tucker? (Hurd and Linsley 1959, adults entering nests), Melissodes (Eumelissodes) trinodis Robertson (Graenicher 1905, observations of host and parasite larvae). Floral Records.—Aster sp. (= Symphyotrichum), Chrysanthemum sp., Chrysopsis mariana (L.) Ell., Chrysothamnus sp., Gaillardia sp., Grindelia squarrosa (Pursh) Dunal, Helianthus annuus L., H. grosseserratus Martens, H. petiolaris Nutt., H. tuberosus L., Heterotheca latifolia [= He. subaxillaris (Lam.) Britt. & Rusby], Phacelia sp., Polygonum sp. Salvia sp., Verbena stricta Vent., Verbesina sp., Vernonia baldwinii Torr., V. m i ss u ri ca Raf., Viguiera sp. Seasonal Records.—20 June to 26 October. Specimens examined.—329 &, 37 % (AUSTIN, BERLIN, BERKELEY, BOULDER, CHAMELA, CORVALLIS, DAVIS, GAINESVILLE, LAWRENCE, LOGAN, LOS ANGELES, NEW YORK, NEW YORK-ASCHER, RIVERSIDE, SAN DIEGO, SAN FRANCISCO, STARKVILLE, URBANA, WASHINGTON D.C.).

Epeolus heterurus Cockerell and Sandhouse 1924: 316–317 [Holotype: California Academy of Sciences No. 1611; &, Redding (Shasta Co.), California; 6 July 1918. Epeolus piscatoris Cockerell 1939: 432–433 [Holotype: California Academy of Sciences No. 6652; &, Fisherman’s Cove, Santa Catalina Island, California; 9 June 1933; Sinapis]; Rust 1984: 120 [synonymy]. Epeolus utahensis Cockerell 1921: 15–16 [Holotype: American Museum of Natural History No. 25112; %, Huntsville (near Ogden, Weber Co.), Utah; 26 July 1920; about 41° 17’N, 110° 46’W]. new synonymy Triepeolus piscatoris; Brumley 1965: 73. Triepeolus heterurus; Hurd 1979: 2093.

Description.—Length ca. 7–9 mm; ITW 1.3–1.9 mm. Integument black, with red on apical half of mandible; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus lacking midline and larger punctures, sometimes covered with sparse, pale setae. Paramedian band clearly separated from other pale setae on mesoscutum (some females), or contiguous with pale setae on anterior margin of mesoscutum (males and most females). Scutellum weakly bigibbous; axillar spine triangular, reaching or not attaining midpoint of scutellum. Mesepisternum with dorsal half covered with white, suberect to erect branched setae, anterior margin with sparse, erect, simple setae, ventral half with punctures nearly contiguous, often covered with sparse, brown, branched setae (females), or covered with erect to suberect, dense, white, branched setae (males). T1 discal patch subovate to trapezoidal; T2 with LLB forming acute angle with ATB. Female: Pseudopygidial area with long setae, parted medially and directed laterally, apical margin concave; S5 slightly downcurved at apical margin. Male: Pygidial plate very narrow and long, apparently lacking transverse basal ridge; S2–S3 with apical bands of white setae (S3 with white setae slightly surpassing apical margin); S4–S5 with brown apical fringes. Comments.—This species is very similar to T. melanarius. Females can easily be distinguished by the pseudopygidial area, which is parted medially and apically concave in T. heterurus. Males of the two species are extremely similar, but can be differentiated by the apical bands of white setae on S2–S3 in males of T. heterurus (metasomal sterna entirely brown in T. melanarius). In addition, in males, the ventral half of the lateral surface of the mesepisternum is darker in T. melanarius than in T. heterurus, and the paramedian bands are clearly separated from other pale setae on the mesoscutum in T. melanarius, while in T. heterurus they are usually surrounded by diffuse pale setae on the anterior margin of the mesoscutum. A variety of this species exists [represented by four females, one from Baja California (repository: San Francisco), two from Westwood Hills, Los Angeles Co., California (repository: Berkeley), and one from Glendora, Los Angeles Co., California (repository: Urbana)], in which the pseudopygidial area has setae that are much shorter than that of the typical T. heterurus. In addition, in these females, the integument is often brownish red instead of black, and the mesepisternum has a distinct, medioventral spot of brown setae. Addi- tional material may support recognizing these specimens as another species. Timberlake gave the specimens from Westwood Hills two different manuscript names, one honoring E. Gorton Linsley, and the other drawing attention to the red legs of the specimen. Distribution.—MéXICO: Baja California; USA: California (northern counties, continuing south through the Central Valley and Coast Range Mountains, and further south through coastal counties to San Diego), Idaho, Nevada, Oregon, Utah, Wyoming. Floral Records.—Chrysothamnus sp., Eriogonum sp. Grindelia camporum Greene, Hemizonia pugens (Hook. and Arn.) Torrey and Gray, Mentha spicata L., Sinapis sp., Solidago californica Nutt., Wislizenia refracta Engelm. Seasonal Records.—5 May to 16 October.

70 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Specimens examined.—71 &, 90 % (BERKELEY, BOULDER, CORVALLIS, DAVIS, GAINESVILLE, LAWRENCE, LOGAN, LOS ANGELES, NEW YORK, NEW YORK-ASCHER, SAN FRANCISCO, URBANA, WASHING- TON D.C.).

TRIEPEOLUS INTERRUPTUS new species (Figs. 92, 93)

Diagnosis.—This species can be distinguished by the combination of the absence of the LLB and usually the BTB on T1, the medially interrupted ATB on T1, the absence of the LLB on T2, and the nearly uniform setae on the pseudopygidial area. This species strongly resembles T. parkeri, but can be distinguished from that species by its medially interrupted ATB on T1 (medially continuous in T. parkeri), and by the presence of a paramedian band on the mesoscutum in T. parkeri (usually absent in T. interruptus). Description of holotype female.—Length ca. 8 mm (7–10.5 mm); ITW 1.4 mm (1.4–1.8 mm). Integu- ment black, with red on apical half of mandible, brownish orange on F1 (sometimes on tegula and part or most of legs); dorsum of mesosoma and metasoma with bands of yellow to yellow-orange setae. Clypeus with faint (sometimes strong) midline and faint larger punctures, covered with sparse white setae (less so in some females, or entirely covered by dense, white setae in males). Paramedian band absent (present in specimen from Hidalgo). Scutellum weakly bigibbous; axillar spine triangular, almost reaching (or reaching) midpoint of scutellum. Mesepisternum with very sparse, short, erect, simple setae (apparently absent in some specimens); with sparse, pale, branched setae near pronotal lobe and beneath scrobal groove (sometimes absent); ventrally asetose, with integument shining, punctures nearly contiguous to separated by ca. 0.5 puncture width (or entire mesepisternum except hypoepimeron covered with dense, white intermixed with brown posteroventrally, branched setae in males). T1 lacking BTB (sometimes with BTB reduced to patch on anterolateral corner); ATB interrupted medially; T2 with LLB absent. Pseudopygidial area semicircular (to subquadrate or subtriangular), with vague basal crescent of shining setae; S5 straight in profile. (Male: Pygidial plate keyhole shaped, with somewhat weak basal transverse ridge and apical downturned plate; S2–S3 with apical bands of white setae, S3 with white setae slightly surpassing apical margin; S4–S5 with apical fringes of setae white, intermixed with brown on S5.) Comments.—This species was labeled “PCAM 32” by D. Yanega. Distribution.—MéXICO: Guanajuato, Hidalgo, Michoacán, Morelos, Zacatecas. Host Records.—Atoposmia sp.? (1 specimen: “México: Hidalgo 4 km SW Metzquititlan, 11 Nov 1991. Noguera, 1580m, rocky hillside, composites, Anthocopa on rock wall”). Seasonal Records.—7 September to 6 December. Holotype.—“MÉXICO Michoacán, 3 km N Arteaga, 850 m, I-XI [November]-1987, T. Griswold // Native Bee Survey USDA Logan, Utah BBSL581418 // Triepeolus sp. 30 T. Griswold det // M. G. Rightmyer Triepeolus Database Specimen No. 3551 // HOLOTYPE & Triepeolus interruptus Rightmyer 2007” (LOGAN). Paratypes.—MÉXICO: GUANAJUATO: Cuerámaro, San Gregorio, 7 September 1989, L. Godinez, 1700 m (1%, MEXICO CITY); HIDALGO: 4 km N Metzquititlan, 11 November 1991, Noguera, 1580 m, rocky hillsides, composites, Anthocopa on rock wall (1&, LAWRENCE); 22 km SW Metzquititlan, 11 Novem- ber 1991, T. Griswold, 1750 m (1&, LAWRENCE); MORELOS: Cuernavaca, 8 September–6 November 1987, F. D. Parker (1&, LOGAN); ZACATECAS: 5 mi Tabasco, 18 September 1970, R. M. Bohart (1&, DAVIS); G. E. & R. M. Bohart (1&, LOGAN). Etymology.—This name is in reference to the medially interrupted bands of pale setae on the metasomal terga. From the Latin, “interruptus” (broken apart).

Epeolus intrepidus Smith 1879: 102 [Holotype: The Natural History Museum, London No. 17B.521; %, México]. Triepeolus digueti Cockerell 1905b: 165 [Holotype: U. S. National Museum of Natural History No. 23290; &, Oaxaca, México]. new synonymy Epeolus nobilis Friese 1908: 85 [Lectotype: Zoologisches Museum, Humboldt-Universität; %, Argentini (= Argentina, Chiapas?), M. (= México?); 1900]. new synonymy, new lectotype designation Triepeolus intrepidus; Cockerell 1949: 460.

Description.—Length ca. 12–17 mm; ITW 2.7–3.2 mm. Integument black, with red to orange on basal mandible, labrum, apical or entire clypeus, scape, pedicel, F1, basal F2, tegula, and legs (excluding coxae and sometimes trochanters and spurs), usually on part or entire scutellum (contrasting with black axillar spine), sometimes partially on mesoscutum; dorsum of mesosoma and metasoma with bands of bright yellow setae. Clypeus lacking or with weak midline and larger punctures, often covered with sparse golden setae. Parame- dian band absent; mesoscutum densely covered with erect, golden setae. Scutellum moderately bigibbous; axillar spine rounded apically, not attaining midpoint of scutellum. Mesepisternum with erect, golden, simple and minutely branched setae (usually denser in males); ventral half with punctures nearly contiguous to separated by two puncture diameters. T1 discal patch triangular; T2 with LLB absent or forming acute angle with ATB. Female: Pseudopygidial area golden, poorly differentiated from rest of T5, indicated by flattened, ovate plane; S5 very slightly downcurved. Male: Pygidial plate longitudinally elongate, slightly keyhole shaped, lacking distinct apical downturned plate; S3–S5 with well-developed apical fringes of golden setae. Comments.—Triepeolus intrepidus is more commonly known by its junior synonym, T. digueti. It is readily recognized by the bright red scutellum, contrasting with the short, black axillar spines; by the erect, golden setae covering much of the head and mesosoma; and by the pattern of the bands of yellow setae on the metasoma. The name “Triepeolus nobilis,” a synonymous name with T. intrepidus, has long been confused with the species T. osiriformis, possibly due to the fact that both T. intrepidus and T. osiriformis usually have a bright red scutellum, and because of the confusing collection locality of the T. nobilis lectotype (i.e., “M. Argen- tini”), which is suggestive of the country Argentina, rather than what is presumably a locality in México. In his original description of T. osiriformis, Schrottky (1910) notes the fact that the species is quite distinct from T. nobilis, despite running to that species in the key provided by Friese (1908). In fact, T. intrepidus appears to be restricted to Central and North America, while T. osiriformis is widely distributed in South America, including Argentina. Friese’s original description of Epeolus nobilis was based on both males and females. In the Berlin Museum, both a male and female of this species identified by Friese are present, but only the male specimen bears a type label; thus I recognize the male as the lectotype specimen. The lectotype label data are as follows: “M // Argentini 1900 // Epeolus nobilis Fr. % 1907 Friese det. // Type [red label] // Lectotype % Epeolus nobilis Friese des. M. Rightmyer 2006” Distribution.—MéXICO: Chiapas, Chihuahua, Guerrero, Hidalgo, Jalisco, México, Michoacán, Morelos, Nuevo León, Oaxaca, Zacatecas; USA: Arizona, New Mexico, Texas (Culberson Co.). Host Records.—Melissodes (= Syntrichalonia) exquisita (Cresson) (Cockerell, 1905b, flying near nest entrance). Floral Records.—Asclepias subverticillata (A. Gray) Vail, Encelia farinosa Gray ex Torr., Helianthus sp., Senecio longilobus (=Senecio flaccidus Less. var. flaccidus), S. salignus [=Barkleyanthus salicifolius (Kunth) H.E. Robins. & Brett.], Simsia exaristata (= Simsia lagasceiformis DC.), Tagetes erecta L., Viguiera dentata (Cav.) Spreng. Seasonal Records.—26 July to 13 November.

TRIEPEOLUS ISOHEDRUS new species (Figs. 96, 97)

Diagnosis.—This species primarily can be distinguished by the combination of the strongly triangular pseudopygidial area, which has a distinct basal region of silvery setae; the paramedian band, which is laterally contiguous with pale yellow setae on the anterior margin of the mesoscutum; relatively short clypeus; the absence of erect, simple setae on the mesepisternum; and the reddish-tinged axillar spine. Triepeolus partitus resembles T. isohedrus, but in T. partitus the pseudopygidial area is not so distinctly triangular, and there are coarse setae present apically that are slightly longer than those in T. isohedrus; the T1 discal patch is more ovate (as opposed to trapezoidal or subtriangular in T. isohedrus); and the paramedian bands are not laterally contiguous with other pale yellow setae on the anterior margin of the mesoscutum. Description of holotype female.—Length ca. 11.5 mm (10.5–13.5 mm); ITW 2.3 mm (2.2–2.6 mm). Integument black, with reddish orange on basal half of mandible, labrum, apical clypeus, scape, pedicel, F1, pronotal lobe, tegula, tip of axillar spine (most specimens) and legs excluding basal margins of coxae and spurs; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus relatively flat in profile, with faint midline and larger punctures (sometimes covered with sparse, pale setae). Paramedian band slightly curving outward anteriorly, connected with yellow lateral setae on subapical margin of mesoscutum, forming anchor shape. Scutellum moderately bigibbous; axillar spine triangular, slightly exceeding (or only reaching) scutellar midpoint, apical point directed inward. Mesepisternum lacking erect, simple setae; dorsal half covered with dense, pale yellow, branched setae, slightly sparser (to absent) on hypoepimeron; ventral integument shining, with punctures relatively small, separated by up to 1–2 puncture widths. T1 discal patch trapezoidal (to subtriangular, or subovate); T2 with LLB forming acute angle with ATB. Pseudopygidial area strongly triangular, with faint inverted V-shaped region of silvery reflectance basally, and with paler-reflecting, slightly upturned setae on apical margin; S5 straight in profile. (Male: Unknown.) Comments.—One specimen of this species was given a manuscript name in reference to its collection locality in Kansas by Timberlake. Distribution.—USA: Arizona, Kansas, New Mexico, Texas. Floral Records.—Grindelia sp. Seasonal Records.—1 May to 15 September. Holotype.—“NEW MEXICO: Hidalgo Co., 25–28 mi. S. Animas, VIII [August]-16-94 [1994], J. G. Rozen & J. S. Ascher // M. G. Rightmyer Triepeolus Database No. 6970 // HOLOTYPE & Triepeolus isohedrus Rightmyer 2007” (NEW YORK). Paratypes.—USA: ARIZONA: Cochise Co., 35 mi. S of Animas, 15 September 1999, Rozen, LeBuhn, Rightmyer (1&, NEW YORK); 1 mi. E. Douglas, 29 August 1973, J. G. Rozen, M. Favreau, R. McGinley (1&, NEW YORK); Apache, 19 August 1979, R. M. Bohart (1&, DAVIS); Santa Cruz Co., Duquesne Rd., 2.6 mi E San Rafael Rd., 28 August 1995, R. R. Snelling (2&, LOS ANGELES); 5 mi. NNE Sonoita, 19 August 1969, J. G. & B. L. Rozen (1&, NEW YORK); Sonoita, 6 September 1957, T. R. Haig (1&, DAVIS); KANSAS: Scott Co., 16 August 1925, Beamer, Lawson (1&, LAWRENCE); (Kearny Co.), 4 mi E Lakin, 7 September 1930, Timberlake, at flowers of Grindelia (1&, BERKELEY); NEW MEXICO: Dona Ana Co., 5 mi. NE Las Cruces, 21 August 1962, H. A. Scullen, 4200 ft. (1&, CORVALLIS); Hidalgo Co., 29 mi. S. Animas, 29 August 1982, J. G. & B. L. Rozen (1&, NEW YORK); 25–28 mi. S. Animas, 16 August 1994, J. G. Rozen, J. S. Ascher (1&, NEW YORK); 27–32 mi. S. Animas, 24 August 1994, J. G. Rozen, J. S. Ascher (1&, NEW YORK); 26 mi. S. Animas, 19 August 2001, J. G. Rozen (1&, NEW YORK); 33–37 mi. S. Animas, 22 August 1997, J. G. Rozen,

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 73 B. McAdams (1&, NEW YORK); TEXAS: Lubbock Co., Lubbock, 1 May 1967, P. L. Wink (1&, LOS ANGE- LES); (Hale Co.), Plainview, Playa, 25 August 1969, D. Bennett, R. Kirby, Sweep Net (1&, LOS ANGELES). Etymology.—This species is named for the nearly equally sided, triangular pseudopygidial area. From the Greek, “isos” (equal) and “hedra” (plane or side).

TRIEPEOLUS JENNIEAE new species (Figs. 98, 99)

Diagnosis.—This species is known by the combination of the absence of the LLB on T1; the poorly differentiated pseudopygidial area, which forms a wide, slightly downturned area at the apical margin of T5; the absence of erect setae on the mesepisternum; and the broadly interrupted transverse bands of pale setae on the metasoma. Description of holotype female.—Length ca. 8 mm; ITW 2.1 mm. Integument black, with orange on mandible, labrum, scape, pedicel, F1 (rest of antenna dark reddish brown), pronotal lobe, tegula, and patches on femora and tibiae; dorsum of mesosoma and metasoma with bands of yellow setae. Clypeus lacking midline, with very faint larger punctures. Paramedian band tapering anteriorly, clearly separated from other pale setae on mesoscutum. Scutellum weakly bigibbous; axillar spine triangular, not attaining midpoint of scutellum. Mesepisternum lacking erect, simple setae; dorsally with longitudinal patch of pale yellow, branched setae below scrobal groove, extending between anterior and posterior margins and continuing along posterior margin, also with patch of pale setae near pronotal lobe and on anterior surface of mesepisternum at same level; ventrally with integument shining, with very small, weakly impressed punctures, separated by one or two puncture diameters (one side of mesepisternum with linear area of impunctate integument spanning ca. five puncture diameters). T1 with BTB and ATB parallel; both T1 and T2 lacking LLB; T1–T3 with ATBs widely interrupted medially (rest of terga not visible). Pseudopygidial area quadrate, poorly differentiated from basal setae on T5, setae fine, apically with patch of golden-reflecting setae (apparently differentiated due to downturn in integument); S5 straight in profile. (Male: Unknown.) Comments.—This species was labeled “PCAM-27” by D. Yanega. Distribution.—MéXICO: Hidalgo. Host Records.—Tetraloniella (Pectinapis) sp.? (from holotype specimen). Seasonal Records.—11 November. Holotype.—“MÉXICO: Hidalgo, 25 km SW Matzquititlan, 11 Nov [November] 1991. Noguera, 1860 m, near top of valley, Pectinapis in Salvia woods // Triepeolus PCAM-27 // Triepeolus PCAM-27 det D. Yanega // female // SM0326440 KUNHM-ENT // M. G. Rightmyer Triepeolus Database Specimen No. 1274 // HOLOTYPE & Triepeolus jennieae Rightmyer 2007” (LAWRENCE). Etymology.—This species is named in honor of my mother, Jennie Anne Rightmyer, who taught me to love the natural world.

TRIEPEOLUS JOLIAE new species (Figs. 100, 101)

Diagnosis.—This species can be distinguished from all but T. californicus by the combination of the shape of the T1 discal patch, which is transversely ovate and has an enlarged circular region medially, and by the nearly uniform setae of the pseudopygidial area. Triepeolus joliae and T. californicus can be separated based on the angle formed by the LLB and ATB on T2 (acute in T. californicus, obtuse or absent in T. joliae), by the paramedian bands (long and distinct in T. californicus, shorter and often connected laterally to diffuse pale setae on anterior margin of the mesoscutum in T. joliae), and by the color of the setae on the pseudopygidial area

74 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER (golden in T. californicus, coppery in T. joliae). In addition, the flight season of T. californicus is in the summer and early fall, while that of T. joliae is in the spring. Description of holotype female.—Length ca. 10 mm (10–11.5 mm); ITW 2.5 mm (2.2–3.0 mm). Integ- ument black to reddish brown, with red on basal half of mandible, entire (or part of) labrum, and apical margin of clypeus (most specimens); orange on scape (most specimens), pedicel (most specimens), F1 and legs excluding basal halves of coxae and spurs (sometimes on entire coxae, or only on legs distal to femora on front leg); brownish orange on tegula (most specimens); dorsum of mesosoma and metasoma with bands of yellow to pale yellow setae. Clypeus with faint midline and larger punctures (sometimes obscured by sparse white setae; these setae denser in males). Paramedian band laterally contiguous with diffuse, pale setae on anterior of mesoscutum (especially in males; or distinct; worn away in one specimen). Scutellum weakly bigibbous, posterior surface somewhat extended into posterior ridge; axillar spine triangular, not (or barely) reaching midpoint of scutellum. Mesepisternum lacking erect, simple setae; densely (to sparsely) covered with pale yellow, branched setae (or dorsally with dense, pale yellow, branched setae, ventrally with dark brown, branched setae in male). T1 discal patch very wide, narrowly ovate (to rectangular), with median circular patch dark setae; T2 with LLB nearly absent (or forming very obtuse angle with ATB). Pseudopygidial area subquadrate (to semicircular) with uniform, coppery, long, fine setae; S5 straight in profile. (Male: Pygidial plate long and moderately narrow, lacking distinct transverse basal ridge and apical downturned plate; S2–S3 with white apical setae, S3 with white setae slightly surpassing apical margin; S4–S5 with golden to white apical fringes.) Distribution.—USA: Arizona, California, Nevada, Utah. Floral Records.—Argemone sp., Coreopsis sp., Ephedra sp. Larrea sp., Senecio douglasii [= Senecio flaccidus Less. var. douglasii (DC.) B.L. Turner & T.M. Barkl.]. Seasonal Records.—10 April to 15 June. Holotype.—“Usa [USA], Arizona: Maricopa Co., 10 mi SE Aguila, May 2 1990, J. G. Rozen // M. G. Rightmyer Triepeolus Database Specimen No. 3558 // HOLOTYPE & Triepeolus joliae Rightmyer 2007” (NEW YORK). Paratypes.—USA: ARIZONA: La Paz Co., Hwy 93, MP 143, 23 April 1994, W. R. Bowlin, 12–12:30, Argemone sp. (1&, LOGAN); Yavapai Co., 8 mi NW Wickenburg, 4 May 1992, J. G. Rozen (1&, NEW YORK); Yuma Co., Telegraph Pass, 11 April 1983, J. C. & E. M. Hall (1&, RIVERSIDE); CALIFORNIA: (Imperial Co.), Holtville, 10 April 1947, G. H. & J. L. Sperry, Coreopsis sp. (1&, LAWRENCE); Inyo Co., 5 mi NW Pan- amint Spr., 8 May 1958, A. E. Menke (1&, DAVIS); 2 mi E Lone Pine, 19 May 1970, R. M. Bohart (1%, DAVIS); Lone Pine, Whitney Road, 4 June 1937, E. C. VanDyke (1%, SAN FRANCISCO); San Bernardino Co., 10 mi S Adelanto, 27 May 1978, L. G. Bezark, Ephedra sp. (2%, DAVIS); Dumont, 33 mi N Baker St. Hwy. 127, Mojave Des., 11 May 1982, K. V. Krombein (1&, WASHINGTON D.C.). Additional specimens.—NEVADA: Clark Co., 29.1 mi SW Mesquite, 2 May 1986, P. F. & D. M. Tor- chio (8&, LOGAN); 33 mi S Mesquite, 17 May 1995, P. F. Torchio, Senecio douglassi (1&, LOGAN); UTAH: Washington Co., Santa Clara, F. Parker, P. Torchio, 30 May 1973, Larrea sp. (1&, 1%, LOGAN); Paradise Can- yon, 30 May–4 June 1983, D. Beck (1%, LOGAN); Snow Canyon, 15 June 1983, W. J. Hanson (2&, LOGAN). Etymology.—This species is named in honor of my sister, Joli K. Rightmyer, for her support, encourage- ment, and friendship.

TRIEPEOLUS LATERALIS new species (Figs. 102, 103)

Diagnosis.—This species can be distinguished by the combination of the absence of erect, simple setae on the mesepisternum, which has pale, plumose setae restricted to the dorsal third excluding the hypoepimeron, or to

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 75 small patches below the scrobal groove and posterior to the pronotal lobe; the metanotum, which has pale setae restricted to the lateral corners, such that the median patch of dark setae is much wider than 1 OD; the parallel BTB and ATB on T1; and the ovate pseudopygidial area with long, darkly shining setae. Triepeolus lateralis strongly resembles T. scelestus, but differs in the following characters: on the female T5, there is a well-differentiated patch of white setae lateral to the pseudopygidial area in T. lateralis, while in T. scelestus these white setae are typically reduced to the apical margin of the tergum, or fully present but rather sparse and not so strongly differentiated from the pseudopygidial area; the metanotum is entirely covered with pale yellow setae in T. scelestus, while in T. lateralis these pale setae are restricted to the lateral margins of the metanotum; the ATBs on the metasoma are medially interrupted on only T1–T2 in T. scelestus, while in T. lateralis these bands are often medially interrupted at least on T1–T3; and the venter of the metasoma lacks white setae in T. scelestus, while in T. lateralis white setae are present on the apical margins of S2– S4. Description of holotype female.—Length ca. 10.5 mm (9–11 mm); ITW 2.0 mm (1.9–2.5 mm). Integu- ment black with red on basal half of mandible; and orange on F1 (rarely entirely black; less commonly also with orange on labrum, apical clypeus, tegula, and parts of legs); dorsum of mesosoma and metasoma with bands of yellow-orange (to yellow) setae. Clypeus lacking (or with very faint) midline and larger punctures (sometimes covered with sparse setae). Paramedian band relatively short and very narrow (sometimes less narrow), clearly separated from other pale setae on mesoscutum. Scutellum weakly (to moderately) bigibbous; axillar spine triangular, not (or nearly) reaching midpoint of scutellum; posterior margin of scutellum with pale setae restricted to lateral margins. Mesepisternum lacking erect, simple setae; dorsally with region of white, branched setae limited to regions below scrobal groove and posterior to pronotal lobe (or on dorsal third excluding hypoepimeron); integument shining, punctation small, somewhat weakly impressed, separated by up to one puncture diameter in some places. T1 discal patch strongly rectangular (to quadrate), forming plus-shaped sign with medially-interrupted BTB and ATB (sometimes this pattern less obvious; sometimes lacking pale setae at basolateral corner of T1); T2 with LLB absent (or forming weakly acute angle with ATB); at least T1–T2 (often also T3 and T4) with apical bands medially interrupted. Pseudopygidial area widely ovate (to circular), with nearly uniform, relatively long, fine, golden setae; T5 with distinct patch white setae lateral to pseudopygidial area; S2–S4 with apical transverse bands of white setae; S5 faintly downcurved apically. (Male: Unknown.) Comments.— This species was labeled “PCAM 28” by D. Yanega. Distribution.—MéXICO: Guerrero, Michoacán, Morelos, Oaxaca, Puebla. Floral Records.—Gymnosperma glutinosum (Spreng.) Less. Seasonal Records.—6 September to 25 January. Holotype.—“MÉXICO: Morelos, Sierra de Huatla, C.E.A.M.I.S.H. [Center for Environmental Educa- tion and Research of Huatla Sierra] Huatla, 2.5 km N, 4.0 km W, 18°27’0”N, 99°2’0”W, 1050 m, 6 SEP [Sep- tember] 1996, R. Brooks, MEX 1B96 024, ex. plant #144 RA // SM0253481 KUNHM-ENT // M. G. Rightmyer Triepeolus Database Specimen No. 6809 // HOLOTYPE & Triepeolus lateralis Rightmyer 2007” (LAWRENCE). Paratypes.—MÉXICO: MICHOACÁN: Morelia, 20 September 1957, H. A. Scullen, 6500 ft. (1&, CORVALLIS); MORELOS: Cuernavaca, 4–6 December 1987, F. D. Parker (1&, LOGAN); OAXACA: 1.4 km SW Puebla/Oaxaca border, Hwy 125, km 58, 14 September 1996, R. Brooks, 18°12’0”N, 97°39’0”W, 1910 m. (1&, LAWRENCE); PUEBLA: 16 km NE Tehuacán, 4 November 1991, R. Ayala, 1820 m. (1&, LAWRENCE); Zapotitlán, Salinas, 29 October 1996, C. V., M. C., 18°20’50”N, 97°34’35”W (1&, PUEBLA); 25 January 1997, 18°18’20”N, 97°32’20”W, Gymnosperma glutinosum (1&, PUEBLA). Etymology.—This name is in reference to the yellow setae, which are restricted to the lateral margins of the metanotum, and which help differentiate this species from T. scelestus. From the Latin, “lateralis” (of the side).

Triepeolus laticaudus Cockerell 1921: 12–13 [Holotype: American Museum of Natural History No. 25096; &, Ute Pass, Cascade (El Paso Co.), Colorado; 22 August 1914].

Description.—Length ca. 7.5–12 mm; ITW 1.6–2.7 mm. Integument black, with dark reddish brown to orange on apical portion of mandible, F1, and tegula; orange on part or entire legs (excluding basal coxae and spurs), often on labrum, apical clypeus, scape, and pedicel; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint to strong midline and larger punctures. Paramedian band clearly separated from other pale setae on mesoscutum. Scutellum moderately bigibbous; axillar spine triangular, reaching midpoint of scutellum. Mesepisternum lacking erect, simple setae; dorsal half covered with dense, pale yellow, branched setae (absent or sparser on hypoepimeron); ventrally sometimes with pale, branched setae also on margins, enclosing vague black region medially (sometimes with sparse pale setae medially as well); punctures nearly contiguous to separated by a puncture diameter, with integument between shining, somewhat tuberculate in appearance. T1 widely trapezoidal; T2 with LLB forming weakly acute angle with ATB, mostly on lateral surface of T2. Female: Pseudopygidial area subtriangular to subquadrate, with distinct, relatively long, basal shining region, extending laterally to apical margin of area, with median circular region of stout setae; S3–S4 (and sometimes S2) with patches of white setae laterally; S5 straight in profile to slightly downcurved. Male: Unknown. Comments.—This species is superficially similar to T. cressonii, but can be differentiated from that species by the mesepisternum (in T. laticaudus the dorsal third is mostly covered by dense, pale setae, and medioventrally the integument between the punctures is distinctly raised in places, appearing tuberculate, while in T. cressonii the pale setae is more restricted dorsally, and the integument is much flatter between punctures medioventrally). Also, the basal shining setae of the pseudopygidial area are longer in T. laticaudus than in T. cressonii. Triepeolus laticaudus is also similar to T. circumculus, but can be differentiated from that species by the mesepisternum, which has sparser punctures and the integument between the punctures appears more tuberculate. Finally, this species is similar to T. townsendi, but can be separated from that species by the punctation of the mesepisternum (denser in T. townsendi), and by the relative differentiation of the basal shining setae on the pseudopygidial area (less differentiated in T. townsendi). Distribution.—MéXICO: Coahuila, Durango; USA: Arizona, Colorado, Kansas, New Mexico, South Dakota, Texas. Floral Records.—Amphiachyris dracunculoides (DC.) Nutt., Aster sp. (= Symphyotrichum), Bidens pilosa L., Calyptocarpus vialis Less., Gaillardia sp., Grindelia squarrosa (Pursh) Dunal, Gutierrezia sarothrae (Pursh) Britt. & Rusby, Haplopappus gracilis [= Machaeranthera gracilis (Nutt.) Shinners], Heteroth- eca subaxillaris (Lam.) Britt. & Rusby, H. villosa (Pursh) Shinners, Melilotus officinalis (L.) Lam., Solidago serotina (= S. gigantea Ait.), Verbesina sp. Seasonal Records.—11 June to 6 November. Specimens examined.—87 & (AUSTIN, BOULDER, CHAMELA, DAVIS, GAINESVILLE, LAWRENCE, LOGAN, LOS ANGELES, NEW YORK, TEMPE, WASHINGTON D.C.).

TRIEPEOLUS LATICEPS (Friese) new combination (Figs. 106, 107)

Epeolus laticeps Friese 1917 [1916]: 338 [Holotype: Zoologisches Museum, Humboldt-Universität; %, Jacubaya (=Tacu- baya?, see Comments under T. grandis), México; 1900].

T. laticeps T. tepanecus T. mauropygus

Clypeus midline strong on entire length of strong on entire length of present dorsally clypeus, or rarely present only clypeus dorsally Paramedian band of present, narrow or wide absent or present, narrow present, narrow setae Mesoscutum puncta- punctures irregular but very punctures mostly separated punctures regularly spaced tion closely spaced, mostly nearly by a puncture width or more and nearly contiguous contiguous (mesoscutum shining) Mesepisternum punctures irregularly spaced punctures irregularly spaced punctures nearly contiguous, (nearly contiguous to one (nearly contiguous to 2–3 with few intervening spaces puncture diameter apart), puncture diameters apart), of puncture diameter; erect, integument between often integument shining; erect, simple setae short and sparse raised; erect, simple setae simple setae short and sparse short yet conspicuous Axillar spine triangular, often with small triangular to sharply pointed; triangular, reaching scutellar incurved free apical point; not reaching or well surpassing midpoint attaining to slightly surpassing scutellar midpoint scutellar midpoint Propodeum larger punctures on lateral larger punctures on lateral larger punctures on lateral fourth to fifth fifth third Leg color black, often with red mid and black black hind legs T1 discal patch triangular, rarely transversely triangular transversely ovate to widely ovate trapezoidal T2 LLB present or absent absent present or rarely absent Metasomal bands of relatively wide, but sometimes relatively narrow relatively narrow pale setae narrow on T1 Pseudopygidial area subquadrate to subovate, setae poorly defined basal bound- subquadrate, setae very slightly denser basally ary (ovate?), setae slightly slightly denser basally suberect basally Distribution Arizona to Texas, south to El Nayarit southeast to Chiapas Arizona south to Sinaloa Salvador

Description.—Length ca. 9–14 mm; ITW 2.0–3.1 mm (the larger size more common); integument black, with red on basal half of mandible; orange on F1, less commonly on tegula and part or most of legs (especially on middle and hind legs, not on coxae or spurs); dorsum of mesosoma and metasoma with bands of yellow- orange to pale yellow setae. Clypeus with strong midline, usually covered with white setae, with tuberculate- like setal bases apparent beneath (denser in male). Upper face with rough punctation, integument shining. Paramedian band clearly separated from other pale setae on mesoscutum. Scutellum moderately to weakly bigibbous; axillar spine triangular, reaching midpoint of scutellum. Mesepisternum lacking erect, simple setae or with very sparse, short, erect, simple setae; with region of dense, suberect setae (both dark and pale) beneath scrobal groove, along posterior margin, and often posterior to pronotal lobe and on anterior surface of mesepisternum; ventral integument sometimes sparsely covered with dark brown, branched setae; integument shining, but irregularly and densely punctate, integument between slightly raised, punctures at most one puncture diameter apart (females), or entire mesepisternum covered with pale yellow, branched setae (male). T1 discal patch relatively wide, trapezoidal to subtriangular or subovate, BTB and ATB interrupted medially,

78 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER ATB narrowed in some specimens from México, more robust in specimens further south; T2 with LLB absent (some specimens from México) or forming weakly acute to acute angle with ATB (México and other localities). Female: Pseudopygidial area semicircular, often darkly shining, with vague basal crescent of finer setae; S2–S4 with apicolateral bands pale yellow setae (often connected medially on S2–S3); S5 straight in profile. Male: Pygidial plate keyhole shaped, with moderately distinct apical downturned plate (basal ridge obscured by long, brown setae); S2–S3 with apical bands of white setae (S3 with white setae slightly surpassing apical margin); S4–S5 with apical fringes of pale golden setae on S4, brown on S5. Comments.—This species is very similar to T. tepanecus and, slightly less so, to T. mauropygus; all three species share the darkly shining pseudopygidial area, with a poorly differentiated basal region of finer, denser setae, and the sparse, short, erect, simple setae of the mesepisternum. Due to these similarities, T. tepanecus, T. laticeps, T. mauropygus are grouped together in a group here termed the “T. tepanecus species group.” See Table 2 for further characteristics of each species. Triepeolus laticeps additionally resembles T. medusa, but can be separated from that species by the more widely spaced, larger punctures on the mesepisternum of T. laticeps (versus the nearly contiguous, fine punctures on the mesepisternum of T. medusa). The original description of Epeolus laticeps is based on one male specimen; thus it is the holotype specimen by default. A specimen of this species was labeled “PCAM 8” by D. Yanega. Distribution.—EL SALVADOR: Quezaltepeque; GUATEMALA: Rabinal; MéXICO: Chihuahua, Coahuila, Colima, Durango, Guanajuato, Guerrero, Jalisco, México, Michoacán, Morelos, Puebla, Querétaro, San Luis Potosí, Sonora, Zacatecas; USA: Arizona, New Mexico, Texas (Brewster Co.). Floral Records.—Simsia amplexicaulis (Cav.) Pers., Zinnia sp., “Leguminosae”, “blue aster”, “Santa Maria weed”. Seasonal Records.—31 May to 30 October. Specimens examined.—81 &, 3 % (BERLIN, BERKELEY, CORVALLIS, DAVIS, GAINESVILLE, LAWRENCE, LOGAN, MEXICO CITY, NEW YORK, RIVERSIDE, SAN FRANCISCO, WASHINGTON D.C.).

TRIEPEOLUS LOOMISORUM Rozen (Figs. 108, 109)

Triepeolus loomisorum Rozen 1989a: 14–18, Figs. 9, 22, 33–36 [Holotype: American Museum of Natural History; &, 13 mi SW Apache, Cochise Co., Arizona; 1 September 1988], 17, 18, Fig. 25 [description of biology; description, illustration of egg]; Rozen 1989b: 14, 15, Figs. 19–23 [description, illustrations of first instar]; Rozen 2003: Fig. 53 [egg compared with others at same scale].

Description.—Length ca. 6–8.5 mm; ITW 1.3–1.8 mm. Integument black, with red to orange on basal half of mandible, labrum, apical clypeus, scape, pedicel, F1 (sometimes also remaining flagellomeres), pronotal lobe, tegula, legs (excluding basal coxae, spurs, and sometimes scattered areas on femora and tibiae), and parts of ventral metasoma; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with weak midline and very weak larger punctures, densely covered with white setae in males. Paramedian band tapering to anterior margin, clearly separated from other pale setae on mesoscutum. Scutellum weakly bigibbous; axillar spine triangular, not or nearly reaching midpoint of scutellum. Mesepisternum lacking erect, simple setae; mostly covered with dense, pale, branched setae except for mediobasally with asetose, circular region of shining integument; with punctures nearly contiguous to separated by up to 3 puncture diameters in shining mediobasal region (females), or entirely covered with dense, white, branched setae (males). T1 discal patch transversely diamond-shaped; T2 with LLB forming acute angle with ATB. Female: pseudopygidial area basally elongate into tear-shaped region of uniformly golden reflectance; S5 strongly downcurved. Male: Pygidial plate keyhole shaped, lacking distinct transverse basal ridge, but with weakly downturned apical plane; S2 with apical band of white setae; S3–S5 with apical fringes of setae (white on S3–S4, brown on S5).

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 79 Comments.—This species is well characterized by the unique pseudopygidial area and the unusually- shaped T1 discal patch, along with its relatively small size. Distribution.—MéXICO: Durango; USA: Arizona, New Mexico. Rozen (1989a) also examined specimens of this species from Texas. Host Records.—Xenoglossa eriocarpi (Cockerell) [= Tetraloniella (Tetraloniella) eriocarpi (Cockerell)] (1 female specimen pinned with host specimen, apparently taken from nest entrance, Cochise Co., Arizona; Rozen, 1989a, adult parasite-host interactions at nest entrance, egg and larvae from cells; Rozen, 1989b, larva from cell); Dieunomia (Epinomia) nevadensis (Cresson) (1 specimen from nesting area, Cochise Co., Ari- zona). Floral Records.—Aster tanacetifolius [= Machaeranthera tanacetifolia (H.B.K.) Nees], Haplopappus sp., Helianthus petiolaris Nutt., Heterotheca sp., Sida abutifolia P. Mill. Seasonal Records.—14 August to 9 October. Specimens examined.—86 &, 1 % (DAVIS, LAWRENCE, LOS ANGELES, NEW YORK, RIVERSIDE, TEMPE, TUCSON).

TRIEPEOLUS LUNATUS (Say) (Figs. 110–113, 256)

Epeolus lunatus Say 1825: 8516 [%, &; Missouri and Prairie des Chiens] [Type lost]. Epeolus lunatus concolor Robertson 1898: 51 [Lectotype: Illinois Natural History Survey No. 8174; &, Carlinville, Macoupin Co., Illinois; 24 July 1888]; Webb 1980: 108 [lectotype designation (by W. E. LaBerge)]. Triepeolus lunatus; Robertson 1901: 231. Triepeolus concolor; Robertson 1903: 285. Triepeolus nautlanus Cockerell 1904: 36 [Holotype: U. S. National Museum of Natural History No. 9705; %, (vicinity of San Rafael, Rio Nautla, Vera Cruz, México)]. new synonymy Triepeolus lunatus lunatus; Mitchell 1962: 472, Fig. 112 [redescription, illustration of scutellum, axillae].

Description.—Length ca. 9–13 mm (rarely as small as 7.5 mm); ITW 2.0–2.7 mm (rarely as small as 1.6 mm). Integument black, sometimes with the following red: basal half of mandible, labrum, apical clypeus, pronotal lobe, tegula, axillar spine, and/or legs (red especially in southern locales, blacker especially in specimens from Illinois, Kansas, and neighboring states); dorsum of mesosoma and metasoma with bands of pale yellow setae (deeper yellow in some specimens from México), usually with banding becoming progressively paler yellow on more posterior terga. Clypeus with strong or rarely weak midline; with weak larger punctures; often sparsely (females) or densely (males) covered with medially-directed, white setae. Paramedian band clearly separated from other pale setae on mesoscutum (somewhat reduced in Midwestern specimens). Scutellum moderately to strongly bigibbous; axillar spine surpassing or rarely only reaching midpoint of scutellum, apical tip slightly incurved, sometimes with reddish coloration. Mesepisternum lacking erect, simple setae or with relatively short (ca. 0.5 OD or less), suberect, simple setae; females with dense, pale yellow, branched setae below scrobal groove, usually also between pronotal lobe and hypoepimeron, and sometimes

16. The original description of Epeolus lunatus Say has been cited by Dalla Torre (1896) and Hurd (1979) as being from the year 1824, page 354. I have seen a microfiche reproduction of the original book in which Say’s description is found. The book, the second volume of Keating’s account of the Long Expedition, contains a series of appendi- ces, the page numbers of which start at p. 1, despite their appearance after p. 248 of the main text, with the addition of two plates of figures at the beginning. I suspect that Dalla Torre continued counting the pages of the appendix from the main text, thus arriving at p. 354 for the original description of E. lunatus. It is unclear if the book that I saw was a later printing of an 1824 edition of this work, or if Dalla Torre and subsequent authors simply mis-cited the date of the publication; however, I can find no indication that the 1825 book is a second printing or new edition to a previous work.

80 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER on anterior surface of mesepisternum (Southwestern specimens); males with more evenly dispersed, sparse, pale yellow, branched setae; punctation usually fairly dense, separated by up to one or two puncture diameter in some places. T1 discal patch strongly triangular to trapezoidal; T2 with LLB absent (esp. Midwest) or present (esp. Southwest), forming acute or weakly acute angle with ATB. Female: Pseudopygidial area semicircular to subquadrate, with distinct, flat, basal crescent; S2–S4 with apical or apicolateral bands of pale setae; S5 very slightly downcurved. Male: Pygidial plate of moderate size, keyhole shaped, with strong basal transverse ridge; S2–S3 with white apical bands of appressed setae (often slightly surpassing apical margin of S3); S4–S5 with brown apical fringes. Comments.—Although the type specimen of this species is no longer available for study, the meaning is clear from the original description and well understood by most workers making determinations in the collections that I have examined. Based on the reduced, triangular T1 discal patch, males of this species might be confused with T. q. atlanticus (separated by the shorter pronotal collar in T. lunatus—only ca. 1 OD in length in T. lunatus, versus the ca. 2 OD length of the pronotal collar in T. q. atlanticus), T. simplex (separated by the apical fringes on both S4 and S5 in T. lunatus, only on S4 in T. simplex), or T. concavus, T. nevadensis, or T. remigatus (separated by the isolated paramedian bands on the mesoscutum in T. lunatus). This species is extremely similar to T. rufithorax; the only notable difference between the two species is the greater amount of red coloration on the thorax in T. rufithorax. One specimen of T. lunatus from Chiapas, México, was labeled “PCAM 40” by D. Yanega; another from Sonora, México, was labeled “PCAM 3” by T. Griswold. Distribution.—MéXICO: Chiapas, Chihuahua, Coahuila, Durango, San Luis Potosí, Sonora, Veracruz, Zacatecas; USA: Alabama, Arizona, Arkansas, California (Modoc Co.), Colorado, Connecticut, Delaware, Florida, Georgia, Illinois, Indiana, Iowa, Kansas, Louisiana, Maryland, Michigan, Mississippi, Missouri, Nebraska, New Jersey, New Mexico, New York, North Carolina, Ohio, Pennsylvania, South Carolina, South Dakota, Tennessee, Texas, Utah, Virginia, Washington D.C., West Virginia, Wisconsin. Host Records.—Melissodes (Melissodes) bimaculata (Lepeletier)? (Mitchell 1962, evidence not given; Hurd et al. 1980, evidence not given; John S. Ascher, in lit., 2003, 2006, observation of nests in numerous localities); Melissodes sp. (1 specimen, emerging from host nest, Champaign, Illinois). Floral Records.—Aplopappus spinulosus [= Machaeranthera pinnatifida (Hook.) Shinners ssp. pinnatifida var. pinnatifida], Asclepias syriaca L., A. tuberosa L., Aster pilosus [= Symphyotrichum pilosum (Willd.) Nesom var. pilosum], A. tanacetifolium [= Machaeranthera tanacetifolia (H.B.K.) Nees], Baccharis sp., Bahia absinthifolia var. dealbata (Gray) Gray, Baileya pleniradiata Harvey & Gray ex Gray, Bidens alba (L.) DC. var. radiata (Schultz-Bip.) Ballard ex T.E. Melchert, Boltonia sp., Brauneria pallida [= Echinacea pallida (Nutt.) Nutt.], Callirhoe sp., Campanulastrum americanum (L.) Small, Ceanothus sp., Cirsium sp., Core- opsis major Walt., Chrysopsis camporum [= Heterotheca camporum (Greene) Shinners var. camporum], Cyrilla parvifolia Raf., Eriogonum deflexum Torr., Eupatorium purpureum L., Euphorbia marginata Pursh, Gaillardia pulchella Foug., Grindelia squarrosa (Pursh) Dunal, Helenium tenuifolium [= Helenium amarum (Raf.) H. Rock var. amarum], Helianthus annuus L., Helianthus atrorubens L., H. petiolaris Nutt., H. tuberosus L., Heliopsis helianthoides (L.) Sweet, Heterotheca subaxillaris (Lam.) Britt. & Rusby, Kallstroemia sp., Koellia flexuosa (= Pycnanthemum tenuifolium Schrad.), Lobelia siphilitica L., Lygodesmia juncea (Pursh) D. Don ex Hook., Medicago sp., Melilotus officinalis (L.) Lam., Monarda punctata L., Opuntia sp., Petaloste- mon occidentalis [= Dalea candida Michx. ex Willd. var. oligophylla (Torr.) Shinners], Platycodon grandiflo- rus (Jacq.) A. DC., Ratibida pinnata (Vent.) Barnh., Rudbeckia hirta L., R. laciniata L., Silphium perfoliatum L., Solidago sp., Spermacoce verticillata L., Sphaeralcea sp., Verbena brasiliensis Vell., V. hastata L., V. stricta Vent., Vernonia baldwinii interior (Small) Faust, Ve. fasciculata Michx., Veronicastrum virginicum (L.) Farw., cantaloupe (= Cucumis melo L.), China aster [= Callistephus chinensis (L.) Nees], cotton (= Gossypium sp.), red clover (= Trifolium pratense L.), smartweed (= Polygonum), soybean (= Glycine).

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 81 Seasonal Records.—20 March to 7 October. Specimens examined.—639 &, 313 % (ANN ARBOR, AUSTIN, BERKELEY, BOULDER, DAVIS, GAINES- VILLE, ITHACA, LAWRENCE, LOGAN, LOS ANGELES, NEW YORK, RIVERSIDE, SAN FRANCISCO, STARKVILLE, TEMPE, URBANA, WASHINGTON D.C.).

TRIEPEOLUS MARGARETAE new species (Figs. 114, 115)

Diagnosis.—This species can be distinguished from all others except T. rohweri and T. claytoni by the very unusual pseudopygidial area, which is entirely clothed with uniformly fine, dense setae, the basal half silvery and apical half pale golden However, in T. margaretae the apical margin is more truncate than in T. claytoni, giving the pseudopygidial area a quadrate appearance, while in T. claytoni the apical margin is rounded, giving it a more elongated, ovate shape; also in T. margaretae the line separating the basal, silvery setae and the apical, golden setae is indented medially, while in T. claytoni this line is straight. Triepeolus margaretae can be differentiated from T. rohweri by the presence of erect, simple setae on the mesepisternum in T. rohweri, by the large, trapezoidal T1 discal patch shape in T. margaretae, and the diffuse setae on the anterior margin of the mesoscutum in T. margaretae. Description of holotype female.—Length ca. 10.5 mm (10–11 mm); ITW 2.3 mm (2.2–2.3 mm). Integ- ument black, with orange on basal half of mandible, labrum, and apical margin of clypeus; orange-brown on scape, pedicel and F1; reddish brown on pronotal lobe, tegula, and legs excluding coxae and spurs. Clypeus with very faint midline. Paramedian band long, poorly distinguished from diffuse apical setae. Scutellum moderately (to strongly) bigibbous; axillar spine triangular, not attaining midpoint of scutellum. Mesepister- num lacking erect, simple setae (but with suberect, branched setae near pronotal lobe and tegula, especially in male), dorsally with rather sparse, white, branched setae; punctures small, weakly impressed, separated by 1 to 2 puncture widths (occasionally up to four puncture diameters ventrally; integument shining). T1 discal patch widely quadrate, BTB and ATB slightly interrupted medially; T2 with LLB mostly on lateral surface of T2, forming acute angle with ATB. Pseudopygidial area subquadrate, entirely composed of fine, long setae; basal half to third with setae forming inverted V-shaped area of silvery reflectance; apically with setae slightly coarser, reflecting golden; S5 straight in profile, slightly convex along longitudinal midline. (Male: Pygidial plate red, strongly keyhole shaped, with faint basal transverse ridge; S2–S3 with apical bands of white setae, slightly surpassing apical margin on S3; S4 with white and golden brown, and S5 with brown apical fringes.) Distribution.—USA: Texas (Goliad Co., Blanco Co.). Floral Records.—Gaillardia sp. Seasonal Records.—7 May to 15 May. Holotype.—“16 mi E. Goliad, Tex. [Texas], V [May]-7-1953 // Taken on Gaillardia // M. G. Rightmyer Triepeolus Database Specimen No. 1143 // HOLOTYPE & Triepeolus margaretae Rightmyer 2007” (LAWRENCE). Paratypes.—USA: TEXAS: Blanco Co., Pedernales Falls State Park, 15 May 1994, A. W. Hook (1%, AUSTIN); (Goliad Co.), Goliad, 8 May 1953, L. D. Beamer, Gaillardia sp. (1&, LAWRENCE). Etymology.—This species is named in honor of my aunt, Margaret V. Depue, for her guidance and care throughout my life.

TRIEPEOLUS MARTINI (Cockerell) (Figs. 116, 117, 247, 257)

Epeolus remigatus var. martini Cockerell 1900: 362 [Holotype: U. S. National Museum of Natural History No. 18965; &, Romeroville (San Miguel Co.), New Mexico; 6 August 1899].

82 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Triepeolus fortis Cockerell 1921: 3 [Holotype: American Museum of Natural History No. 25082; &, (cottonwood area at head of Dry Willow Creek), Wray (Yuma Co.), Colorado; about 40° 0’N, 102° 10’W; 3700 ft; (18) August 1919]. new synonymy Triepeolus trilobatus Cockerell 1921: 7–8 [Holotype: American Museum of Natural History No. 25088; %, White Rock, near Boulder (Boulder Co.), Colorado; about 40° 3’N 105° 8’W; 5200 ft; 13 August 1919]. new synonymy Triepeolus (Synepeolus) insolitus Cockerell 1921: 6 [Holotype: American Museum of Natural History No. 25086; %, Pueblo (Pueblo Co.), Colorado; about 38° 10’N 104° 36’W; 4700 ft; 9 August 1920]. new synonymy Triepeolus martini; Hurd 1979: 2094.

Description.—Length ca. 12–16 mm; ITW 2.6–3.1 mm. Integument black, with red on basal half of mandible and F1, sometimes with red to orange on labrum, scape, pedicel, and part or most of legs (excluding coxae and spurs); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with strong midline and weak larger punctures, sometimes covered with sparse, brown setae. Paramedian band strongly demarcated, joined laterally to apicolateral setae, forming anchor shape. Scutellum strongly bigibbous; axillar spine surpassing midpoint of scutellum, slightly incurved. Mesepisternum lacking erect, simple setae; with dense, pale, branched setae between hypoepimeron and pronotal lobe (sometimes also beneath scrobal groove); punctures rough, nearly contiguous, integument between punctures raised, somewhat tuberculate. T1 discal patch triangular to subrectangular; T2 with LLB forming weakly acute angle with ATB. Mesosoma and metasoma venter entirely black. Female: Pseudopygidial area poorly differentiated from rest of T5, formed by flattened, triangular plane of coarser, dark setae, with golden reflectance along apical margin; T5 lacking lateral patch of white setae (rarely with small patch of sparse white setae lateral to pseudopygidial area); S5 straight in profile. Male: Pygidial plate relatively wide, keyhole shaped, with distinct basal transverse ridge and moderately downturned apical plate; S2–S3 mostly dark brown, with faint bands of white setae on apical margins; S4–S5 with apical fringes of dark brown setae. Comments.—This species was placed in the subgenus Synepeolus by Cockerell, based on the presence of only two submarginal cells in the holotype specimen; however, this is a variable character within the species. Females of T. martini are easily recognizable by the distinctive, dark, perfectly triangular pseudopygidial area. Distribution.—MéXICO: Chihuahua; USA: Arizona, Colorado, Kansas, Nebraska, New Mexico, Texas (Gaines Co.). Host Records.—Nomia bakeri Cockerell [= Dieunomia (Epinomia) nevadensis bakeri (Cockerell)] (1 specimen, noted by C. Hicks as “flying over ground where many nest, especially Nomia bakeri,” from Colo- rado). Floral Records.—Gutierrezia sarothrae (Pursh) Britt. & Rusby, Helianthus annuus L., H. petiolaris Nutt., Melilotus alba [= Melilotus officinalis (L.) Lam.], Prionopsis ciliata [= Grindelia papposa Nesom & Suh]. Seasonal Records.—17 July to 17 September. Specimens examined.—56 &, 3 % (BOULDER, DAVIS, LAWRENCE, LOS ANGELES, NEW YORK, NEW YORK-ASCHER, SAN DIEGO, URBANA, WASHINGTON D.C.).

TRIEPEOLUS MAUROPYGUS new species (Figs. 118, 119)

Diagnosis.—This species can be distinguished by the combination of the relatively short clypeus; the very short and sparse (easily overlooked and therefore apparently absent) erect, simple setae on the mesepisternum; the transversely ovate to widely trapezoidal T1 discal patch; the transverse bands of T1–T4, which are medially interrupted or nearly so; the presence of pale setae lateral to the pseudopygidial area; and the almost entirely dark setae of the pseudopygidial area, combined with the presence of a weakly shining basal crescent of silvery setae.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 83 Triepeolus mauropygus is similar to T. laticeps and T. tepanecus (in particular, a large specimen from Sonora strongly resembles T. laticeps); however, in T. mauropygus the axillar spine is smaller than that of T. tepanecus; the T1 discal patch is more distinctly ovate (as opposed to usually subtriangular in both T. laticeps and T. tepanecus), and the apical transverse bands of pale setae on T2–T3 are interrupted medially (uninterrupted medially on T3 and sometimes T2 in T. laticeps). See Table 2 for further characteristics of these species. Description of holotype female.—Length ca. 10 mm (8–10 mm); ITW 1.8 mm (1.6–1.8 mm). Integu- ment black, with red on apical half of mandible; brownish orange on F1 (sometimes on entire anterior surface of antenna, when directed dorsally) and tegula (darker in some specimens); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint (to dorsally well-defined) midline and distinct larger punctures. Paramedian band clearly separated from other pale setae on mesoscutum. Scutellum weakly bigibbous; axillar spine triangular, almost reaching midpoint of scutellum. Mesepisternum apparently lacking (but with very short and sparse) erect, simple setae; dorsal third, except hypoepimeron, covered with dense, pale yellow, branched setae, faintly extending along anterior margin of mesocoxa, lacking on anterior surface of mesepisternum; ventral integument shining, with punctures nearly contiguous to separated by one puncture diameter. T1 widely ovate (to trapezoidal), with ATB and BTB interrupted medially; T2 with LLB mostly on lateral surface of T2, forming very weakly acute angle with ATB (or rarely absent). Pseudopygidial area subquadrate (to semicircular), with setae nearly uniformly glossy, dark, and fine except for vague basal crescent of slightly denser, finer, shorter setae; S5 straight in profile. (Male: Unknown.) Distribution.—MéXICO: Sinaloa, Sonora; USA: Arizona. Floral Records.—“blue Malvaceae”. Seasonal Records.—26 August to 28 November. Holotype.—“ARIZONA – Cochise Co. 25 mi NE Douglas, 26 August 1969, T. Hsiao, W. J. Hanson // UTAH STATE UNIV Intermountain Insect Survey // Native Bee Survey USDA Logan, Utah BBSL581802 // M. G. Rightmyer Triepeolus Database Specimen No. 1268 // HOLOTYPE & Triepeolus mauropygus Right- myer 2007” (LOGAN). Paratypes.—MÉXICO: SINALOA: 10 mi W Concordia, 30 October 1965, G. E. & A. S. Bohart (2&, LOGAN); 20 mi N Los Mochis, 9 October 1968, G. E. Bohart (1&, LOGAN); small blue Malvaceae (2&, LOGAN); Mazatlán, 28 November 1962, T. Gantenbein (1&, LOS ANGELES); USA: ARIZONA: Cochise Co., 25 mi NE Douglas, 26 August 1969, T. Hsiao, W. J. Hanson (3&, LOGAN). Etymology.—This name is in reference to the darkly shining pseudopygidial area found in this species and the closely related T. laticeps and T. tepanecus. From the Greek, “mauros” (dark) and “pygos” (rump).

TRIEPEOLUS MEDUSA Cockerell (Figs. 120, 121)

Triepeolus medusa Cockerell 1917b: 301–302 [Holotype: U. S. National Museum of Natural History No. 22895; &, Dis- trito Federal, México; “7 + 8.10” (7–8 October?)].

Description.—Length ca. 10.5–11.5 mm; ITW 2.2–2.7 mm. Integument black, with red on apical half of mandible; orange on most of middle and hind legs (excluding spurs), partially on front leg, antenna entirely brown; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint midline and larger punctures, covered with dense white setae in males. Frons with erect, minutely-branched, pale setae (more noticeable in profile). Paramedian band clearly separated from other pale setae on mesoscutum (most females) or laterally contiguous with diffuse pale setae on anterior of mesoscutum. Scutellum moderately to weakly bigibbous; axillar spine triangular, not attaining midpoint of scutellum. Mesepisternum with very sparse, short, erect, simple setae; dorsal half, excluding hypoepimeron, with well-defined area of dense, yel-

84 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER low, branched setae; ventrally with sparser, black, branched setae; integument very densely, roughly punctate, punctures small and nearly contiguous (females), or mesepisternum covered with dense, pale, branched setae (sparser on hypoepimeron; males). T1 widely trapezoidal to ovate; T2 with LLB forming acute angle with ATB. Female: Pseudopygidial area semicircular, with basal region of strongly shining, silvery, fine setae, gradually grading to coarser, less dense apical setae; venter of metasoma dark brown except for patches of white setae laterally on S4–S5, extended medially, sparser on S5, and sometimes sparse medially on S3; S5 straight in profile. Male: Pygidial plate keyhole shaped, with distinct basal transverse ridge and apical downturned plate; S2–S3 with apical bands of white setae; S4–S5 with apical fringes of brown and white setae. Comments.—This species is similar to T. townsendi, but T. medusa has longer erect, simple setae on the mesepisternum and the pseudopygidial area has a vague basal shining crescent. Triepeolus medusa is also similar to T. laticeps, but the ventral half of the lateral surface of the mesepisternum is not as shiny as in that species, and the clypeus has only a faint midline. Finally, T. medusa is similar to T. fulgidus, but the setae of the pseudopygidial area are not as uniform as in that species, there are more terga that have the apical transverse bands of pale yellow setae medially interrupted, and the erect setae on the frons are more prevalent in T. medusa. Distribution.—MéXICO: Guanajuato, Hidalgo, Jalisco, México, Nayarit, Sonora, Distrito Federal; USA: New Mexico. Floral Records.—Bidens pilosa L., Cosmos sp. Heterotheca subaxillaris (Lam.) Britt. & Rusby. Seasonal Records.—12 July to 25 October. Specimens examined.—8 &, 3 % (LAWRENCE, LOGAN, SAN FRANCISCO, WASHINGTON D.C.).

TRIEPEOLUS MELANARIUS new species (Figs. 122, 123)

Diagnosis.—This species can be distinguished from other species in the genus by the pseudopygidial area, which is remarkably similar to that of Epeolus. In overall appearance, it is almost identical to T. heterurus, and is often collected at the same localities; however females of T. heterurus and T. melanarius can easily be distinguished by the pseudopygidial area, which is parted medially and apically concave in T. heterurus, and totally unlike that of Epeolus. Males of the two species are extremely similar, but can be differentiated by the apical bands of white setae on S2 and S3 in T. heterurus (metasomal sterna entirely brown in T. melanarius). In addition, in males, the ventral half of the lateral surface of the mesepisternum is darker in T. melanarius than in T. heterurus, and the paramedian bands are clearly separated from other pale setae on the mesoscutum in T. melanarius, while in T. heterurus they are usually surrounded by diffuse pale setae on the anterior margin of the mesoscutum. Description of holotype female.—Length ca. 8.5 mm (7–8.5 mm); ITW 1.9 mm (1.5–2.0 mm). Integu- ment black, with red on apical half of mandible; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus lacking midline and larger punctures (sometimes covered with sparse, white setae). Paramedian band clearly separated from other pale setae on mesoscutum, reaching anterior margin, slightly curving outwards anteriorly (sometimes appearing straight; rarely absent, possibly due to wear?). Scutellum weakly bigibbous; axillar spine triangular, not attaining midpoint of scutellum. Mesepisternum with dorsal third covered with white, branched, suberect setae; anterior margin with long, erect to suberect, brown, branched and simple setae; ventrally with punctures nearly contiguous, covered with sparse, brown, branched setae (denser in males). T1 discal patch subtriangular; T2 with LLB forming acute angle with ATB. Pseudopygidial area subquadrate, poorly differentiated from rest of T5 (except for dark color contrasting with white setae laterally on T5), apical margin with transverse patch shining, silvery setae; S5 straight in profile. (Male: Pygidial plate extremely narrow; metasomal sterna lacking white setae; S4–S5 with dark brown apical fringes.)

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 85 Comments.—A specimen of this species bears a manuscript name (authorship not given) relating to the locality Mariposa. Distribution.—USA: California. Floral Records.—Brassica nigra (L.) W.D.J. Koch, Cistanthe umbellata (Torr.) Hershkovitz. Grindelia camporum Greene, Hemizonia fasciculata (DC.) Torr. & Gray, He. wrightii (= He. kelloggii Greene), Marrubium vulgare L., Petroselinum crispum (P. Mill.) Nyman ex A.W. Hill, Salvia sp., Solidago sp. Seasonal Records.—24 May to 22 October. Holotype.—“Twain-Harte, Tuolumne Co, 4000 ft., Cal. [California], July 1937 // on Grindelia // Pres. By F. E. Blaisdell Collector // M. G. Rightmyer Triepeolus Database Specimen No. 6668 // HOLOTYPE & Trie- peolus melanarius Rightmyer 2007” [California Academy of Sciences #18247] (SAN FRANCISCO). Paratypes.—USA: CALIFORNIA: Alameda Co., 4 September 1956 (1&, 2%, URBANA); Berkeley, 16 June 1952, R. H. & L. D. Beamer et al. (1&, LAWRENCE); U. C. Berkeley campus, 6 August 1910 (1%, WASH- INGTON D.C.); 13 August 1910, J. C. Bridwell (3%, WASHINGTON D.C.); Berkeley Hills, 9 August 1908, J. C. Bridwell (3%, WASHINGTON D.C.); Piedmont, 19 June 1912, F. W. Nunenmacher (12&, WASHINGTON D.C.); 27 June 1912, F. W. Nunenmacher (1&, WASHINGTON D.C.); Mariposa Co., 5.3 mi NE Bagby, 30 July 1953, Salvia sp. (1&, WASHINGTON D.C.); Mariposa, 17 June 1968, R. M. Bohart (1%, DAVIS); Merced Co., 6 mi NE Ballico, 17 June 1970, R. R. Snelling, Grindelia (10&, 1%, LOS ANGELES); Merced Falls, 30 July 1953, Grindelia camporum (1&, WASHINGTON D.C.); 15 mi S Merced, 12 August 1960, R. R. Snelling (3&, 12%, LOS ANGELES); Santa Clara Co., Campbell, 6 June 1975, R. W. Ribble, Brassica nigra (1&, URBANA); 17 August 1974, R. W. Ribble, Peteoselinum erispum (1&, URBANA); Milpitas, 14 June 1956, D. Ribble (2&, 7%, URBANA); 25 June 1956, D. Ribble (10&, 6%, URBANA); 23 July 1956, D. Ribble (2&, 1%, URBANA); San Antonio Valley, 27 June 1953, G. A. Marsh (1%, LOS ANGELES); Stanford University, July 1910 (1%, WASH- INGTON D.C.); 8 July 1957, P. H. Arnaud (1%, SAN FRANCISCO); August 1917, Thompson (1&, WASHINGTON D.C.); 30 August 1953, P. H. Arnaud (1&, NEW YORK); Stanislaus Co., Del Puerto Canyon, 8 September 1956, R. R. Snelling, (1&, LOS ANGELES); La Grange, 28 May 1961, R. R. & M. D. Snelling, Marrubium vulgare (1%, LOS ANGELES); Salado Creek Canyon, 4 October 1956, R. R. Snelling (1%, WASHINGTON D.C.); 10 mi SW Turlock, 1 September 1961, R. R. Snelling (1&, LOS ANGELES); Tuolumne Co., June 1912, J. G. Grundel (1&, SAN FRANCISCO); Twain-Harte, July 1937, F. E. Blaisdell, 4000 ft, Grindelia sp. (6&, 4%, SAN FRANCISCO). Additional specimens.—USA: CALIFORNIA: Alameda Co., hills in back of Oakland, 2 August 1908, E. C. VanDyke (1&, WASHINGTON D.C.); Butte Co., Eliot Springs, 750, Neal Road, 5.1 mi NE junction Hwy 99, 21 June 1979, R. R. Snelling, P. Melhop (1%, LOS ANGELES); Calaveras Co., 5 km W Copperopolis, 24 June 1963, P. H. Arnaud (2&, SAN FRANCISCO); Contra Costa Co., Lafayette, 2 July 2001, J. S. Ascher (1&, NEW YORK-ASCHER); Orinda, 10 June 1934, B. E. White (1&, SAN FRANCISCO); El Dorado Co., Kyburz, 13–21 June 1981, S. C. Kuba (1&, SAN FRANCISCO); Fresno Co., Little Panoche Road, 1 mi W of I- 5, 3 June 2001, J. S. Ascher (3%, NEW YORK-ASCHER); Glenn Co., Plaskett Meadows, 14 June 1972, J. Doyen, 6300 ft, Calyptridium umbellatum (1&, BERKELEY); Los Angeles Co., Santa Monica, 30 June 1935, E. G. Linsley (1%, BERKELEY); Taubark Flat, 2 June 1976, B. Villegas (1&, DAVIS); Westwood Hills, 27 July 1955, E. G. Linsley (1&, BERKELEY); Madera Co., 5 May 1942, E. G. Linsley (6%, BERKELEY); 5 June 1942, E. G. Linsley (1%, 1& BERKELEY); Mendocino Co., Eel River Camp, 11 July 1948, U. N. Lanham (1&, BOUL- DER); Napa Co., Pope Valley, 1/3 km S Aetna Springs, 3–14 October 1993, L. S. Kimsey (1&, DAVIS); Nevada Co., Grass Valley, 10 July 1954, E. C. Zimmerman (2&, SAN FRANCISCO); Riverside Co., 10 October 1927, Timberlake, Hemizonia wrightii (1&, BERKELEY); 11 October 1927, Timberlake, Hemizonia wrightii (1&, BERKELEY); 12 October 1927, Timberlake, Hemizonia wrightii (1&, BERKELEY); Sierra Co., Sattley, 22 June 1984, R. M. Bohart (1&, DAVIS); Solano Co., Benecia, 14 August 1910, J. C. Bridwell (1%, WASHING- TON D.C.); Mix Canyon, 4 April 1975, R. J. Skalaski (1&, DAVIS); (San Bernardino Co.), Redlands, 31 July 1908, H. A. Scullen (1%, SAN FRANCISCO); San Diego Co., 1 mi E Del Mar, 17 July 1963, J. C. Hall (1&, RIV-

86 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER ERSIDE); 5 mi S Del Mar, 10 July 1963, E. I. Schlinger, Hemizonia sp. (1&, RIVERSIDE); Julian, 19 June 1962, J. A. Froebe (1&, DAVIS); La Mesa, 22 October 1954, F. X. Williams (1&, SAN FRANCISCO); La Jolla, 14 August 1911, J. C. Bridwell (4%, WASHINGTON D.C.); Oceanside, 17 August 1946, Timberlake (1&, BERKE- LEY); Grindelia alata (2&, BERKELEY); Hemizonia fasciculata (1&, BERKELEY); (San Francisco Co.), Glen Canyon, Santa Cruz Mountains, 16 August 1935, Timberlake, Solidago (1%, BERKELEY); San Mateo Co., Redwood City, 8 July 1961, R. R. Snelling (1&, LOS ANGELES); Santa Cruz Co., 1 July 1955 (1&, URBANA); Loma Prieta Road, 11 June 1960, D. W. Ribble (1%, LAWRENCE; 1%, URBANA); Shasta Co., 2 mi W Shingle- town, 5–10 August 1985, R. Miller, 2750 ft (2&, GAINESVILLE); Tulare Co., Badger, 29 June 1957, R. R. Snelling, 3000 ft (1&, 1%, LOS ANGELES); Lindsay, 19 June 1929, Timberlake, Michaelmas daisy (1%, BERKE- LEY; 1%, WASHINGTON D.C.); Yolo Co., Davis, J. J. duBois (1&, LOS ANGELES); June 1926, J. J. duBois (1%, WASHINGTON D.C.); 2 September 1956, A. T. McClay (1%, DAVIS). Etymology.—I follow Timberlake’s suggested manuscript name for this species, as indicated on a specimen from Riverside County. This name is appropriate, as the metasomal sterna and mesepisternum of these bees are darker than those of T. heterurus, which they resemble. From the Greek, “melanos” (black) and “arius” (pertaining to).

TRIEPEOLUS MICHENERI new species (Figs. 124–126)

Diagnosis.—This species can be distinguished by the combination of the distinctive ventral half of the mesepisternum, which lacks erect, simple setae and which is intensely shining, almost entirely glabrous, and usually has punctures separated by 4 to 5 puncture diameters; the axillar spines, which are pointed, slightly incurved apically, and surpass the scutellar midpoint; the ATB of T1, which is medially interrupted by at least 1 OD; and the acute angle formed by the LLB and ATB on T2. Specimens from the more southern locales (i.e., Jalisco and Oaxaca) tend to have bands of orange setae, while those of Sinaloa and Sonora have bands of rather pale yellow setae. Description of holotype female.—Length ca. 10 mm (9–10 mm); ITW 2.1 mm (2.1–2.2 mm). Integu- ment black, with red on apical half of mandible; orange on F1, tegula, and middle and hind legs distal to coxae (sometimes black on tegula and entire legs); dorsum of mesosoma and metasoma with bands of lemon yellow setae (ranging from orange-yellow to pale yellow). Clypeus with elevated midline and faint larger punctures laterally, partially obscured by sparse white setae (these sparse setae absent in some specimens). Paramedian band clearly separated from other pale setae on mesoscutum. Scutellum moderately (to weakly) bigibbous; axillar spine relatively long and pointed, surpassing midpoint of scutellum. Mesepisternum lacking erect, simple setae; dorsal half with well-defined region of dense, pale yellow (or white), branched setae, absent (or less dense) on hypoepimeron; ventrally with very shining integument, punctures minute, spaced by as much as 5 to 7 puncture diameters (as much as ca. ten puncture diameters apart in some specimens). T1 discal patch subtriangular (to subovate or trapezoidal); T2 with LLB forming acute angle with ATB. Pseudopygidial area subquadrate (to semicircular), with distinct, narrow, basal silvery crescent; S5 straight in profile. (Male: Unknown.) Distribution.—MéXICO: Jalisco, Oaxaca, Sinaloa, Sonora. Seasonal Records.—19 June to 28 October. Holotype.—“Villa Guadalupe, Jalisco, México, 26 July 1951 // H. E. Evans Collector // M. G. Rightmyer Triepeolus Database Specimen No. 1260 // HOLOTYPE & Triepeolus micheneri Rightmyer 2007” (LAWRENCE). Paratypes.—MÉXICO: JALISCO: 8 km W. Tequila, 18 July 1951, H. E. Evans (1&, LAWRENCE); Chapala, 19 June 1963, Scullen, Bolinger (1&, CORVALLIS); OAXACA: 23 mi S Matías Romero, 14 August

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 87 1963, F. D. Parker, L. A. Stange (2&, DAVIS); SINALOA: Choix, Rio Choix, 19 July 1968, T. A. Sears, R. C. Garner, C. S. Glaser (1&, DAVIS); Mazatlán, 28 October 1965, G. E. & A. S. Bohart (2&, LOGAN); SONORA: Navajo, 27 September 1966, G. E. & A. S. Bohart (1&, LOGAN). Etymology.—This species is named in honor of my doctoral advisor, Charles D. Michener, for his inspiring mentorship and kind friendship.

TRIEPEOLUS MICHIGANENSIS Mitchell (Figs. 127, 128)

Triepeolus michiganensis Mitchell 1962: 473–474 [Holotype: U. S. National Museum of Natural History No.75244; &, Shiwassee Co., Michigan; 3 September 1950].

Description.—Length ca. 8–9 mm; ITW 1.9–2.0 mm. Integument black, with red to orange on apical mandible, labrum, entirely or basally on antenna, and partially on legs, sometimes on tegula, rarely on pronotal lobe; dorsum of mesosoma and metasoma with bands of pale gray to white setae. Clypeus with distinct midline and vague larger punctures, asetose or covered with medially-directed white setae (denser on basal half, especially in males). Paramedian band clearly separated from other pale setae on mesoscutum (females) or connected laterally to areas of diffuse setae on anterior margin of mesoscutum (males). Scutellum moderately to strongly bigibbous; axillar spine triangular, not incurved apically, reaching scutellar midpoint. Mesepister- num lacking erect, simple setae; punctures nearly contiguous to separated by one puncture diameter; with white, branched setae mostly restricted to dorsal half to fourth of lateral surface (absent on hypoepimeron) as well as anterior surface (males with setae denser). T1 discal patch widely trapezoidal; T2 with LLB forming acute angle with ATB; T1–T3 with ATBs interrupted medially. Female: Pseudopygidial area subquadrate, with distinct basal crescent of silvery shining setae; S2–S4 with white appressed setae on apical margin; S5 very slightly downcurved apically or straight. Male: Pygidial plate of moderate size with slight emargination on lateral margins and weak or strong basal transverse ridge; S2–S4 with white setae on apical margins; S4– S5 with brown apical fringes (white laterally on S4). Comments.—The males of this species are similar to T. obliteratus, but T. obliteratus lacks the midline on the clypeus, and the apical transverse setae on T1–2 are only slightly interrupted medially (in T. michiganensis, the setae are well-separated medially on T1–2, and slightly interrupted medially on T3). Females are recognized by the combination of their small size, bands of white setae on the metasoma, and the distinct basal crescent of shining setae on the pseudopygidial area. Distribution.—USA: Michigan, New Hampshire, New York. Host Records.—Melissodes (Eumelissodes) denticulata Smith (John S. Ascher, in lit., 2003, observation of nesting site). Floral Records.—Solidago sp. Seasonal Records.—26 July to 3 September. Specimens examined.—5 &, 2 % (ITHACA, NEW YORK-ASCHER, PHILADELPHIA, WASHINGTON D.C.).

TRIEPEOLUS MICROPYGIUS Robertson (Figs. 129, 130)

Triepeolus micropygius Robertson 1903: 286 [Lectotype: Illinois Natural History Survey No. 23531; &, Carlinville, Macoupin Co., Illinois; 29 September 1902]; Webb 1980: 110 [lectotype designation (by W. E. LaBerge)]. Triepeolus micropygius micropygius; Mitchell 1962: 474–475.

88 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Description.—Length ca. 10–12 mm (rarely as small as 8 mm); ITW 1.9–2.4 mm (rarely as small as 1.4 mm). Integument black, with red apical half of mandible; sometimes with orange labrum and clypeus, usually with orange F1, tegula, and legs (black in specimen from Mississippi); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint midline (stronger dorsally) and faint larger punctures, clypeus sparsely covered with white setae (sometimes asetose in more eastern specimens). Paramedian band narrow, clearly separated from other pale setae on mesoscutum (lectotype specimen and a few others) or joined laterally to appressed setae on anterior margin of mesoscutum. Scutellum weakly bigibbous; axillar spine triangular, reaching midpoint of scutellum. Mesepisternum with relatively short, erect, simple setae (ca. 0.25 OD); usually mostly covered with pale yellow, appressed, branched setae, this setae sparser ventrally (lectotype specimen mostly lacking appressed setae ventrally); punctures dense and small, ventrally nearly contiguous to separated by one puncture diameter (up to three puncture diameters in lectotype). T1 discal patch transversely ovate to widely subrectangular; T2 with LLB forming weakly to strongly acute angle with ATB, this angle correlated with overall width of tergal bands [some specimens’ banding enlarged by diffuse pale setae, and others (including lectotype) with banding rather narrow (reminiscent of T. helianthi)]. Female: Pseudopygid- ial area (and T5 in general) narrowed apically, basally with dense, silvery, pilose setae, this silvery setae extending laterally towards apical margin, partially enclosing small, rounded apical region of sparse, coarse setae (apical margin straight in specimen from Mississippi); S2–S4 with apical bands white setae (rarely only on S2–S3 or S3–S4); S5 distinctly downturned, also narrowed apically. Male: Unknown. Comments.—This species resembles T. fraserae due to the small, rounded pseudopygidial area with the distinct basal region of silvery shining setae; however the S5 and T5 of T. micropygius are conspicuously narrowed distally, S5 is much more strongly downcurved, the pseudopygidial area is even more distinctly rounded, and the erect, simple setae of the mesepisternum are much shorter in T. micropygius (0.25 OD vs. greater than 0.5 OD in T. fraserae). The rounded pseudopygidial area and the downcurved S5 resemble those of the T. verbesinae species group; however the small size of the pseudopygidial area and the general banding pattern of the metasoma (i.e., often the T1 transverse bands of pale setae are medially interrupted, and the T2 LLB forms only a weakly acute angle with the ATB—although in some exceptionally setose specimens this angle is strongly acute) differentiate this species from the T. verbesinae group. The distribution of this species is largely in the Western U. S., with only one known specimen from Illi- nois (the lectotype specimen). The lectotype specimen is less setose than many, but not all, of the other examined specimens. In addition, I have seen one specimen from Mississippi (repository: Starkville); this specimen agrees with T. micropygius in most characters, except it has black legs and tegula, and is also quite sparsely setose. Distribution.—USA: Arizona, California (Mono Co.), Colorado, Idaho, Illinois (Macoupin Co.), Missis- sippi (Oktibbeha Co.), Montana, Nebraska, Nevada, New Mexico, Oregon, Utah. Floral Records.—Aster tanacetifolius [= Machaeranthera tanacetifolia (H.B.K.) Nees], Chrysothamnus sp., Grindelia squarrosa (Pursh) Dunal, Melilotus sp., Oligoneuron rigidum (L.) Small, Solidago sp. Seasonal Records.—19 August to 23 October. Specimens examined.—45 & (BOULDER, CORVALLIS, DAVIS, LAWRENCE, LOGAN, NEW YORK, SAN FRANCISCO, STARKVILLE, URBANA, WASHINGTON D.C.).

TRIEPEOLUS MITCHELLI Hurd (Fig. 131)

Triepeolus sublunatus Mitchell 1962: 483–484 [nec Cockerell] [Holotype: U. S. National Museum of Natural History (on indefinite loan from North Carolina State University); %, Marion (McDowell Co.), North Carolina; 27 July 1924; Coreopsis stelista]. Triepeolus mitchelli Hurd 1979: 2094 [replacement name].

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 89 Description.—Length ca. 11 mm; ITW 1.9 mm. Integument black; dorsum of mesosoma and metasoma with bands of pale gray to white setae. Clypeus with strong midline, mostly covered with white setae. Paramedian band barely contiguous with diffuse, weak area of white setae on anterior margin of mesoscutum. Scutellum moderately bigibbous; axillar spine triangular, reaching scutellar midpoint. Mesepisternum lacking erect, simple setae; with dense, white, branched setae on dorsal third (excluding hypoepimeron), less dense on anterior surface of mesepisternum, relatively sparse on rest of lateral surface; with punctures nearly contiguous to spaced up to three puncture diameters apart, integument between relatively flat. T1 discal patch relatively small, subtriangular/subovate. T2 with LLB forming 90 degree angle with ATB. Female: Unknown. Male: Pygidial plate of moderate size, with distinct basal transverse ridge; S2–S3 with apical bands of white setae; S4–S5 with brown apical fringes plus white setae laterally on S4. Comments.—This species is very similar to T. georgicus Mitchell, due to the white setal banding on the black body, the clypeus of normal length with strong midline, the mesepisternum lacking erect, simple setae and the shining integument which is moderately punctate and smooth (unlike T. atripes Mitchell, which it also superficially resembles). It differs from T. georgicus by the broader metasomal banding and the triangular T1 discal patch; however this is based on the male specimen, and it is not particularly rare to find male specimens with greatly expanded areas of pale setae on the metasoma in Triepeolus. Additionally, the scutellum is not nearly as biconvex in T. mitchelli as in T. georgicus, although this character also can also vary within a species. Of note is the wide spacing of the mesepisternal punctures in T. mitchelli, which is rare among male specimens of Triepeolus species (particularly those with pale gray to white banding). More specimens of T. georgicus and/or T. mitchelli, especially from more northerly localities, may provide further evidence on whether the species should be synonymized. Distribution.—USA: North Carolina. Floral Records.—Coreopsis major stellata [= Coreopsis major Walt.]. Seasonal Records.—27 July. Specimens examined.—1 % (WASHINGTON D.C.).

TRIEPEOLUS MOJAVENSIS Linsley (Figs. 132, 133, 258)

Triepeolus mojavensis Linsley 1939: 2–4 [Holotype: California Academy of Sciences No. 4801; &, 1 mi N Deep Creek (Mojave Desert near Hesperia, San Bernardino Co.) California; 26 April 1936].

Description.—Length ca. 10–12 mm; ITW 2.2–2.8 mm. Integument black, with red on apical half of mandible; dorsum of mesosoma and metasoma with bands of white setae. Preoccipital carina very reduced. Clypeus with weak midline dorsally, lacking larger punctures, mostly asetose except for few long, brown setae. Head and mesosoma prominently covered with erect, simple setae. Paramedian band poorly discernable, diffuse, joined with diffuse, white setae on anterior mesoscutum. Scutellum moderately bigibbous; axillar spine rounded, not attaining midpoint of scutellum. Mesepisternum densely covered with dark, long, erect, simple setae; punctures beneath not distinct, integument furrowed (rugose). T1 almost entirely covered by white setae except for vague black oval mediobasally (BTB present); rest of terga with ATBs uninterrupted; T2 with LLB absent, but ATB slightly widened laterally, resulting in vague, black, semicircular region at base of tergum. Female: Pseudopygidial area subrectangular, with apical, transverse row of stout setae, basally with fine black setae, poorly differentiated from rest of T5; S5 straight in profile. Mesosoma and metasoma venters entirely black. Male: Unknown. Comments.—This species is very similar to T. dacotensis, but the BTB on T1 (present in T. mojavensis and absent in T. dacotensis) and the punctation of the mesepisternum are different. Distribution.—USA: California (San Bernardino Co., Riverside Co.).

90 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Host Records.—Anthophora linsleyi Timberlake (Linsley and MacSwain 1942, adults observed explor- ing burrows, and presumably nests were excavated to obtain parasitism rate in a population near Bakersfield, California). Seasonal Records.—26 April to 31 May. Specimens examined.—3 & (SAN FRANCISCO, WASHINGTON D.C.).

TRIEPEOLUS MONARDAE Mitchell (Figs. 134, 135)

Triepeolus monardae Mitchell 1962: 475, Fig. 112, [Holotype: U. S. National Museum of Natural History No. 75246; %, Wilmington (New Hanover Co.), North Carolina; 12 September 1932].

Description.—Length ca. 15–18 mm; ITW 2.9–3.5 mm. Integument black, with red on medially on mandible; brown on outer F1; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with strong midline and weak larger punctures, densely covered with white setae in males. Paramedian band clearly separated from other pale setae on mesoscutum in both sexes. Scutellum strongly bigibbous; axillar spine strongly pointed, weakly incurved, surpassing midpoint of scutellum. Mesepisternum with short, erect, simple setae; punctures nearly contiguous to separated by up to 0.5 puncture diameter; integument between punctures raised; with dense, white, branched setae on dorsal third (sparser on hypoepimeron in females). T1 discal patch widely rectangular/trapezoidal; T2 with LLB forming acute angle with ATB. Female: Pseudopy- gidial area subovate, with uniformly shining, dark, “glossy” setae; S2–S3 (sometimes also S4) with white setae on apicolateral margins; S5 not downcurved. Male: Pygidial plate relatively wide, keyhole shaped, with distinct basal transverse ridge; S2–S3 with white apical bands of setae; S4–S5 with apical fringes setae brown (paler brown with white laterally on S4, sometimes with small area of white setae on S5). Comments.—This species is distinctive for its large size and pattern of pale setal bands on the metasoma. Distribution.—USA: Florida, Georgia, North Carolina. Floral Records.—Monarda punctata L., white goldenrod (= Solidago bicolor L.). Seasonal Records.—28 August to 12 September. Specimens examined.—3 &, 3 % (RALEIGH, NEW YORK, WASHINGTON D.C.).

TRIEPEOLUS NAYARITENSIS new species (Figs. 136, 137)

Diagnosis.—This species can be distinguished from all other species of Triepeolus except T. concavus and T. penicilliferus by the pseudopygidial area, which has long, erect setae forming a rounded, three-dimensional structure similar to that of T. concavus, and the strongly downcurved female S5. It can be distinguished from T. concavus and T. penicilliferus by the erect, simple setae on the mesepisternum (lacking in those species). T. nayaritensis is also similar to T. flavigradus, but in T. flavigradus the paramedian bands are poorly distinguishable from other pale setae on the mesoscutum (clearly isolated in T. nayaritensis), the upper face has erect setae such that clypeus appears recessed (such facial setae absent in T. nayaritensis), and the metasomal band coloration grades from yellow to white in T. flavigradus (evenly pale yellow in T. nayaritensis). Description of holotype female.—Length ca. 12 mm (10.5–12 mm); ITW 2.4 mm (2.1–2.5 mm). Integ- ument black to brown, with red medially on mandible; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint (to distinct) midline, larger punctures faint (or covered with appressed white setae in male); clypeus in lateral view slightly convex, mirroring slope of face. Paramedian band relatively narrow, clearly separated from other pale setae on mesoscutum. Scutellum very weakly bigibbous, somewhat

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 91 extended posteriorly; axillar spine triangular, rounded apically, not attaining midpoint of scutellum. Mesepis- ternum with erect, simple setae (sparser and shorter than that of T. flavigradus); dorsally and along posterior margin with dense, white, appressed, branched setae (also along anterior surface and ventrally in males); punctures moderately small, in places separated by 2–3 puncture diameter or less (more densely punctate in males). T1 discal patch widely trapezoidal, with BTB and ATB interrupted medially; T2 with LLB forming acute angle with ATB. Pseudopygidial area similar to that of T. concavus, but setae less distinctly forming posterior plate, and basal setae more erect than that of T. flavigradus; S5 strongly downcurved. (Male: Pygid- ial plate relatively wide, with transverse basal ridge often hidden beneath long setae and downturned apical plate; S2 with apical band of white setae; S3 with white apical setae slightly extended beyond apical margin; S4–S5 with dark brown apical fringes, sometimes with white laterally.) Comments.—All of the known specimens of this species were obtained from one collection event (see distributional and seasonal records, below). Distribution.—MéXICO: Nayarit. Seasonal Records.—28 September. Holotype.—“Méx. [México] Nay. [Nayarit]: (mi. 75157.7) 29 km N. Peñitas, 90 m. elev. 28 Sept. [Sep- tember] 1976, C. D. George & R. R. Snelling, colls.// M. G. Rightmyer Triepeolus Database No. 6968 // HOLOTYPE & Triepeolus nayaritensis Rightmyer 2007” (WASHINGTON D.C.). Paratypes.—Label data for all paratypes exactly as in holotype (13 &, 10 %, WASHINGTON D.C.). Etymology.—This species is named after the collection locality, the state of Nayarit, México.

TRIEPEOLUS NEVADENSIS (Cresson) (Figs. 138–140)

Epeolus nevadensis Cresson 1878: 86 [Lectotype: Academy of Natural Sciences No. 2220; &, Nevada]; Cresson 1916: 125 [lectotype designation]. Triepeolus nevadensis; Robertson 1901: 231.

Description.—Length ca. 12.5–14 mm; ITW 2.6–2.9 mm. Integument black, with red on basal half of mandible; sometimes with orange labrum and F1; dorsum of mesosoma and metasoma with bands of yellow setae. Clypeus shining, mostly asetose, lacking midline, with large punctures. Preoccipital carinae on gena and posterior margin of head behind vertex. Mesoscutum and scutellum shining; mesoscutum with dense band of setae on anterior fourth, with paramedian band barely distinct from dense anterior setae. Scutellum flattened and somewhat extended posteriorly; axillar spine not attaining midpoint of scutellum, rounded apically. Mesepisternum lacking erect, simple setae; with distinct area of yellow, branched setae on dorsal half to third (including hypoepimeron); ventrally with brown, branched setae; punctures small and contiguous. T1 discal patch distinctly rectangular. T2 with LLB forming conspicuous right angle with ATB. Mesosoma and metasoma venter entirely dark brown (except males with white apical band of setae on S3). Female: Pseudopygid- ial area uniformly dark, “glossy”; poorly differentiated from rest of T5 (T5 usually lacking pale setae, but sometimes with pale setae lateral to pseudopygidial area), S5 not downcurved. Male: Pygidial plate of moderate size, keyhole shaped, with strong basal transverse ridge, forming downturned apical plate; S3 with white apical band of setae (sometimes brown medioapically), setae medially weakly surpassing apical margin; S4– S5 with brown apical fringes. Comments.—Triepeolus nevadensis is superficially similar to T. concavus, but can be separated by the following characters of T. nevadensis: the clypeus is shining with distinct larger punctures, the mesoscutum is shining, the paramedian bands are isolated from the yellow setae on the anterior margin of the mesoscutum (though weakly developed), the scutellum is flattened and somewhat extended posteriorly, the T1 discal patch is strongly rectangular, and the T2 LLB forms a 90 degree angle with the ATB. The female pseudopygidial

92 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER area of T. nevadensis is completely different from that of T. concavus, appearing poorly differentiated from the rest of T5, darkly shining, flat, and quadrangular; also, S5 is not downcurved. Triepeolus nevadensis is also similar to T. remigatus, but in T. nevadensis the paramedian bands are not so strongly developed as in T. remigatus, the mesoscutum is more distinctly shining, the scutellum is more flattened and extended posteriorly, and the T1 discal patch is distinctly rectangular (as opposed to ovate or subtriangular in T. remigatus). The males have white banding only on S3 (as opposed to both S2 and S3). The female pseudopygidial areas are similar, but the species can be differentiated by the other characters listed above. Triepeolus nevadensis is also similar to T. robustus in overall appearance, but can be distinguished from that species by the lack of dense, erect setae on the upper face, which, in T. robustus, causes the clypeus to appear to be recessed. Distribution.—MéXICO: Chihuahua, Durango; USA: Georgia, Nevada, New Mexico, North Carolina, Oklahoma, Texas. Floral Records.—Monarda citriodora Cerv. ex Lag., M. punctata L., Vernonia glauca (L.) Willd. Seasonal Records.—3 May to 27 September. Specimens examined.—21 &, 2 % (BOULDER, DAVIS, LAWRENCE, NEW YORK, PHILADELPHIA, RALEIGH, SAN FRANCISCO, WASHINGTON D.C.).

TRIEPEOLUS NIGRIHIRTUS Mitchell (Figs. 141–144)

Triepeolus nigrihirtus Mitchell 1962: 476–477 [Holotype: U. S. National Museum of Natural History No. 400195 (on indefinite loan from North Carolina State University); %, Merry Oaks (Clatham Co.), North Carolina; 27 May 1926].

Description.—Length ca. 10 mm; ITW 2.3–2.5 mm. Integument black (& specimen) or reddish brown (% HT, possibly due to preservation?); tegula with transparent outer margin; dorsum of mesosoma and metasoma with bands of pale white setae. Clypeus with slight midline dorsally; shining with rather deeply impressed punctures (no larger punctures present); very sparsely setose in female, male with long white setae medioapically. Supraclypeal area between antennae rather swollen, produced laterally (especially in & specimen). Vertex with punctures small and distantly spaced, especially near lateral ocellus. Pronotal collar dorsally with long, sparse, erect setae directed laterally and parted at midpoint. Paramedian band apparently absent; anterior third to half of mesoscutum with diffuse, erect to suberect setae. Scutellum moderately bigibbous; axillar spine triangular, reaching midpoint of scutellum. Mesepisternum with long, minutely-branched, erect to semi-erect, white setae on anterior and posterior margins of mesepisternum, between pronotal lobe and hypoepimeron, and ventrally below hypoepimeron; integument shining, with small, deeply impressed punctures, separated by up to one puncture diameter. T1 discal patch transversely ovate, T2 with LLB forming acute angle with ATB (worn in & specimen). Female: Pseudopygidial area longitudinally ovate, basally well-defined from rest of T5, with dark, short setae; apically with tuft of longer, golden, shining setae; S2–S4 with white setae on apical margins; S5 faintly downcurved apically. Male: Pygidial plate keyhole shaped, with distinct basal transverse ridge; S2–S3 with white apical bands (medially curving basally on S3); S4–S5 with pale golden brown apical fringes. Comments.—Despite the disparate collection localities for the male holotype and the sole female specimen from Texas, I believe that these two specimens are conspecific based on the following unusual, shared morphological features: the diffuse, erect setae anteriorly on the mesoscutum, the carinate posterior margin of the head, and the shape of the T1 discal patch. Both specimens were collected relatively early in the year (late April and late May). Triepeolus nigrihirtus superficially resembles T. mojavensis and T. dacotensis in overall appearance, but can be separated from the latter two species by the strong preoccipital carina on the gena and posterior margin of the head, and by the completely different pseudopygidial area, which is unique among Triepeolus species.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 93 Distribution.—USA: North Carolina (holotype specimen), Texas (“Magnolia, April 30 1953, L D Beamer”) Floral Records.—(Texas specimen): Verbena officinalis L. Seasonal Records.—30 April to 27 May. Specimens examined.—1 &, 1 % (LAWRENCE, WASHINGTON D.C.).

TRIEPEOLUS NISIBONENSIS Genaro (Figs. 145, 146)

Triepeolus nisibonensis Genaro 2001: 1033–1034, Figs. 18–20 [Holotype: Florida State Collection of Arthropods; %, Nisibón, La Altagracia, Dominican Republic; 5–7 May 1978; near cacao planting; flight trap].

Description.—Length ca. 8.5–11.5 mm; ITW 2.1–2.2 mm. Integument black, with dark red medially on mandible; orange brown on F1; dorsum of mesosoma and metasoma with bands of yellow to white setae. Clypeus with faint midline (covered with pale yellow to white setae). Posterior margin of head at vertex with distinct medial notch. Paramedian band clearly separated from other pale setae on mesoscutum , well separated from anterior margin in white-banded specimen, or tapering to anterior margin in yellow-banded (i.e., holotype) specimen. Scutellum strongly bigibbous; axillar spine strongly pointed and incurved, apical point reaching or slightly exceeding posterior margin of scutellum. Mesepisternum lacking erect, simple setae; with very distinct circular area of dark, branched setae surrounded by pale yellow to white, branched setae; punctation mostly obscured by setae but apparently nearly contiguous. T1 discal patch very widely subovate (apical bands suggestive of T. verbesinae); T2 with LLB forming weakly acute angle with ATB. Female: Unknown. Male: Pygidial plate narrow and long, with strong basal transverse ridge and apical downturned plate; S2–S3 with apical bands of white setae; S4–S5 with apical fringes of setae white on S4, brown on S5 (sometimes with white laterally). Comments.—This species can be differentiated from the other species of Triepeolus by the notched dorsal margin of the head. Distribution.—DOMINICAN REPUBLIC: La Altagracia. Seasonal Records.—5 to 7 May. Specimens examined.—2 % (LAWRENCE, GAINESVILLE).

TRIEPEOLUS NORAE Cockerell (Figs. 147, 148)

Triepeolus norae Cockerell 1907b: 59 [Holotype: U. S. National Museum of Natural History No. 100034; &, Mesilla Park (Dona Ana Co.), New Mexico; 16 May; Sphaeralcea lobata]; Linsley 1962: 150, 152, 161 [biological data on sleeping aggregations]; Hurd et al. 1980: 131 [Table of flight times].

Description.—Length ca. 7–11 mm; ITW 1.6–2.2 mm. Integument black, with dark reddish brown to orange on basal half of mandible, part or entire labrum, apical margin of clypeus, part or entire scape, pedicel, and F1, rarely on pronotal lobe; orange on tegula and legs (except basally on coxae and usually spurs); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus shining, with glabrous midline present and distinct larger punctures. Paramedian band clearly separated or barely contiguous with pale setae on anterior margin of mesoscutum. Scutellum strongly to moderately bigibbous; axillar spine reaching midpoint of scutellum, triangular, usually with apical point directed inward. Mesepisternum with sparse, short, erect, simple setae; mostly covered with appressed, white, branched setae, but medioventrally with circular area (asetose or covered with diffuse, black, branched setae); punctures nearly contiguous to separated by one puncture

94 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER diameter in some areas, integument between punctures raised, tuberculate. T1 discal patch transversely ovate; T2 with LLB forming strongly acute angle with ATB. Female: Pseudopygidial area subtriangular, with two distinct regions usually of approximately equal size (dorsally with shining silvery setae, ventrally with coarse, longer setae); S5 straight in profile or faintly downcurved apically. Male: Unknown. Comments.—Triepeolus norae is distinguishable by its pseudopygidial area, which has a relatively large basal region of silvery reflectance, the apical margin of which is nearly straight in most specimens, causing this entire basal region to have a somewhat triangular appearance (as opposed to the typical crescent-shaped basal region of silvery setae in many Triepeolus species). This species was labeled “PCAM 20” by T. Griswold. Distribution.—México: Chihuahua, Coahuila, Durango, Zacatecas; USA: Arizona, California (Riverside Co.), Nevada, New Mexico, Texas (Armstrong Co., and counties west of and including Brewster Co. and Cul- berson Co.), Utah. Floral Records.—Acacia greggii Gray, Aplopappus gracilis [= Machaeranthera gracilis (Nutt.) Shin- ners], Aster spinosus [= Chloracantha spinosa (Benth.) Nesom], Baileya multiradiata Harv., Dyssodia aurea [= Thymophylla aurea (Gray) Greene ex Britt. var. aurea], Eriogonum sp., Haplopappus sp., Helianthella sp., Larrea tridentata (Sessé & Moc. ex DC.) Coville, Lepachys tagetes [= Ratibida tagetes (James) Barnh.], Melilotus alba [= Melilotus officinalis (L.) Lam.], Pectis angustifolia Torr., Poliomintha incana (Torr.) Gray, Prosopis juliflora (Sw.) DC., Psorothamnus scoparius (Gray) Rydb., Sida sp., Sphaeralcea ambigua Gray, S. angustifolia (Cav.) G. Don. Seasonal Records.—15 April to 21 October. Specimens examined.—158 & (AUSTIN, BERKELEY, CHAMELA, CORVALLIS, DAVIS, GAINESVILLE, ITH- ACA, LAWRENCE, LOGAN, LOS ANGELES, MEXICO CITY, NEW YORK, RIVERSIDE, SAN FRANCISCO, TEMPE, URBANA, WASHINGTON D.C.).

TRIEPEOLUS OCCIDENTALIS (Cresson) (Figs. 149, 150, 259)

Epeolus occidentalis Cresson 1878: 87–88 [Lectotype: Academy of Natural Sciences No. 2224; &, Colorado]; Cresson 1916: 126 [lectotype designation]. Triepeolus occidentalis; Cockerell 1904: 38.

Description.—Length ca. 9.5–13 mm; ITW 1.8–2.8 mm. Integument black, with dark reddish brown to orange on basal half of mandible, part or entire labrum, and F1, sometimes on apical margin of clypeus, scape, and pronotal lobe; orange on tegula and legs (except basally on coxae and spurs); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint midline and faint large punctures. Paramedian band long, clearly separated from other pale setae on mesoscutum. Scutellum moderately bigibbous; axillar spine triangular, surpassing scutellum midpoint, with apical point directed inwardly. Mesepisternum with very short, sparse, erect, simple setae; with dense, white, branched setae on upper third (lacking or sparse on hypoepimeron) and sometimes along margins ventrally; entire ventral half or medioventral area asetose or covered with sparse, brown, branched setae; with contiguous, minute punctures. T1 discal patch transversely ovate, with BTB and ATB not or little interrupted medially; T2 LLB forming acute angle with ATB. Female: Pseudopygidial area semicircular to subtriangular with large amount of basal and lateral shining silvery setae, mostly enclosing median region of coarse setae; shining setae mediobasally often with slight notch; T5 lateral white setae extending basally, surrounding on either side a black median band basal to pseudopygidial area; S2–S4 with white setae (sometimes faint) at apex; S5 slightly downcurved at apex. Male: Unknown. Comments.—This species is similar to T. circumculus; the two can be differentiated by the pseudopygidial area, in which the basal shining setae is less distinctly differentiated from the apical coarse setae in T. occi-

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 95 dentalis; the mesepisternum, which is more densely punctate in T. occidentalis; and by the axillar spine, which is more robust in T. occidentalis. Distribution.—USA: Colorado, Idaho, Kansas, North Dakota. Host Records.—Melissodes mizeae Cockerell [= Melissodes (Eumelissodes) menuachus Cresson]? (Hicks, 1926, two females observed entering and spending time in nests of host). Floral Records.—Grindelia squarrosa (Pursh) Dunal. Seasonal Records.—21 August to October (day unspecified). Specimens examined.—25 & (BOULDER, ITHACA, LOGAN, NEW YORK, WASHINGTON D.C.).

TRIEPEOLUS PAENEPECTORALIS Viereck (Figs. 151–154)

Triepeolus paenepectoralis Viereck 1905: 278–280 [Holotype: Academy of Natural Sciences No. 10124; &, Vancouver, British Columbia), Canada]. Triepeolus amandus Cockerell 1921: 10 [Holotype: American Museum of Natural History No. 25092; %, Meeker (Rio Blanco Co.), Colorado; about 40° 2’N 107° 55’W; 6200 ft; (21) July 1919; (Grindelia serrulata)]. new synonymy Triepeolus alpestris Cockerell 1921: 13 [Holotype: American Museum of Natural History No. 25097; &, Leadville (Lake Co.), Colorado; about 39° 15’N 106° 16’W; 10,300 ft; (4) August 1919; (Lepidium alyssoides)]. new synonymy Triepeolus vandykei Cockerell and Sandhouse 1924: 307 [Holotype: California Academy of Sciences No. 1599; &, Mill- brae (San Mateo Co.), California; 1 September 1912]. new synonymy

Description.—Length ca. 7–12 mm; ITW 1.4–2.6 mm. Integument black, with dark red on apical half of mandible; dark reddish brown rarely on legs; orange on F1; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint to absent midline, with larger punctures visible if not covered with brown setae; or entirely covered with white setae in males. Paramedian band separated from or barely contiguous with lateral setae. Scutellum weakly bigibbous, slightly extended posteriorly; axillar spine triangular, not attaining midpoint of scutellum. Mesepisternum with erect, simple setae (sometimes sparse and short); dorsal half covered with pale yellow, branched setae (sparser or with brown, branched setae on hypoepimeron); ventrally with punctures relatively small, dense (separated by less than one puncture diameter in some specimens), integument shining, sometimes medioventrally with short, relatively dense brown setae interspersed with golden, short, suberect setae (females), or entirely covered with white setae in males. T1 discal patch rectangular, BTB and ATB not or only slightly interrupted medially; T2 with LLB forming 90 degree angle with ATB. Female: Pseudopygidial area semicircular with distinct shining basal crescent, extending apically along lateral margins to partially enclose circular region of stout setae; S5 not or very slightly downcurved. Male: Pygidial plate triangular, with indistinct transverse basal ridge and apical downturned plate; S2–S3 with apical bands of white setae (see comments, below); S4 with brown or white, and S5 with brown, apical fringes of setae. Comments.—Triepeolus paenepectoralis is a variable species, encompassing a number of intergrading forms. The most widespread form is smaller in size, resembling T. subalpinus in overall appearance (ca. 1.4– 2.0 mm ITW), while a couple of other forms are larger, approaching the general appearance of T. texanus (ca. 1.8–2.6 mm ITW). Of the two larger forms, one is most abundantly found in California, and has relatively broad bands of yellow setae on the metasoma; this form corresponds to the synonymized name T. vandykei. The other larger form is found mostly in Oregon and Washington, and has slightly narrower bands of yellow setae on the metasoma. This species is very similar to T. eldoradensis and T. argyreus; Table 1 gives distinguishing features of each species. Also, T. paenepectoralis is similar to T. subalpinus and T. texanus. Females of T. subalpinus are easily differentiated from those of T. paenepectoralis by their small, triangular, silvery pseudopygidial area. Triepeolus texanus can be distinguished from T. paenepectoralis by the clypeus, which is elongate and has a relatively strong midline in T. texanus (clypeus relatively short in T. paenepectoralis).

96 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER The description of the male is based on one specimen from Colorado (the holotype of T. amandus) and 12 males from Morongo, California, and caution should be taken before applying this description to the males of T. paenepectoralis as a whole. Timberlake placed a manuscript name on a specimen of this species in reference to the relatively strong genal carina. Distribution.—CANADA: Alberta, British Columbia; USA: California (Siskiyou Co. and Lassen Co. south to San Diego Co. and Riverside Co.), Colorado, Idaho, Nevada, Oregon, Utah, Washington, Wyoming. Host Records.—Melissodes (Eumelissodes) microsticta Cockerell? (2 specimens at nest site, Kitsap Co., Washington). Floral Records.—Chrysothamnus sp., Cirsium lanceolatus [= Cirsium vulgare (Savi) Ten.], C. undulatum (Nutt.) Spreng. var. undulatum, Eriogonum fasciculatum Benth., Grindelia squarrosa (Pursh) Dunal, Gutierrezia lucida [= Gutierrezia microcephala (DC.) Gray], Haplopappus sp., Hemizonia paniculata Gray, Monardella sp., Solidago spectabilis (D.C. Eat.) Gray var. confinis (Gray) Cronq., “tall cone flower”. Seasonal Records.—31 May to 12 October (most commonly July through September). Specimens examined.—207 &, 13 % (BERKELEY, BERLIN, BOULDER, CORVALLIS, DAVIS, GAINESVILLE, LAWRENCE, LOGAN, LOS ANGELES, PHILADELPHIA, NEW YORK, RIVERSIDE, SAN FRANCISCO, TUCSON, URBANA, WASHINGTON D.C., ZILLAH).

TRIEPEOLUS PARKERI new species (Figs. 155, 156)

Diagnosis.—This species primarily can be distinguished from all other species of the genus except T. interruptus by the absence of the BTB on T1 and can be distinguished from that species by the uninterrupted ATB on T1 (medially interrupted in T. interruptus). In addition, the species usually can be differentiated due to the presence of paramedian bands on the mesoscutum in T. parkeri (absent in T. interruptus). Other diagnostic characters of T. parkeri include the strong midline and relatively short length of the clypeus. Description of holotype female.—Length ca. 7.5 mm (7–8.5 mm); ITW 1.5 mm (1.4–1.8 mm). Integu- ment black, with red on apical half of mandible, orange on F1–5 (sometimes only on F1); dorsum of mesosoma and metasoma with bands of yellow-orange setae. Clypeus with strong midline and faint larger punctures (covered with dense white setae in males). Paramedian band present but very narrow and short, clearly separated from other pale setae on mesoscutum . Scutellum weakly bigibbous; axillar spine triangular, not (or nearly) reaching midpoint of scutellum. Mesepisternum with dense but very short, erect, simple setae (possibly overlooked without careful scrutiny); with punctures small, nearly contiguous; with small dorsal patch of pale yellow, branched setae below scrobal groove (males with denser branched setae covering much of mesepisternum, white dorsally and anteriorly, remainder brown). T1 lacking BTB (sometimes very faintly present), with LLB sparse and thin but present on basolateral corner; T1–T4 each with straight, uninterrupted ATB; T2 with LLB absent. Pseudopygidial area subquadrate, with very faintly differentiated basal crescent of silvery setae, mostly composed of relatively long, coarse, golden setae; S5 straight in profile. (Male: Pygidial plate keyhole shaped, with distinct transverse basal ridge and apical downturned plate; S2–S3 with white apical bands, S3 white setae slightly surpassing apical margin; S4–S5 with pale golden apical fringes.) Comments.—In addition to the specimens recorded below as paratypes, there are hundreds of additional specimens of this species collected by Frank Parker housed at the Wild Bee Lab in Logan. Distribution.—COSTA RICA: Guanacaste, San José. Seasonal Records.—18 November to 23 March. Holotype.—“COSTA RICA Guan. [Guanajuato] Finca Montezuma, 3 km SE Río Naranjo, 3–8 March 1992, F. Parker // USDA-ARS Bee Biol. & Syst. Lab, Logan Utah, Faunal Survey No. 000 026 317 // M. G.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 97 Rightmyer Triepeolus Database Specimen No. 3552 // HOLOTYPE & Triepeolus parkeri Rightmyer 2007” (LOGAN). Paratypes.—COSTA RICA: GUANAJUATO: Finca Montezuma, 3 km SE Río Naranjo, December 1991, F. Parker (6%, LOGAN), 18–22 January 1993, F. Parker (2%, LOGAN), 29 January–8 February 1993, F. Parker (4&, LOGAN), 19–24 February 1992, F. Parker (1&, LOGAN), 25 February–2 March 1992, F. Parker (1&, LOGAN), 1–15 March 1992, F. D. Parker (1&, LOGAN), 3–8 March 1992, F. Parker (1&, LOGAN); SAN JOSE: March 1923 (1&, BERLIN); 3 mi S Santa Ana, 28 December 1988, F. D. Parker (2&, LOGAN). Etymology.—This species is named in honor of Frank Parker, who, with the exception of nine specimens housed in the Berlin Museum, single-handedly collected the numerous specimens of this species.

TRIEPEOLUS PARTITUS new species (Figs. 157, 158)

Diagnosis.—This species can be distinguished by the combination of the clypeus, which is relatively short, flat, shining, and black, usually strongly contrasting with the red antennal scape and labrum; the pseudopygidial area, which is triangular to subquadrate, with a distinct, narrow, basal crescent of silvery shining setae; the BTB and ATB on T1, which are nearly parallel and surround a transversely ovate discal patch; and the very short and sparse (apparently lacking unless carefully examined) erect, simple setae of the mesepisternum. Triepeolus partitus resembles T. isohedrus, but in T. partitus the pseudopygidial area is not so distinctly triangular and the coarse apical setae are slightly longer, the T1 discal patch is more ovate, and the paramedian bands are not laterally contiguous with other pale yellow setae on the anterior margin of the mesoscutum. Description of holotype female.—Length ca. 12.5 mm (10–13.5 mm); ITW 2.7 mm (2.0–2.9 mm). Integument black, with bright reddish orange on basal half of mandible, labrum, apical margin of clypeus, scape, pedicel, F1, base of F2, tegula, and legs excluding basal fore- and midcoxae and spurs (legs sometimes black) (red coloration sometimes also medially on pronotal lobe and apically on axillar spine); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus relatively flat in profile, shining, with midline and larger punctures present. Paramedian band clearly separated from other pale setae on mesoscutum (in some specimens slightly curving outward on anterior margin). Scutellum strongly (to moderately) bigibbous; axillar spine triangular, reaching scutellar midpoint (often with pale yellow setae in inner lateral margin). Mesepisternum apparently lacking or with very sparse, short, erect, simple setae; dorsal half covered with pale yellow, branched setae, lacking (or brown) on hypoepimeron; ventrally with integument mostly asetose, shining, with punctures nearly contiguous separated by up to 1.5 puncture widths. T1 discal patch transversely ovate (to trapezoidal, rarely subtriangular), BTB and ATB interrupted medially; T2 with LLB mostly on lateral surface, forming acute angle with ATB. Pseudopygidial area longitudinally elongate, subquadrate (to triangular), mostly composed of narrow, suberect, relatively sparse setae, with narrow basal region of fine, silvery setae; S5 straight in profile. (Male: Unknown.) Comments.—This species was labeled “PCAM 7” by T. Griswold; it was also given the morphospecies number 6 by Paul Hurd. Timberlake gave a specimen of this species a manuscript name in reference to its collection locality in Arizona. Distribution.—MéXICO: Baja California Sur, Chihuahua, Durango, Zacatecas; USA: Arizona, New Mexico. Floral Records.—Eriogonum abertianum Torr., E. deflexum Torr., Gaillardia sp., Helianthus annuus L., Heterotheca subaxillaris (Lam.) Britt. & Rusby, Kallstroemia grandiflora Torr. ex Gray, Verbesina encelioides (Cav.) Benth. & Hook. f. ex Gray, Verbesina encelioides (Cav.) Benth. & Hook. f. ex Gray ssp. exauriculata (Robins. & Greenm.) J.R. Coleman. Seasonal Records.—4 August to 12 October.

98 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Holotype.—“AZ [Arizona]: Cochise Co., 5 mi S San Simon, VIII [August]-23-96 [1996], J. G. & B. L. Rozen, A. Pence // M. G. Rightmyer Triepeolus Database Specimen No. 3565 // HOLOTYPE & Triepeolus partitus Rightmyer 2007” (NEW YORK). Paratypes.—USA: ARIZONA: Cochise Co., Apache, 4 September 1986, F. Parker (2&, LOGAN); 13.3 m SW Apache, 31°32’28”N 109°16’30”W, 23 August 2003, J. G. Rozen (1&, NEW YORK); 18 mi SW Apache, 22–23 August 1994, J. G. Rozen, J. S. Ascher (2&, NEW YORK); 7 mi W Benson, 3 September 1961, Timber- lake, Heterotheca subaxillaris (2&, WASHINGTON D.C.); Chiricahua Mts., 27 August 1997, J. G. Rozen, B. McAdams, 7600 ft. (1&, NEW YORK); Dos Cabezas, 27 August 1974, R. M. Bohart (2&, DAVIS); Dos Cabezas Mts., 2 km N Fort Bowie, 3 September 1986, F. Parker (1&, LOGAN); Double Adobe, 23 August 1990, J. G. Rozen, J. Krieger (1&, NEW YORK); Douglas, 7 September 1975, E. S. Ross, 3900 ft. (1&, SAN FRANCISCO); 1 mi E Douglas, 20–22 August 1994, J. G. Rozen, J. S. Ascher (1&, NEW YORK); 28 August 1967, E. I. Schlinger (1&, RIVERSIDE); 29 August 1967, J. H. & J. M. Davidson, M. A. Cazier (3&, TEMPE); 1 September 1993, J. G. Rozen (1&, NEW YORK), 3 September 1992, J. G. & B. L. Rozen (1&, NEW YORK); 1 mi N Douglas 29 August 1967, J. G. Rozen, G. Krueger (1&, NEW YORK); 12 mi NW Douglas, 30 August 1989, J. G. & B. L. Rozen, R. Foster (1&, NEW YORK); 25 mi NE Douglas, 26 August 1969, W. J. Hanson, T. Hsiao (2&, LOGAN); Portal, 29 August 1967, E. I. Schlinger (1&, RIVERSIDE); 14 September 1980, Hanson, Knowlton (1&, LOGAN); near Portal, 30 August 1967, W. J. Hason (1&, LOGAN); 1 mi NE Portal, 29 August 1970, R. A. Men- dez (1&, NEW YORK); 2 mi E Portal, 12 September 1985, D. K. Broemeling (1&, LOGAN); 5 mi E Portal, 14 September 1955, C. & M. Cazier (1&, NEW YORK); G. E. Bohart, Verbesina encelioides (1&, LOGAN); 5 mi W Portal, 30 August 2003, J. S. Ascher (1&, NEW YORK); 5–7 mi SE Portal, 12 September 1978, G. Gordh (1&, RIVERSIDE); near mouth of Ramsay Canyon, 10 August 1940, Timberlake, Verbesina exauriculata (1&, BER- KELEY); Skeleton Canyon, 22 August 1962, J. G. Rozen, M. Statham, S. J. Hessel, Gaillardia (1&, NEW YORK); Skeleton Canyon, 6 mi SE Apache, 1 September 1958, P. D. Hurd, Eriogonum abertianum neomexicanum (1&, DAVIS); 2 mi SE Willcox, 22 August 2002, J. G. Rozen (2&, NEW YORK); 4 mi E Willcox, 28 August 1985, J. G. & B. L. Rozen (1&, NEW YORK); 1 mi S Jct. 181-666, 1 August 1961, M. A. Cazier (1&, NEW YORK). Additional specimens.—MÉXICO: BAJA CALIFORNIA SUR: Las Animas, Sierra Laguna, 12 October 1941, Ross, Bohart (2&, SAN FRANCISCO); CHIHUAHUA: Salaices, 20 August 1947, G. M. Bradt, 5200 ft. (1&, NEW YORK); DURANGO: Mapimí Reserva Biósfera, Presón Dos Amigos, 26°43’20”N 103°40’59”W, 28 September 1994, R. López, 1178 m. (1&, MEXICO CITY); Reserva Biósfera Mapimí near Ceballos, 23 August 1991, J. G. Rozen (1&, LAWRENCE); ZACATECAS: 5 mi N Tobasco, 18 September 1970, G. E. & R. M. Bohart (1&, LOGAN); USA: ARIZONA: Mojave Co., 5 mi N Kingman, Rt. 93, 11 Sep- tember 1976, R. C. Miller (1&, DAVIS); Pima Co., SE Continental, 31°46’98”N 110°53’18”W, 14 September 1994, T. Griswold (1&, LOGAN); W of Santa Rita Mountains, 31°47’26”N 110°49’28”W, 15 September 1994, T. Griswold (1&, LOGAN); 5 mi E Continental, Sta. Rita Mountains, 24 August 1950, T. Cohn, P. Boone, M. Cazier, 3500 ft. (1&, NEW YORK); 7 mi E Continental, 23 September 1962, J. Willcox (2&, RIVERSIDE); N of Madera Canyon, 31°47’91”N 110°53’34”W, 15 September 1994, T. Griswold, Kallstroemia (1&, LOGAN); Santa Cerrita Mts., 7 September 1963, V. L. Vesterby (1&, DAVIS); Santa Clara Co., 12 mi E Nogales, 14 September 1966, R. W. Thorp (1&, DAVIS); NEW MEXICO: Grant Co., 23 mi SW Silver City, 25 August 1969, W. J. Hanson, T. Hsiao (1&, LOGAN); Hidalgo Co., Animas, 4–5 August 1974, E. G. & J. M. Linsley, Helianthus annuus (2&, BERKELEY); 20 mi S Animas, 6–8 September 1977, R. J. McGinley (2&, WASHING- TON D.C.); 26 mi S Animas, 19 August 2001, J. G. Rozen (1&, NEW YORK); 13 September 1976, J. G. Rozen (1&, NEW YORK); 27–32 mi S. Animas, 24 August 1994, J. G. Rozen, J. S. Ascher (2&, NEW YORK); Lords- burg, 29 September 1960 (1&, LOGAN); Rodeo, 15 August 1974, E. G. & J. M. Linsley, Kallstroemia grandiflora (1&, BERKELEY); 10 September 1976, R. M. Bohart (1&, DAVIS); 1 mi N Rodeo, 19 August 1974, Rozen, Favreau (1&, NEW YORK); 25 August 1996, J. G. & B. L. Rozen, A. Pence (1&, NEW YORK); 2 mi N Rodeo, 20 September 1999, M. G. Rightmyer (1&, NEW YORK); 2 mi W Rodeo, 26 August 1982, R. M. Bohart, Eriogonum deflexum (1&, DAVIS); 20 mi S Rodeo, 28 August 1965, E. I. Schlinger (1&, RIVERSIDE).

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 99 Etymology.—This name is in reference to the paramedian bands, which are isolated from other bands of pale setae on the mesoscutum, and which help differentiate this species from T. isohedrus. From the Latin, “partitus” (divided).

TRIEPEOLUS PARVIDIVERSIPES new species (Figs. 159, 160)

Diagnosis.—This species can be distinguished from all other Triepeolus except T. diversipes by the special form of the pseudopygidial area, which is widely triangular and has dense, pilose, suberect, golden setae. Triepeolus diversipes and T. parvidiversipes can be distinguished by the mesepisternum (with much longer erect, simple setae and much more widely-spaced punctures in T. parvidiversipes) and by the much smaller size (ca. 7–8 mm) of T. parvidiversipes. Description of holotype female.—Length ca. 7 mm (7–8 mm); ITW 1.5 mm. Integument black, with reddish orange on basal half of mandible; orange on F1, tegula, and legs excluding basal halves of coxae and spurs; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus shining, lacking midline, with larger punctures present. Paramedian band somewhat diffuse, contiguous with lateral bands. Scutellum weakly bigibbous; axillar spine triangular, not (or nearly) reaching midpoint of scutellum. Mesepisternum with erect, simple setae; dorsal half with dense, white, branched setae, sparser on hypoepimeron; ventrally with sparse, brown, branched setae; punctures small, relatively dense and strongly impressed, punctures separated by up to one puncture diameter. T1 discal patch transversely ovate/subrectangular; T2 with LLB forming weakly acute angle with ATB. Pseudopygidial area triangular, with pilose, coarse setae, elongate on apical margin, and basal region with silvery setae; S5 straight in profile. (Male: Unknown.) Distribution.—USA: Arizona (Cochise Co.). Seasonal Records.—17 September to 21 September. Holotype.—“USA, Arizona: Cochise Co., Willcox, Sept. [September] 21, 1976, J. G. Rozen // M. G. Rightmyer Triepeolus Database Specimen No. 3563 // HOLOTYPE & Triepeolus parvidiversipes Rightmyer 2007” (NEW YORK). Paratype.—USA: ARIZONA: Cochise Co., 6 mi SE Willcox, 17 September 1967, S. A. Gerodenski, M. A. Cazier, on grass stems (TEMPE). Etymology.—This name is in reference to the smaller size of this species in comparison to T. diversipes, a species which it resembles. From the Latin, “parvus” (little), and the specific epithet diversipes, which is based on the Latin “diversus” (different), perhaps due to the diverging setae of the pseudopygidial area.

TRIEPEOLUS PARVUS new species (Figs. 161, 162)

Diagnosis.—This species can be identified by its small, widely triangular pseudopygidial area, which is mostly composed of long, coarse setae; and by the paramedian bands, which are laterally contiguous with pale setae on the anterior margin of the mesoscutum. Additional distinguishing features are the relatively short clypeus and the apparent absence of erect, simple setae on the mesepisternum (these setae observable under high magnification). Description of holotype female.—Length ca. 7 mm (7–8 mm); ITW 1.4 mm (1.4–1.9 mm). Integument black (to reddish brown, due to preservation?), with orange on basal half of mandible, margins of (or entire) labrum, apical margin of clypeus, scape, pedicel, F1, tegula, and legs excluding spurs (sometimes also medially on pronotal lobe) (sometimes orange coloration lacking on apical margin of clypeus, scape, pedicel, and foreleg); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus somewhat protuber-

100 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER ant, with faint midline and larger punctures, with sparse white setae directed medially (these setae sometimes lacking). Paramedian band laterally contiguous with pale band of setae on anterior margin of mesoscutum. Scutellum moderately (to weakly) bigibbous; axillar spine triangular, reaching midpoint of scutellum. Mesepisternum apparently lacking or with sparse, short, erect, simple setae under close observation; dorsal half with dense, white, branched setae, sparser on hypoepimeron; ventrally grading to sparse, white, branched setae; with punctures rather deeply impressed, nearly contiguous to separated by 0.5 puncture diameter. T1 discal patch transversely subovate (to rectangular); T2 with LLB forming acute angle with ATB. Pseudopy- gidial area widely triangular, mostly composed of coarse, long setae, basally with small area of silvery shining setae; S5 straight in profile. (Male: Unknown.) Distribution.—USA: Arizona, Colorado, New Mexico. Floral Records.—Asclepias subverticillata (A. Gray) Vail, Heterotheca sp. Seasonal Records.—15 July to 9 September. Holotype.—“USA, New Mexico: Hidalgo Co., Rodeo, IX [September]-9-1973 // M. Favreau Collector // M. G. Rightmyer Triepeolus Database Specimen No. 3581 // HOLOTYPE & Triepeolus parvus Rightmyer 2007” (NEW YORK). Paratypes.—USA: ARIZONA: Cochise Co., 3 mi SE Portal, 23 July 1959, M. Statham (1&, NEW YORK); COLORADO: (Boulder Co.), Boulder, 3 August 1935, C. D. Michener (1&, LAWRENCE); (Jeffer- son Co.), Golden, 15 July, D. Stoner (1&, WASHINGTON D.C.); NEW MEXICO: Hidalgo Co., 2 mi NE Rodeo, 8 August 1966, J. M. Davidson, M. A. Cazier, Asclepias subverticillata (1&, TEMPE); 7 mi SW Rodeo, 22 August 1964, J. G. & B. L. Rozen, Heterotheca (&, NEW YORK). Etymology.—This name is in reference to the small size of this bee species. From the Latin, “parvus” (little).

TRIEPEOLUS PECTORALIS (Robertson) (Figs. 163–166)

Epeolus pectoralis Robertson 1897: 345 [Lectotype Illinois Natural History Survey No. 15626; &, Carlinville, Macoupin Co., Illinois; 18 September 1893]; Webb 1980: 109 [lectotype designation (by W. E. LaBerge)]. Triepeolus pectoralis; Robertson 1901: 231; Mitchell 1962: 478–479, Fig. 112 [redescription, illustration of scutellum, axillae]. Epeolus virginiensis Cockerell 1907c: 137 [Holotype: U. S. National Museum of Natural History No. 40107; %, Falls Church, Virginia; 7 September]; Mitchell 1962: 478 [synonymy]. Triepeolus virginiensis; Brumley 1965: 73. Triepeolus heterurus; Hurd 1979: 2093. Epeolus oswegoensis Mitchell 1962: 453 [Holotype: U. S. National Museum of Natural History No. 75202; %, Oswego, New York; 26 August 1936]. new synonymy

Description.—Length ca. 8–11 mm; ITW 1.8–2.1 mm. Integument black to dark brown, with red basally on mandible; sometimes with orange on labrum, apical margin of clypeus, basally on antenna, tegula, and legs (at least in part); dorsum of mesosoma and metasoma with bands of pale yellow setae (setae, especially banding of metasoma, denser in specimens from Utah and Idaho). Clypeus with strong midline and weak larger punctures; with sparse (females) or dense (males) white, medially directed setae. Paramedian band clearly separated from other pale setae on mesoscutum (most females and some males), or contiguous with lateral setae (females from Southwest and some males). Scutellum moderately bigibbous; axillar spine triangular, reaching midpoint of scutellum. Mesepisternum with long, erect, simple setae; females dorsally with dense, white, branched setae on upper fourth, sparser on hypoepimeron; ventral three-fourths with punctation small and sparse (separated by up to 5–10 puncture diameters in most specimens), integument especially shining; males with entire mesepisternum densely covered with white, branched setae, obscuring integument. T1 discal

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 101 patch widely rectangular to ovate; T2 with LLB mostly on lateral surface of T2, forming acute to weakly acute angle with ATB. Female: Pseudopygidial area subovate to subquadrate, with relatively long, distinct basal crescent of silvery setae; S2–S4 with white setae mostly on apicolateral margins (sometimes on entire apical margins); S5 slightly downcurved. Male: Pygidial plate of moderate size or relatively narrow, with distinct basal transverse ridge; S2–S3 with white apical bands of appressed setae; S4–S5 with brown apical fringes (sometimes tinged with white on S4). Comments.—This species is similar to T. brittaini in that the mesepisternum has erect, simple setae, but in females of T. pectoralis the mesepisternum is much less densely punctate (and therefore is more distinctly shining), the mesoscutum has paramedian bands present, and the pseudopygidial area has a distinct basal crescent of shining setae. Males of T. pectoralis and T. brittaini may be separated by the clypeal midline, which is stronger in T. pectoralis, and the leg coloration, which is partially orange in T. pectoralis. Distribution.—CANADA: New Brunswick; USA: Arizona, Arkansas, Colorado, Connecticut, Delaware, Georgia, Idaho, Illinois, Indiana, Kansas, Maine, Maryland, Massachusetts, Michigan, Minnesota, Missis- sippi, Missouri, Nebraska, New Hampshire, New Jersey, New Mexico, New York, North Carolina, North Dakota, Ohio, Pennsylvania, South Dakota, Utah, Vermont, Virginia, Washington D.C., Wisconsin. Host Records.—Melissodes (Eumelissodes) rustica (Say) [= Melissodes druriella (Kirby)] (Mitchell 1962: 479, at nesting site; Hurd et al. 1980, evidence not given). According to John S. Ascher (in lit., 2004), this species is common in Ithaca, NY, on Melissodes druriella; at this location, both bees commonly go to Sol- idago flowers. Floral Records.—Aster ericoides [= Symphyotrichum ericoides (L.) Nesom var. ericoides], Aster pilosus [= Symphyotrichum pilosum (Willd.) Nesom var. pilosum], Bidens bipinnata L., Chrysopsis mariana (L.) Ell., Erigeron sp., Eupatorium coelestinum [= Conoclinium coelestinum (L.) DC.], Grindelia squarrosa (Pursh) Dunal, Heterotheca sp., Myosotis sp., Solidago canadensis L., S. graminifolia [= Euthamia graminifolia (L.) Nutt. var. graminifolia], “blue aster”. Seasonal Records.—3 May (specimens from Washington State) or 1 August to 11 November. Specimens examined.—163 &, 123 % (AUSTIN, BOULDER, CORVALLIS, DAVIS, GAINESVILLE, ITHACA, LAWRENCE, LOGAN, LOS ANGELES, NEW YORK, PHILADELPHIA, RIVERSIDE, SAN FRANCISCO, STARKVILLE, URBANA, WASHINGTON D.C.).

TRIEPEOLUS PENICILLIFERUS (Brues) (Figs. 167–169, 262)

Epeolus penicilliferus Brues 1903: 81–82 [& %, Austin (Travis Co.), Fedor (Lee Co.), Texas] [Types lost?]. Triepeolus penicilliferus; Cockerell 1904: 34. Triepeolus perelegans Cockerell 1921: 8 [Holotype: American Museum of Natural History No. 25089; %, Comobabi Mountains (road from Haynes Well to Combabi, Pima Co.), Arizona; 32° (1)’N 111° (42)’W; (about 3400 ft); (9) August 1916]. new synonymy Triepeolus trichopygus Cockerell and Timberlake 1929: 169 [Holotype: American Museum of Natural History; &, River- side, California; 25 July 1927; Senecio douglasii]. new synonymy

Description.—Length ca. 9–14.5 mm; ITW 1.8–2.5 mm. Integument black to brown, with red or orange on base of mandible, entire labrum, and legs (excluding basal coxae and spine), usually on part or all of clypeus, entire scape, pedicel, and F1, pronotal lobe, and tegula, sometimes on mesepisternum and venter of mesosoma and metasoma; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus shining, lacking or with very weak midline, with distinct large punctures and minute, weakly impressed punctures. Para- median band clearly separated from other pale setae on mesoscutum (females and some males) or joined laterally to pale setae on anterior margin of mesoscutum (most males); mesoscutum shining. Scutellum weakly bigibbous, slightly flattened and extended posteriorly; axillar spine triangular, not or barely reaching

102 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER scutellum midpoint. Mesepisternum lacking erect, simple setae; upper third (including hypoepimeron) covered with dense, pale yellow, branched setae (females), or with pale, branched setae also on ventral margin, sometimes medially with sparse, pale, branched setae (males); ventrally mostly asetose with relatively small, weakly impressed punctures, separated by up to two puncture diameters. T1 discal patch widely rectangular to quadrate; T2 with LLB forming weakly acute to acute angle with ATB. Female: Pseudopygidial area similar to that of T. concavus (i.e., with long, stout setae forming posterior plate-like structure); T5 with narrow band of white setae on apical margin lateral to pseudopygidial area; S5 strongly downcurved. Male: Pygidial plate keyhole shaped, with distinct transverse basal ridge and downturned apical plate; S2–S3 with white apical bands of setae, (S2 with white setae surpassing apical margin); S4–S5 with apical fringes entirely white, brown, or (most commonly) white laterally and brown medially. Comments.—Although I was unable to locate the type of T. penicilliferus, the original description leaves no doubt as to the identity of this species. Triepeolus penicilliferus is extremely similar to T. subnitens, but can be separated from that species by the pseudopygidial area (very short and wide, seemingly formed by the downturned integument in T. subnitens), and by the female S5, which is strongly downcurved in T. penicilliferus but only very slightly downcurved in T. subnitens. Although the distributions of the two species overlap, T. penicilliferus appears to be most abundant in Texas, while T. subnitens appears to be most abundant in California. Males of the two species are difficult to differentiate, but may be separated based on the following characters: the pronotal lobe tends to be red in T. penicilliferus and brown in T. subnitens; the clypeus in profile tends to be flatter in T. penicilliferus than in T. subnitens; the mesepisternum tends to be less densely punctate and less setose in T. penicilliferus than in T. subnitens; and the pygidial plate is narrower, with a more distinct transverse basal ridge in T. penicilliferus than in T. subnitens. Distribution.—MéXICO: Chihuahua, Coahuila, Durango, Sonora, Tamaulipas; USA: Arizona, Califor- nia (Riverside Co., San Bernardino Co.), Colorado, Kansas, Missouri, New Mexico, Oklahoma, Texas. Host Records.—Svastra (Epimelissodes) sabinensis sabinensis (Cockerell) (Rozen, 1983, adult entering nest). Floral Records.—Bahia absinthifolia var. dealbata (Gray) Gray, Bebbia juncea (Benth.) Greene, Core- opsis sp., Eriogonum deflexum Torr., Gaillardia suavis (Gray & Engelm.) Britt. & Rusby, Gutierrezia sarothrae (Pursh) Britt. & Rusby, Helenium microcephalum DC., Heterotheca subaxillaris (Lam.) Britt. & Rusby, Kallstroemia grandiflora Torr. ex Gray, Larrea tridentata (Sessé & Moc. ex DC.) Coville, Nepeta cataria L., Opuntia sp., Psilostrophe cooperi (Gray) Greene, Ratibida columnifera (Nutt.) Woot. & Standl., Senecio douglasii [= Senecio flaccidus Less. var. douglasii (DC.) B.L. Turner & T.M. Barkl.], Silphium asperrimum (= Silphium radula Nutt.), Sphaeralcea sp., Verbena halei Small, Verbesina helianthoides Michx., Viguiera stenoloba Blake. Seasonal Records.—12 April to 16 October. Specimens examined.—111 &, 70 % (AUSTIN, BERKELEY, BOULDER, CORVALLIS, DAVIS, GAINESVILLE, ITHACA, LAWRENCE, LOGAN, LOS ANGELES, MEXICO CITY, NEW YORK, RIVERSIDE, SAN FRANCISCO, STARKVILLE, TEMPE, WASHINGTON D.C.).

TRIEPEOLUS PERPICTUS new species (Figs. 170, 171)

Diagnosis.—This species is known by the diagnostic combination of reddish brown integument of the mesepisternum, dorsal metasoma, and scutellum; the paramedian bands, which reach the anterior margin of the mesoscutum and are often laterally contiguous with pale setae on the anterior margin of the mesoscutum; and the transversely ovate T1 discal patch.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 103 Description of holotype female.—Length ca. 8.5 mm (7–9 mm); ITW 2.0 mm (1.6–2.0 mm). Integu- ment black, with brownish red to orange on basal half of mandible, labrum, apical half of clypeus (sometimes clypeus entirely orange or black), antenna, pronotal lobe, tegula, scutellum (in most specimens), mesepisternum, legs, and dorsal and ventral metasoma (dorsal metasoma black in some specimens) (sometimes also orange on mesoscutum); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus lacking midline, with faint larger punctures, dorsally covered with sparse white setae (most females; denser in males). Paramedian band separated from (or laterally contiguous with) pale setae on anterior of mesoscutum. Scutellum weakly bigibbous; axillar spine triangular, not attaining midpoint of scutellum. Mesepisternum lacking erect, simple setae; dorsally with dense, white, branched setae, grading to sparser setae ventrally, mostly obscuring integument beneath; punctures nearly contiguous to separated by nearly one puncture diameter, integument between punctures raised, tuberculate. T1 discal patch transversely ovate, BTB and ATB parallel-sided; T2 with LLB mostly on lateral surface of T2, forming weakly acute angle with ATB. Pseudopygidial area semicircular, with distinct, pilose, basal crescent; S5 slightly downcurved apically. (Male: Pygidial plate relatively long, lateral margins parallel-sided, with distinct transverse basal ridge and apical downturned plate; S2–S3 with white apical bands, slightly extended past apical margin on S3; S4–S5 with pale golden to white apical fringes.) Distribution.—USA: Arizona, California, Colorado, Nevada, Utah. Floral Records.—Baileya sp., Helianthus petiolaris Nutt., Sphaeralcea sp., Verbesina encelioides (Cav.) Benth. & Hook. f. ex A. Gray. Seasonal Records.—16 April to 23 July. Holotype.—“NEVADA Nye Co., Las Vegas, 38 m NW, IV [April]-28-1973, F. Parker & P. Torchio // Native Bee Survey USDA, Logan, Utah BBSL 582235 //M. G. Rightmyer Triepeolus Database Specimen No. 3571 // HOLOTYPE & Triepeolus perpictus Rightmyer 2007” (LOGAN). Paratypes.—USA: ARIZONA: Maricopa Co., 8 mi NW Wickenburg, 8 May 1990, J. G. & B. L. Rozen (1&, NEW YORK); Pima Co., Tucson, La Cienega & Treat, 28 April 1998, R. Minckley, T. Roulston (1&, LAWRENCE); Yavapai Co., 11 mi SW Congress, 2 May 1990, J. G. Rozen (1&, NEW YORK); CALIFOR- NIA: Imperial Co., Black Mt, NE of Glamis, 17 May 1998, G. R. Ballmer (1&, RIVERSIDE); Riverside Co., 3 mi W Shaver’s Well, 16 April 1966, R. R. Snelling (1&, LOS ANGELES); COLORADO: Moffat Co., Arte- sia, 23 July 1950, C. D. Michener, Helianthus petiolaris (1&, LOGAN); NEVADA: Clark Co., 29.1 mi SW Mesquite, 2 May 1986, P. F. & D. M. Torchio (1&, LOGAN); 4.5 mi W Riverside, 1 May 1997, F. D. Parker (1&, LOGAN); Valley of Fire, 17 May 1970, P. F. Torchio, Baileya (1&, LOGAN); UTAH: Emery Co., San Rafael Desert, 3 mi SSE Temple Mt, 3 June 1982, Parker, Griswold (1&, LOGAN); Washington Co., Dixie State Park, 6 June 1964, G. F. Knowlton (1&, LOGAN); Paradise Canyon, 24–28 May 1983, D. Beck (1&, LOGAN); (unknown county): 12 mi E Jenson, 23 June 1939, Timberlake, Sphaeralcea (3&, BERKELEY); Our- day(?) Valley, 17 July 1952, G. F. Knowlton, G. E. Bohart (1&, LOGAN). Additional specimens.—USA: UTAH: Emery Co., North Temple Wash, San Rafael Desert, 14 June 1983, F. & J. Parker, 5300 ft. (1%, LOGAN); Grand Co., 11 June 1963, L. Crows(?) (1%, LOGAN); Salt Valley, NW of Arches National Park, 4 June 1982, T. L. & R. T. Griswold (1%, LOGAN). Etymology.—This species name is in reference to the red coloration found on much of the integument. The manuscript name of this species was left by an anonymous worker on a specimen of the species. From the Latin, “per” (very) and “pictus” (colored).

TRIEPEOLUS PHAEOPYGUS new species (Figs. 172, 173)

Diagnosis.—This species can be distinguished by the combination of the pseudopygidial area, which is subquadrate, with uniformly golden or coppery, suberect setae, and which has a straight apical margin (some-

104 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER times appearing slightly concave due to long setae laterally on each side of the apical margin); the relatively short clypeus; the laterally contiguous paramedian band on the mesoscutum (rarely isolated); the absence of erect, simple setae on the mesepisternum; the transversely ovate to subrectangular (almost transversely diamond-shaped) discal patch of setae on T1; the 90 degree angle formed by the LLB and ATB of T2; and the medially continuous ATB of that same tergum. Triepeolus phaeopygus somewhat resembles T. fulgidus due to the uniformly golden brown pseudopygidial area, but T. fulgidus has isolated paramedian bands and erect, simple setae on the mesepisternum Description of holotype female.—Length ca. 8.5 mm (7–9 mm); ITW 1.6 mm (1.4–1.9 mm). Integu- ment black or reddish brown, with reddish orange on basal half of mandible, labrum, apical margin of clypeus (sometimes entire clypeus), scape, pedicel, F1, pronotal lobe, tegula, and legs excluding spurs (sometimes excluding basal halves of coxae) (black to reddish brown sometimes on labrum, part or entire clypeus, pronotal lobe, and tegula); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint midline (sometimes lacking), with faint larger punctures. Paramedian band relatively long, laterally contiguous with pale setae on anterior margin of mesoscutum (sometimes poorly distinct from diffuse pale setae on lateral and anterior margins of mesoscutum; rarely isolated from other pale setae). Scutellum moderately (to weakly) bigibbous (often with longitudinal line of pale yellow setae between biconvexities); axillar spine triangular, reaching (or rarely surpassing) midpoint of scutellum. Mesepisternum lacking erect, simple setae; dorsal half with dense, pale yellow, branched setae, extending along posterior margin of mesepisternum near midcoxa (in some specimens not extending along posterior margin); punctation small, moderately impressed, spaced almost contiguously to nearly one puncture width apart in some areas. T1 discal patch nearly diamond-shaped (sometimes transversely ovate or subrectangular), BTB and ATB narrowly interrupted (rarely uninterrupted); T2 with LLB forming 90 degree angle with ATB (sometimes this angle appearing strongly acute due to sparser pale setae at junction of apical and lateral bands). Pseudopygidial area subquadrate, with setae uniformly semi-erect, dense, fine, and apically downturned, apical margin of pseudopygidial area straight or appearing slightly concave due to long lateral setae at apical margin; S5 straight in profile. (Male: Unknown). Distribution.—MéXICO: Chihuahua, Coahuila; USA: Arizona, Kansas, New Mexico, Oklahoma, Texas, Utah. Host Records.—Dieunomia (Epinomia) nevadensis (Cresson) (6 specimens, from nest area, Cochise Co., Arizona), Protoxaea sp. (2 specimens, from “nest #1”, Cochise Co., Arizona). Seasonal Records.—23 May to 20 September. Holotype.—“USA, Arizona: Cochise Co., 1 mi E Apache, VIII [August]-17-95 [1995], J. G. Rozen & S. A. Budick // nesting area of Nomia nevadensis // M. G. Rightmyer Triepeolus Database Specimen No. 3548 // HOLOTYPE & Triepeolus phaeopygus Rightmyer 2007” (NEW YORK). Paratypes.—USA: ARIZONA: Cochise Co., Apache, 22 August 1972, Rozen, Favreau (1&, NEW YORK); 1 mi E Apache, 17 August 1995, J. G. Rozen, S. A. Budick, nesting area of Nomia nevadensis (5&, NEW YORK); 8 mi NE Portal, 23–25 August 1989, J. G. Rozen, R. L. Foster (1&, NEW YORK); Protoxea nest #1 (2&, NEW YORK); 28 August 1990, J. G. & B. L. Rozen, J. Krieger (3&, NEW YORK); 28–29 August 1989, J. G. & B. L. Rozen, R. Foster (1&, NEW YORK); 4 mi E Willcox, 22 August 2002, J. G. Rozen (1&, NEW YORK); Gila Co., Carrizo, 1 September 1986, F. Parker (1&, LOGAN); NEW MEXICO: Hidalgo Co., Rodeo, 25 August 1974, M. & T. M. Favreau (1&, NEW YORK); 1 mi N Rodeo, 29 August 1994, J. G. Rozen, J. S. Ascher (2&, NEW YORK); 2 mi N Rodeo, 22 August 1996, J. G. & B. L. Rozen, A. Pence (1&, NEW YORK); 2.5 km N Rodeo, 12–20 August 1991 (1&, LAWRENCE). Additional specimens.—MÉXICO: BAJA CALIFORNIA SUR: 18 km N Consitución, 13 April 1994, R. McGinley, Verbesina encelioides (1&, LAWRENCE); CHIHUAHUA: Boquillas, 19 June 1992, R. L. Minckley (2&, LAWRENCE); 5 km N Escalón, 20 September 1970, G. E. & R. M. Bohart (2&, LOGAN); Samal- ayuca, 31 August 1992, B. N. Danforth (1&, WASHINGTON D.C.); COAHUILA: Guadalupe, 23 August 1947,

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 105 Michener (3&, NEW YORK); USA: KANSAS: (Seward Co.), Liberal, 16 August 1945, R. H. Beamer (1&, LAWRENCE); OKLAHOMA: (Marshall Co.), 2 mi E Willis, Lake Texoma, June 1965, R. M. Bohart (1&, DAVIS); July 1965, R. M. Bohart (3&, DAVIS); TEXAS: Dimmit Co., Catarina, 23 May 1987, J. L. Neff (1&, AUSTIN); 12 mi E Catarina, Chaparral W.M.A., 13 June 1992, J. L. Neff, A. Hook (1&, AUSTIN); Reeves Co., 3 mi S Orla, 9 July 1961, R. L. Westcott (1&, LOS ANGELES); UTAH: Garfield Co., 2 mi NE Henrieville, 30 August 1985, D. K. Broemeling (1&, LOGAN). Etymology.—This name is in reference to the uniformly dusky brown setae of the pseudopygidial area. From the Greek, “phaeo” (dusky, brown) and “pygos” (rump).

TRIEPEOLUS PUNCTOCLYPEUS new species (Figs. 174, 175)

Diagnosis.—This species primarily can be distinguished by the unique form of the pseudopygidial area, which is a triangle that is longer than wide, and which has uniformly short, appressed, and dense setae; these setae are silvery shining on all margins, enclosing a median patch of slightly darker setae. The species is additionally known by the combination of the relatively short clypeus; the isolated paramedian bands of the mesoscutum; the erect, simple setae of the mesepisternum; and the 90 degree angle formed by the LLB and ATB of T2. Triepeolus punctoclypeus resembles T. denverensis in the appearance of the pseudopygidial area; however, Triepeolus denverensis can be differentiated from T. punctoclypeus by the distinctly coarser texture of the dark, submedian patch of setae on its pseudopygidial area; by the denser punctation of its mesepisternum (punctures nearly contiguous in T. denverensis, as opposed to punctures separated by two puncture diameters in some places in T. punctoclypeus—the integument of these impunctate regions appearing tuberculate); and by the absence of distinct larger punctures on its clypeus (such larger punctures present in T. punctoclypeus). The specimens from California also closely resemble T. quadratus (and, to a lesser degree, T. antiochensis), but may be distinguished from them by the pseudopygidial area, which is comprised of very differently textured setae in those two species, and by the longer axillar spines in T. quadratus and T. antiochensis. Description of holotype female.—Length ca. 10 mm (10–11 mm); ITW 2.6 mm (2.0–2.6 mm). Integu- ment black, with red on basal half of mandible, labrum, apical margin of clypeus, pedicel, scape, F1, tegula, and legs except forecoxa, basal halves of mid- and hindcoxae and spurs (sometimes black on labrum, clypeus, pedicel, and scape); dorsum of mesosoma and metasoma with bands of yellow (to pale yellow) setae. Clypeus lacking (or with weak) midline, with distinct larger punctures (weak punctures in one specimen from Utah). Paramedian band clearly separated from other pale setae on mesoscutum. Scutellum moderately bigibbous; axillar spine triangular, reaching scutellar midpoint. Mesepisternum with erect, simple setae; dorsal third and anterior surface with dense, pale yellow, branched setae, absent (or sparser) on hypoepimeron (reduced and not on anterior surface in one specimen from Nevada); ventrally mostly asetose; punctures dense and small (occasionally separated by 1–2 puncture diameters), integument between raised, tuberculate. T1 rectangular (to transversely ovate), ATB interrupted medially (uninterrupted in one specimen from Utah); T2 with LLB forming 90 degree angle with ATB (weakly acute in one specimen from Utah); ATB medially interrupted. Pseudopygidial area triangular, with blunt, appressed setae very uniform in density and texture, appearing silvery on all margins; S5 straight along length. (Male: Unknown.) Comments.—Triepeolus punctoclypeus currently consists of two slightly differing forms. One is represented by three females from San Bernardino Co., California, in which the bands of setae on the metasoma are relatively dense and yellow; the other is represented by three females from Utah and Nevada, in which the bands of setae are paler yellow and more diffuse. It is possible that these differences are due to preservation or age of the specimens when collected, or they may represent different geographic forms of the species. Alternatively, additional material may lend support for segregating the two types as two distinct species.

106 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Distribution.—USA: California (San Bernardino Co.), Nevada, Utah. Seasonal Records.—6 September to 30 October. Holotype.—“Cedar Springs, San Bernardino Co., Calif. [California], X [October]-30-1966 // A. R. Gillogly Collector // Univ. Calif. Riverside Ent. Res. Museum UCR ENT 57188 // M. G. Rightmyer Triepeo- lus Database Specimen No. 1282 // HOLOTYPE & Triepeolus punctoclypeus Rightmyer 2007” (RIVERSIDE). Paratypes.—USA: CALIFORNIA: San Bernardino Co., Santa Ana Canyon, 6 September 1931, C. D. Michener (1&, LAWRENCE); Cajon Pass, 6 October 1977, N. J. Smith (1&, DAVIS). Additional specimens.—USA: NEVADA: Douglas Co., 6 mi N Yerington, 30 September 1956, Tim- berlake (1&, WASHINGTON D.C.); UTAH: Kane Co., 16 mi W Glen Canyon, 13 September 1964, Timberlake (1&, WASHINGTON D.C.); San Juan Co., 14 mi SE Moab, 23 September 1974, B. Bohart, W. Hanson (1&, LOGAN). Etymology.—This name is in reference to the larger punctures of the clypeus, which help to differentiate this species from T. denverensis. From the Latin, “punctum” (small hole) and “clypeus” (shield).

TRIEPEOLUS QUADRATUS new species (Figs. 176, 177)

Diagnosis.—This species can be distinguished by the combination of the triangular to subquadrate pseudopygidial area, which is about as long as wide, and which has the entire basal half to third composed of dense, silvery setae, and the apical half or more composed of sparser, coarser, dark setae; the relatively short clypeus; the isolated paramedian band on the mesoscutum; the relatively long axilla, with its medially incurved apical spine; the erect, simple setae of the mesepisternum; the rectangular T1 discal patch; the 90 degree angle formed by the LLB and ATB of T2; and the medially interrupted apical bands of all the metasomal terga. Triepeolus quadratus resembles T. antiochensis, but the two species can be distinguished by the ATBs of the metasomal terga, which are strongly interrupted medially in T. quadratus and continuous or nearly continuous in T. antiochensis. Triepeolus quadratus also resembles specimens of T. punctoclypeus from California, which differ from T. quadratus by the uniformly textured setae of the pseudopygidial area and by the shorter axillar spine. Description of holotype female.—Length ca. 10 mm (10–10.5 mm); ITW 2.2 mm (2.2–2.3 mm). Integ- ument black, with red on most of mandible, labrum, apical clypeus, pronotal lobe (usually), tegula, and legs distal to femora, (sometimes on scape, pedicel, and F1); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint midline, mostly covered with sparse, dark brown setae (white in male); face with transverse area of white setae at level of antennal sockets, excluding supraclypeal area. Paramedian band clearly separated from other pale setae on mesoscutum. Scutellum strongly bigibbous; axillar spine triangular, slightly surpassing midpoint of scutellum, apical point directed inward. Mesepisternum with erect, simple setae; with small patch of pale yellow, branched setae below scrobal grove and beneath pronotal lobe (or sparsely covering most of mesepisternum in male); punctation very small and nearly contiguous. T1 discal patch widely rectangular; T2 with LLB reduced, seemingly forming 90 degree angle with ATB; all metasomal terga with ATBs medially interrupted by ca. 1 OD (0.5–0.7 in male; more closely spaced on posterior- most terga). Pseudopygidial area triangular (to subquadrate), basally with very fine, silvery setae, apical half to two-thirds with coarse, apically-downturned setae; S5 straight in profile. (Male: Pygidial plate keyhole shaped, with distinct subapical ridge; S2–S3 with well developed apical bands of white setae; S4–S5 with white apical fringes, dusky or brown medially on S5.) Comments.—One specimen of this species bears a manuscript name that makes reference to dark pig- mentation at the end of the body.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 107 Distribution.—USA: California (Mono Co., San Luis Obispo Co.). Floral Records.—Hemizonia paniculata Gray. Seasonal Records.—29 August to 12 September. Holotype.—“Black L. [Lake] Cn. [Canyon] Cal. [California] S. [San] Luis Obispo Co. VIII [August].29.1955 // R. M. Bohart Collector // M. G. Rightmyer Triepeolus Database Specimen No. 1291 // HOLOTYPE & Triepeolus quadratus Rightmyer 2007” (DAVIS). Paratypes.—USA: CALIFORNIA: Mono Co., Mono Bay, 30 August 1943, Timberlake, Hemizonia paniculata (2&, 1% BERKELEY); San Luis Obispo Co., 4 mi S Oceano, 12 September 1956, J. C. Hall (1&, LOGAN). Etymology.—This name is in reference to the widely rectangular shape of the T1 discal patch. From the Latin, “quadratus” (four-cornered).

TRIEPEOLUS QUADRIFASCIATUS ATLANTICUS Mitchell (Figs. 178–181)

Triepeolus quadrifasciatus atlanticus Mitchell 1962: 479–480, Fig. 112 [Holotype: U. S. National Museum of Natural History No. 400194 (on indefinite loan from North Carolina State University); &, Bogue (Carteret Co.), North Caro- lina; 31 August 1933; Monarda punctata].

Description.—Length ca. 14–16mm; ITW 3mm; rarely as small as length 12mm, ITW 2.5mm. Integument black, with red to orange on mandible, labrum, apical margin of clypeus (usually), scape, pedicel and F1, pronotal lobe, tegula, and legs distal to coxae (spurs black); dorsum of mesosoma and metasoma with bands of yellow to pale yellow setae. Clypeus with integument shining, asetose to sparsely covered with golden setae (more densely covered in males), lacking midline, and with distinct larger punctures. Pronotal collar long (ca. 2 OD in length), especially medially. Mesoscutum shining with paramedian band narrow and clearly separated from other pale setae on mesoscutum, or absent. Axillar spine pointed, surpassing scutellum midpoint (or very rarely only reaching midpoint), curving slightly inward apically; scutellum somewhat flattened and extended posteriorly. Mesepisternum lacking erect, simple setae; with white, appressed, branched setae dorsally and sometimes posteroventrally; ventral half to three-fourths with brown, appressed, branched setae or asetose; ventral half of mesepisternum with punctures small, separated by up to a puncture diameter (usually less); integument between punctures raised, slightly tuberculate. T1 almost entirely covered with yellow setae except for black longitudinal line medially and often small black oval, triangle, or large rectangle medially; T2 with LLB absent, or rarely present but very reduced (forming 90 degree angle with ATB). Female: Pseudopygidial area subquadrate, covered with darkly shining, coarse setae, these setae very slightly sparser, more erect, and longer on apical half to three-fourths; metasomal venter brown, with lateral patches of white setae on S2–S4 (sometimes extending across entire S3 apical margin; usually faint on S4); S5 slightly downcurved apically. Male: Pygidial plate relatively wide, with distinct basal transverse ridge; S2–S3 with white apical setae laterally or on entire apical margin; S4–S5 with dark brown apical fringes (S4 sometimes with small patch of white setae on apicolateral margin). Comments.—The subspecies T. q. atlanticus has the black T1 discal patch commonly represented by a longitudinal line with a median triangular or subovate region. It is rarely found as a longitudinal line only, or is present as a strongly rectangular area. Specimens from St. Louis, Missouri, range from the T1 discal patch perfectly longitudinally linear, through linear with a small ovate black patch medially, to strongly rectangular medially; thus, the shape of the T1 discal patch is not always consistent for delineating the subspecies of T. quadrifasciatus, especially at this portion of their respective distributions, where the two subspecies may be interbreeding. However, a consistent character is found in the coloration of the face: T. q. atlanticus differs from T. q. quadrifasciatus by the coloration of clypeus and interantennal area, which is entirely red and

108 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER strongly delineated from the black lateral areas of the face in T. q. quadrifasciatus. In T. q. atlanticus, the clypeus is either entirely black or black with red on the apical margin. Distribution.—USA: Alabama, Florida, Georgia, Illinois, Maryland, Mississippi, Missouri, New Jersey, North Carolina, Virginia. Host Records.—Svastra (Epimelissodes) atripes atrimitra (LaBerge) (3 specimens from nesting site, in Alabama; and Cane, 1995, adults inspecting and entering host nests). Floral Records.—Bidens pilosa L., “white goldenrod” (= Solidago bicolor L.). Seasonal Records.—30 June to 19 October. Specimens examined.—26 &, 27 % (BERLIN, CORVALLIS, GAINESVILLE, ITHACA, LAWRENCE, LOS ANGELES, NEW YORK, STARKVILLE, URBANA, WASHINGTON D.C.).

TRIEPEOLUS QUADRIFASCIATUS QUADRIFASCIATUS (Say) (Figs. 182–184)

Epeolus 4 fasciatus17 Say 1823: 81 [&; Arcansa (Arkansas)] [Type lost]. Triepeolus quadrifasciatus; Mitchell 1962: 485.

Description.—Length ca. 13–16mm; ITW 2.4–3.1 mm. Integument black, with red on basal half of mandible, labrum, clypeus, interantennal region, scape, pedicel, T1, pronotal lobe, tegula, and legs distal to coxae, (sometimes including coxae; spurs black), often on mesoscutum, axilla, and scutellum; dorsum of mesosoma and metasoma with bands of yellow to pale yellow setae. Clypeus integument shining, asetose to sparsely covered with golden setae, lacking midline, with distinct or vague larger punctures. Pronotal collar in dorsal view elongate, especially medially (ca. 2 or more OD in length). Mesoscutum shining with paramedian band narrow and clearly separated from other pale setae on mesoscutum, or absent. Scutellum somewhat flattened and extended posteriorly; axillar spine pointed, surpassing scutellum midpoint (or very rarely only reaching midpoint), curving slightly inwards apically. Mesepisternum lacking erect, simple setae; dorsal third covered with yellow, branched setae, except sometimes in females with area between hypoepimeron and pronotal lobe at level of scrobal groove lacking yellow setae, sometimes in males with yellow setae extending further ventrally; ventrally asetose or with brown, branched setae, with small punctures separated by up to a puncture diameter (usually less); integument between raised, slightly tuberculate. T1 almost entirely covered with yellow setae except for black longitudinal line medially and sometimes small black oval or diamond medially; T2 with LLB absent, or rarely reduced, forming 90 degree angle with ATB. Female: Pseudopygidial area subquadrate, uniformly covered with darkly shining, coarse setae; metasomal venter brown, sometimes with small lateral patches white setae on S2–S3 (sometimes vaguely on S4); S5 slightly downcurved apically. Male: Pygidial plate relatively wide; S2–S3 with white apical setae laterally or on entire apical margins; S4– S5 with dark brown apical fringes. Comments.—Although the type of T. quadrifasciatus is no longer available for study, the original description leaves no doubt as to the identity of this species, and is well understood by most workers making determinations in the collections that I have examined. This subspecies differs from T. q. atlanticus by the entirely red clypeus and interantennal area, which strongly contrast with the black lateral areas of the face; in addition the following areas are often reddish: the axillae, scutellum, and sometimes mesoscutum and mesepisternum. The T1 discal patch tends to be a longitudinal line, sometimes with a small medial ovate or diamond area. However, T. q. atlanticus often has the T1 discal patch with a small medial ovate area, or even rarely is represented by a longitudinal line only.

17. Hurd (1979: 2095) cites this species as “Epeolus 4-fasciatus;” however, the original description of this species does not include a hyphen in the specific epithet.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 109 Distribution.—USA: [Arkansas—original description]; Kansas, Louisiana, Oklahoma, Texas (counties east of panhandle). Floral Records.—Gaillardia pulchella Foug., Helenium tenuifolium [= Helenium amarum (Raf.) H. Rock var. amarum], Helianthus petiolaris Nutt., Liatris sp., Nama ovata (= Hydrolea ovata Nutt. ex Choisy), Prionopsis ciliata [= Grindelia papposa Nesom & Suh], Rudbeckia hirta L., cotton (= Gossypium sp.). Seasonal Records.—31 May to 12 October. Specimens examined.—22 &, 11 % (AUSTIN, CORVALLIS, ITHACA, LAWRENCE, LOS ANGELES, NEW YORK, SAN FRANCISCO, STARKVILLE, URBANA, WASHINGTON D.C.).

TRIEPEOLUS REMIGATUS (Fabricius) (Figs. 185–187, 260)

Melecta remigata Fabricius 1804: 387 [Carolina] [Type lost? (See Comments, below)]. Epeolus remigatus; Lepeletier de Saint-Fargeau (in Lepeletier de Saint-Fargeau and Serville 1825): 104. Epeolus superbus Provancher 1895: 190–191 [Holotype: Université Laval, Collection Provancher; &, (Los Angeles, Cal- ifornia); 1734 (See Comments, below)]. new synonymy Triepeolus remigatus; Robertson 1901: 231; Mitchell 1962: 480–481, Fig. 112 [redescription, illustration of scutellum, axillae]; Rozen 1966: 17–19, Fig. 24 [description, illustration of predefecating larva]; Bohart 1966: 255–261, Figs. 1–20 [descriptions, photographs and illustrations of egg, first through fourth instar, prepupa, adult]. Epeolus texanus var. nigripes Cockerell 1898: 61 [Holotype: U. S. National Museum of Natural History No. 18966; %, Mesilla (Dona Ana Co.), New Mexico; 14 August; Helianthus ciliaris]. new synonymy Triepeolus texanus nigripes; Cockerell 1916: 392.

Description.—Length ca. 10.5–15.5 mm; ITW 2.4–3.5 mm. Integument black, with red on basal half of mandible and sometimes on axillar spine; often with orange on labrum, apical margin of clypeus, basal antenna, and legs in Southwestern specimens; dorsum of mesosoma and metasoma with bands of yellow setae. Face sometimes slightly elongate. Clypeus black, with midline absent or rarely weak; larger punctures weak; mostly asetose (some females), basally covered with white setae (some females and some males), or entirely covered with white or brown setae (some males). Paramedian band joining laterally with yellow setae on apical margin of mesoscutum to form strong anchor pattern (both sexes). Scutellum strongly to moderately bigibbous; axillar spine reaching midpoint or, more commonly, surpassing midpoint, with slightly incurved apical point. Mesepisternum lacking erect, simple setae; with distinct dorsal region of dense, yellow, branched setae (lacking on hypoepimeron) and ventral region of black, branched setae; integument beneath with small punctures nearly contiguous to separated by 0.5 puncture diameter. T1 discal patch ovate to subtriangular, sometimes small; T2 with LLB forming 90 degree angle with ATB or forming semicircular, basal black region. Female: Pseudopygidial area subovate to subquadrate, setae almost uniformly glossy, fine, and dark, but slightly denser and finer basally; ventral metasoma lacking pale setae (Eastern and Midwestern distributions) or S2–S4 with bands of pale setae on apicolateral margins (Western distribution); S5 not downcurved or very slightly downturned apically. Male: Pygidial plate of moderate size, with distinct basal transverse ridge; S2–S3 with apicolateral bands of pale setae (S3 medially also with dark brown setae, which slightly extend past apical margin); S4–S5 with apical fringes of dark brown setae, S4 also with white setae on apicolateral margin in specimens from Western distribution. Comments.—According to Zimsen (1964), the holotype of this species and Epeolus mercatus Fabricius should be located in the Bosc collection in the Museum of Natural History in Paris; however, the specimens could not be found (Claire Villemont, in lit., 2005). Nonetheless, the original description mentions an important feature for identifying T. remigatus, namely the distinctly trilobed (i.e., anchor-shaped) black region on the mesoscutum, and this species is consistently understood by workers to mean the one described herein (to judge from numerous previously identified specimens in the collections that I have examined).

110 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER According to the original description, the species name Epeolus superbus was apparently based on a single female type specimen. The holotype specimen, a female from the same collection locality as indicated in the original description, nonetheless has two lectotype labels on it. To my knowledge neither lectotype designation has been published. The full label data for the holotype are as follows: “1734 // Epeolus superbus Prov. Cal. // Lectotype 442 Epeolus superbus (Huart) Provancher Comeau 1944 [red label] // Lectotype Epeo- lus superbus Provancher 1734 Barron ’71 [red label].” Females of T. remigatus can be distinguished by the anchor-shaped region of black setae on the mesoscutum in combination with the nearly uniform, darkly shining setae of the pseudopygidial area. Males of T. remigatus resemble T. concavus; however, in T. remigatus the paramedian bands and yellow setae on the anterior margin of the mesoscutum form a strong anchor pattern, while in T. concavus the paramedian bands are not distinct from the region of dense yellow setae on the anterior third or fourth of the mesoscutum. Triepeolus remigatus is also similar to T. nevadensis, but in T. nevadensis the paramedian bands are not so strongly developed as in T. remigatus, the mesoscutum is more distinctly shining, the scutellum is more flattened and extended posteriorly, and the T1 discal patch is distinctly rectangular (as opposed to ovate or subtriangular in T. remigatus). The males have white banding only on S3 in T. nevadensis (as opposed to both S2 and S3 in T. remigatus). Distribution.—MéXICO: Chihuahua; Coahuila, Durango, Jalisco, San Luis Potosí, Sinaloa, Sonora, Zacatecas; USA: Arizona, California (Tulare Co. south to Riverside Co. and San Diego Co.), Colorado, Del- aware, Georgia, Illinois, Indiana, Iowa, Kansas, Louisiana, Maryland, Mississippi, Missouri, New Jersey, New Mexico, New York, North Carolina, Oklahoma, Pennsylvania, Tennessee, Texas, Virginia, Washington D.C. Host Records.—Dieunomia (Dieunomia) heteropoda (Say) (2 specimens, at nest entrance, Hidalgo Co., New Mexico); Centris sp. (1 specimen with label “Centris Nest #1” from Cochise Co., Arizona), Peponapis pruinosa (Say)? (Mitchell, 1962, collection records; John S. Ascher, in lit., 2003, unpublished data), Xenoglo- ssa strenua (Cresson) (Mitchell, 1962, collection records; Rozen, 1966, larvae taken from nest; Bohart, 1966, eggs and larvae taken from nests). Floral Records.—Bahia absinthifolia var. dealbata (Gray) Gray, Centaurea repens [= Acroptilon repens (L.) DC.], Cichorium intybus L., Cosmos sp., Crotalaria incana L., Eriogonum abertianum neomexicanum (= Eriogonum abertianum Torr.), E. deflexum Torr., Eupatorium linearifolium (= Eupatorium glaucescens Ell.), Gaillardia pulchella Foug., Helianthus annuus L., H. ciliaris DC., Kallstroemia grandiflora Torr. ex Gray, Melilotus alba [= Melilotus officinalis (L.) Lam.], Petalostemon sp. (= Dalea sp.), Sphaeralcea fendleri Gray ssp. elongata Kearney, Verbesina encelioides (Cav.) Benth. & Hook. f. ex Gray, Vernonia noveboracensis (L.) Michx., China aster [= Callistephus chinensis (L.) Nees], “cultivated cucurbits”. Seasonal Records.—9 May to 22 October. Specimens examined.—183 &, 50 % (AUSTIN, BOULDER, CHAMELA, CORVALLIS, DAVIS, ITHACA, LAWRENCE, LOGAN, LOS ANGELES, NEW YORK, QUéBEC, RIVERSIDE, SAN FRANCISCO, URBANA, WASHING- TON D.C.).

TRIEPEOLUS ROBUSTUS (Cresson) (Figs. 188–190)

Epeolus robustus Cresson 1878: 85–86 [Lectotype: Academy of Natural Sciences No. 2218; &, New Mexico]; Cresson 1916: 129 [lectotype designation]. Epeolus nigriceps Smith 1879: 103 [Holotype: The Natural History Museum, London No. 17B.520; &, Texas]. new synonymy Triepeolus nigriceps; Cockerell 1905a: 314. Triepeolus robustus; Cockerell 1906: 304.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 111 Description.—Length ca. 11–18 mm; ITW 2.2–3.6 mm. Integument entirely black to brown; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus shining, with distinct larger punctures (females) or densely covered with long, appressed, pale yellow, setae (most males), appearing recessed due to dense, erect, pale yellow and/or brown setae on upper face. Mesoscutum shining, with paramedian band barely distinct from diffuse, erect and appressed setae on anterior mesoscutum. Scutellum flattened and somewhat extended posteriorly; axillar spine triangular, apex rounded, not attaining midpoint of scutellum. Mesepisternum with erect, simple setae; dorsal third, except hypoepimeron, with dense, pale yellow, branched setae (area expanded on males); ventrally with sparse, black to dark brown, branched setae; punctures separated by up to one puncture diameter in some places. T1 discal patch distinctly rectangular, sometimes subtriangular, rarely filled with diffuse, pale yellow setae; T2 with LLB forming 90 degree angle with ATB. Female: Pseudopygidial area delimited by downturned integument, setae relatively long, fine, appressed, and poorly differentiated from rest of setae on T5 (except for white setae apicolaterally on T5); S5 strongly downcurved. Mesosoma and metasoma venter entirely black (occasionally with small patch white setae apicolaterally on S4). Male: Pygidial plate relatively wide, with distinct transverse basal ridge and downturned apical plate; S2–S3 with white, apicolateral setae; S4–S5 with apical fringes (black and white to varying degrees on S4, black on S5). Comments.—The T. nigriceps holotype (from Texas) differs from the typical T. robustus specimens only in that it has the T1 discal patch partially filled with diffuse pale yellow setae, T5 lacks pale setae lateral to the pseudopygidial area, and has entirely brown setae on face. There are specimens of T. robustus identified by Paul Hurd as a new species, and assigned a manuscript name in honor of Jerome G. Rozen, Jr. This is likely due to the fact that T. robustus appears to have been commonly confused with the new species T. flavigradus; Hurd, who was working on a revision of the genus, likely had not yet seen the lectotype of T. robustus, and thus may not have been aware of the mix-up. Triepeolus flavigradus can be differentiated from T. robustus by the coloration of the transverse bands of pale setae on the metasomal terga (grading posteriorly from yellow to white in T. flavigradus; uniformly pale yellow in T. robustus), as well as by the general shape of the T1 discal patch (quadrate to subtriangular in T. flavigradus; strongly rectangular in T. robustus) and the LLB of T2 (forming a right angle with the ATB in T. robustus, usually absent in T. flavigradus). Distribution.—MéXICO: Chihuahua, Durango; USA: Arizona, California (locality not specified), Nevada, New Mexico, Texas (Culberson Co., Hudspeth Co., Presidio Co.), Utah. Floral Records.—Asclepias subverticillata (A. Gray) Vail, Baccharis glutinosa [= Baccharis salicifolia (Ruiz & Pavón) Pers.], Bahia absinthifolia var. dealbata (Gray) Gray, Eriogonum deflexum Torr., Gaillardia sp., Helianthus annuus L., Kallstroemia grandiflora Torr. ex Gray, Melilotus alba [= Melilotus officinalis (L.) Lam.], Verbesina encelioides (Cav.) Benth. & Hook. f. ex Gray. Seasonal Records.—2 August to 16 October. Specimens examined.—160 &, 90 % (AUSTIN, DAVIS, GAINESVILLE, LAWRENCE, LOGAN, LONDON, LOS ANGELES, MEXICO CITY, NEW YORK, NEW YORK-ASCHER, PHILADELPHIA, RIVERSIDE, SAN FRANCISCO, TEMPE, TUCSON, WASHINGTON D.C.).

TRIEPEOLUS ROHWERI Cockerell (Figs. 191, 192)

Triepeolus rohweri Cockerell 1911: 668–669 [Holotype: U. S. National Museum of Natural History No. 100035; %, Canadian Zone, North Boulder Creek, Boulder Co. Colorado; 23 August 1907].

Description.—Length ca. 10 mm; ITW 2.1–2.4 mm. Integument black, with red on apical half of mandible; orange on F1 and parts of legs (especially middle and hind legs); brownish orange on tegula; dorsum of meso-

112 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER soma and metasoma with bands of pale yellow setae. Clypeus with faint midline and distinct larger punctures, entirely covered with white setae in males. Paramedian band laterally contiguous with diffuse, pale setae on anterior of mesoscutum. Scutellum moderately bigibbous; axillar spine triangular, nearly reaching midpoint of scutellum. Mesepisternum with long, erect, simple setae; dorsal half covered with pale yellow, branched setae (sparser on hypoepimeron); ventrally with sparse, brown, branched setae, with punctures nearly contiguous to separated by almost one puncture diameter, with integument between punctures raised, tuberculate (females), or mesepisternum entirely covered with pale yellow, branched setae (sparser on hypoepimeron; males). T1 discal patch transversely ovate, with BTB and ATB nearly parallel and only separated by little more than 1 OD ; T2 with LLB forming 90 degree angle with ATB. Female: Pseudopygidial area semicircular to subtriangular, with setae of similar texture throughout, but with setae on basal third shining silvery and remaining apical setae shining gold; S5 straight in profile. Male: Pygidial plate keyhole shaped, rather long and narrow, with distinguishable transverse basal ridge; S2–S3 with apical bands of white setae; S4–S5 with apical fringes of setae brown with white laterally. Comments.—The pseudopygidial area of this species is similar to that of T. margaretae; however, the two species differ in that only T. rohweri has erect, simple setae on the mesepisternum. The pseudopygidial area is also similar to that of T. claytoni, but these are distinguishable by the angle formed by the lateral longitudinal and apical transverse bands of pale setae on T2 (forming an acute angle in T. claytoni, forming a 90 degree angle in T. rohweri) Distribution.—USA: Colorado (Boulder Co.) Seasonal Records.—18 to 23 August. Specimens examined.—1 &, 1 % (BOULDER, WASHINGTON D.C.).

TRIEPEOLUS RUFITHORAX Graenicher (Figs. 193–195, 245)

Triepeolus rufithorax Graenicher 1928: 279–281 [Holotype: U. S. National Museum of Natural History No. 41793; &, Miami (Miami-Dade Co.), Florida; 16 July 1927]; Rozen 1966: 4, Fig. 1 [position of egg within host cell]; Mitchell 1962: 481, 482, Fig. 112 [redescription, illustration of scutellum, axillae]. Triepeolus alachuensis Mitchell 1962: 462 [Holotype: Florida State Collection of Arthropods; &, Alachua Co., Florida; 12 May 1959; Melilotus alba]. new synonymy

Description.—Length ca. 10.5–14 mm; ITW 2.0–3.1 mm. Integument black or brown, with the following entirely or partly red: mandible, labrum, clypeus, interantennal area (sometimes), basal or entire antennae, dorsal surface of mesosoma, and often parts of metasomal terga; in addition, integument orange on legs. Clypeus with strong midline and faint larger punctures (sometimes covered by white, medially-directed setae, especially in males). Paramedian band narrow and clearly separated from other pale setae on mesoscutum, or absent, sometimes with dark integumental coloration where the paramedian band would normally be found. Scutellum moderately bigibbous; axillar spine triangular with pointed apex, reaching or surpassing scutellar midpoint. Wings entirely dusky. Mesepisternum lacking erect, simple setae or with short, sparse, suberect, simple setae; with dense, white and/or black, branched setae on upper third (absent on hypoepimeron), mostly asetose especially ventrally (sparsely setose in males); irregularly punctate, punctures almost contiguous to separated by two puncture diameters. T1 discal patch variable, variously appearing trapezoidal, rectangular, or triangular, sometimes with BTB reduced or entirely absent; T2 with LLB absent or reduced, forming weakly acute angle with ATB. (Metasomal tergal bands sometimes very narrow.) Female: Pseudopygidial area subovate to subquadrate, with distinct basal crescent extending laterally to apical margin and partially surrounding medial area of coarse setae; S3–S4 (sometimes also S2) with pale banding apicolaterally; S5 slightly downcurved apically. Male: Pygidial plate of moderate size, keyhole shaped with distinct basal transverse ridge; S2–S3 with white apical setae, somewhat extended medially on S3; S4–S5 with apical fringes brown.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 113 Comments.—Blacker individuals of this species might be confused with T. lunatus because of the similar shape of the T1 discal patch and the pseudopygidial area; see comments under that species. The T. alachuensis holotype has red integumental coloration restricted apically and laterally on the clypeus, the axillar spines, and anterolaterally on the mesoscutum; orange coloration on the antennae basal to the base of F2, the tegula, the pronotal lobes, and the legs; the T1 discal patch is trapezoidal; and the metasomal banding is narrow. It arguably might have also been synonymized under T. lunatus, but I have placed it here due to the similar type localities of T. alachuensis and T. rufithorax. Distribution.—USA: Alabama, Florida, Georgia. Host Records.—Svastra sp. (2 specimens “from burrow,” in Everglades National Park, Florida); Svastra (Epimelissodes) obliqua obliqua (Say) (Rozen, 1964, adults entering nest, egg found in cell). Floral Records.—Bidens pilosa L., Borrichia frutescens (L.) DC., Lythrum lineare L. Seasonal Records.—13 March to 7 September. Specimens examined.—97 &, 38 % (ANN ARBOR, BOULDER, GAINESVILLE, ITHACA, LAWRENCE, LOS ANGELES, NEW YORK, SAN FRANCISCO, URBANA, WASHINGTON D.C.).

TRIEPEOLUS RUFOCLYPEUS (Fox) (Figs. 196, 197)

Epeolus rufoclypeus Fox 1891: 344 [Holotype: Academy of Natural Sciences No. 10122; &, Kingston, Jamaica; (April)]. Triepeolus foxii Cockerell 1919a: 179 [Holotype: U. S. National Museum of Natural History No. 20711; &, Portland, Jamaica]. new synonymy Triepeolus rufoclypeus; Brumley 1965: 73. Triepeolus foxi; Raw 1984: 503 [lapsus calami].

Description.—Length ca. 8.5–13.5 mm; ITW 1.9–2.3 mm. Integument black to reddish brown, with red on basal mandible, entire or outer margins of labrum, apical margin of clypeus, scape, pedicel, F1, pronotal lobe, tegula, and entire or parts of legs (excluding basal coxae but only sometimes excluding spurs); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus lacking or with weak midline dorsally, lacking or with faint larger punctures, sometimes covered with sparse, pale, medially-directed setae (denser in males). Pronotal collar in dorsal view with pale setae usually of the same length, rarely thinner submedially in some males. Paramedian band clearly separated from other pale setae on mesoscutum, truncate or tapering anteriorly and reaching anterior margin of mesoscutum. Scutellum weakly bigibbous; axillar spine triangular, reaching or slightly surpassing scutellum midpoint. Mesepisternum lacking erect, simple setae; dorsally with dense, white, branched setae beneath scrobal groove and pronotal lobe; ventral integument shining, with irregular punctures, nearly contiguous to separated by almost a puncture diameter in some areas, sometimes sparsely covered with pale, branched setae (especially in males). T1 discal patch triangular; with <-shaped band of pale setae mostly on lateral margin, but with ATB present (sometimes reduced, but less reduced than that of T. cameroni); T2 with LLB absent or reduced, forming weakly acute angle with ATB. Female: Pseudopygidial area subquadrate, longitudinally elongate, with nearly uniformly golden setae (basal third with shorter, slightly more shining setae); S5 straight in profile. Male: Pygidial plate keyhole shaped, with weak transverse basal ridge and apical downturned plate; S3–S5 with well-developed apical fringes of setae; metasomal sternal setae uniformly golden. Comments.—This species strongly resembles T. cameroni and may represent a northern variety of that species; see comments under T. cameroni for further discussion. Distribution.—MéXICO: Nuevo León, Sinaloa, Tamaulipas; USA: Texas (counties south and east of Gillespie Co. and Travis Co.); JAMAICA: Kingston, Portland; GRENADA: St. George’s. Floral Records.—Asclepias sp., Cevallia sinuata Lag., Gaillardia suavis (Gray & Engelm.) Britt. & Rusby, Hedyotis nigricans [= Stenaria nigricans (Lam.) Terrell var. nigricans], Helenium microcephalum

114 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER DC., Nepeta cataria L., Palafoxia texana DC., Phacelia laxa Small, Phyla nodiflora (L.) Greene, Prosopis glandulosa Torr., Ratibida columnifera (Nutt.) Woot. & Standl., Rudbeckia bicolor (= Rudbeckia hirta L. var. pulcherrima Farw.), Sapindus drummondii [= Sapindus saponaria L. var. drummondii (Hook. & Arn.) L. Ben- son], Verbesina encelioides (Cav.) Benth. & Hook. f. ex Gray, squash (= Cucurbita). Seasonal Records.—April (day unspecified) to 28 October. Specimens examined.—110 &, 165 % (BERKELEY, DAVIS, GAINESVILLE, LAWRENCE, LOGAN, LOS ANGELES, NEW YORK, PHILADELPHIA, STARKVILLE, URBANA, WASHINGTON D.C.).

TRIEPEOLUS RUGOSUS Mitchell (Figs. 198–200)

Triepeolus rugosus Mitchell 1962: 482–483, Fig. 112 [Holotype: Florida State Collection of Arthropods; &, Highlands Hammock State Park (Highlands Co.), Florida; 31 March 1956].

Description.—Length ca. 8.5–10 mm; ITW 2.0–2.6 mm. Integument black; dorsum of mesosoma and metasoma with bands of setae pale gray to white. Clypeus elongate, lacking midline and larger punctures; covered with sparse white setae. Paramedian band clearly separated from other pale setae on mesoscutum, narrow, reaching anterior margin of mesoscutum and curving slightly outwards anteriorly. Mesoscutum and scutellum shining. Scutellum moderately bigibbous (holotype scutellum somewhat extended posteriorly); axillar spine triangular, reaching or almost reaching midpoint of scutellum. Mesepisternum with dense, erect, simple setae; punctures deeply impressed and generally separated by ca. one puncture diameter, with integument between raised, almost tuberculate; with branched, white setae on dorsal third. T1 discal patch transversely ovate to rectangular, ATB interrupted medially, but T2 and distal terga with transverse bands of pale setae continuous (notched slightly on T2); T2 with LLB forming weakly acute angle with ATB (mostly on lateral surface of T2). Female: Pseudopygidial area subquadrate to subovate, with basal region of silvery reflectance strongly differentiated from relatively long, coarse setae on rest of disk; metasoma with sparse white setae on all sterna, denser apically on S2–S4; S5 straight along length. Male: Unknown. Comments.—This species is distinguishable from the other species with pale gray metasomal banding by the elongate face and erect, simple setae on mesepisternum, and from T. donatus by the rugose mesepisternum. The male of T. rugosus is currently unknown; the mesepisternum may not be as distinctly rugose in the males as in the females of this species. Thus, it may be that the males of T. rugosus and T. donatus cannot be separated based on the rugosity of the mesepisternum, given that the mesepisternum is sexually dimorphic in at least one other species of Triepeolus (i.e., T. pectoralis, in which the female mesepisternum is very sparsely punctate and the male is much more densely punctate and setose). Distribution.—USA: Florida, New Jersey. Floral Records.—Pontederia sp. Seasonal Records.—3 March to 9 July. Specimens examined.—7 & (GAINESVILLE, LAWRENCE, LOS ANGELES, RALEIGH, WASHINGTON D.C.).

TRIEPEOLUS SCELESTUS (Cresson) (Figs. 201, 202)

Epeolus scelestus Cresson 1878: 86–87 [Lectotype: Academy of Natural Sciences No. 2221; &, Texas]; Cresson 1916: 130 [lectotype designation]. Triepeolus scelestus; Brumley 1965: 73.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 115 Description.—Length ca. 10–12 mm; ITW 1.8–2.4 mm. Integument black, with red to orange on basal mandible and F1, sometimes on parts of legs; dorsum of mesosoma and metasoma with bands of yellow to pale yellow setae. Clypeus with faint midline and larger punctures. Paramedian band clearly separated from other pale setae on mesoscutum. Scutellum weakly bigibbous; axillar spine triangular, reaching scutellum midpoint, apically directed slightly inward. Mesepisternum lacking erect, simple setae; with dense, pale yellow, branched setae between hypoepimeron and pronotal lobe; ventrally most asetose, with dense, somewhat irregular punctures (separated by up to 1.5 puncture diameters in some areas), integument raised between punctures. T1 discal patch trapezoidal to rectangular; T2 with LLB absent, rarely much reduced, apparently forming 90 degree angle with ATB; T1–T2 with ATBs interrupted medially, posterior terga with ATBs uninterrupted. Female: Pseudopygidial area ovate, with setae uniformly coarse, darkly shining golden brown, poorly reflective and poorly differentiated from setae of rest of T5 except for rather sparse white setae apicolaterally on T5; S5 faintly downcurved apically. Male: Unknown. Comments.—This species is very similar to T. lateralis but differs in the following characters: on the female T5, there is a well-differentiated patch of white setae lateral to the pseudopygidial area in T. lateralis, while in T. scelestus these white setae are typically reduced to the apical margin of the tergum, or fully present but rather sparse and not so strongly differentiated from the pseudopygidial area; the metanotum is entirely covered with pale yellow setae in T. scelestus, while in T. lateralis these pale setae are restricted to the lateral margins of the metanotum; the ATBs on the metasoma are medially interrupted on only T1–T2 in T. scelestus, while in T. lateralis these bands are often medially interrupted at least on T1–T3; and the venter of the metasoma lacks white setae in T. scelestus, while in T. lateralis white setae are present on the apical margins of S2– S4. Distribution.—USA: Kansas, Texas (as far south as Dimmit Co.). Floral Records.—Helianthus annuus L. Seasonal Records.—10 June to 30 September. Specimens examined.—17 & (CORVALLIS, LAWRENCE, LOGAN, NEW YORK, PHILADELPHIA, SAN FRAN- CISCO).

TRIEPEOLUS SIMULATUS new species (Figs. 203–205)

Diagnosis.—This species primarily can be distinguished from all others in the genus except T. bimorulus by the scutellum, in which extensive pale yellow setae encircle one black or reddish brown region of setae on each dorsal convexity. It can be told from T. bimorulus by the clypeus and ventral half of the mesepisternum, which are shining and sparsely punctate in T. bimorulus (densely punctate and not shining in T. simulatus), and by the pseudopygidial area, the setae of which transition relatively gradually from dense and silvery basally to coarse and sparser apically in T. bimorulus (rather than a relatively abrupt, clear boundary between the two setal types in T. simulatus). Description of holotype female.—Length ca. 10.5 mm (10.0–10.5 mm); ITW 2.1 mm (1.9–2.1 mm). Integument black, with orange on basal half of mandible, labrum, scape, pedicel, F1, tegula, apex of axillar spine, and legs excluding basal halves of coxae and spurs, (also on clypeus, pronotal lobe, metepisternum, and ventral metasoma in paratype); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with faint midline, covered with dense, medially-directed white setae. Paramedian band barely laterally contiguous with pale setae on apical margin of mesoscutum. Scutellum weakly bigibbous, mostly covered with appressed setae so that only biconvexity lacks setae, forming distinctive round spot laterally on scutellum; axillar spine triangular, apical point surpassing midpoint of scutellum. Mesepisternum with sparse, short, erect, simple setae; mostly covered with dense, pale yellow, branched setae; sparser ventrally; also with small

116 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER patch brown, branched setae medioapically; ventral punctures obscured by setae, apparently nearly contiguous. T1 discal patch very widely subtriangular; T2 with LLB forming acute angle with ATB; apical bands of all terga continuous. Pseudopygidial area subquadrate, with distinct basal shining crescent; S5 straight in profile. (Male: Unknown.) Distribution.—USA: Arizona (Yuma Co.), California (Orange Co.). Seasonal Records.—1 May to 17 August. Holotype.—“U.S.A.: California, Orange County, Seal Beach Naval Weapons Station (SBNNWS) // S. Fence, Sweeping, 17 Aug [August] 1975, D. Minnesang [? name unclear] // California State Univ. Long Beach 1996 donation to Calif. Acad. Sci. // M. G. Rightmyer Triepeolus Database Specimen No. 1230 // HOLOTYPE & Triepeolus simulatus Rightmyer 2007” [California Academy of Sciences #18248] (SAN FRANCISCO). Paratype.—USA: ARIZONA: Yuma Co., 15 mi E Yuma, 1 May 1972, P. Torchio, F. Parker, G. Bohart (1&, LOGAN). Etymology.—This name is in reference to the similar appearance of this species and T. bimorulus. From the Latin, “simulatus” (imitate, copy).

TRIEPEOLUS SUBALPINUS Cockerell (Figs. 206, 207)

Triepeolus subalpinus Cockerell 1910b: 245 [Holotype: U. S. National Museum of Natural History No. 100037; &, Eldora (Boulder Co.), Colorado; 18–19 August; Grindelia (subalpina)]. Triepeolus lestes Cockerell 1921: 11 [Holotype: American Museum of Natural History No. 25093; &, Glenwood Springs (Garfield Co.), Colorado; about 39 33’N 107 20’W; 5800 ft; 22–29 July 1919]. new synonymy Triepeolus stricklandi Cockerell 1937: 86–87 [Holotype: Canadian National Collection No. 4172; &, Lethbridge, Alberta, Canada; 5 August 1935]. new synonymy

Description.—Length ca. 7–11 mm; ITW 1.4–2.3 mm. Integument black, with red on apical half of mandible; sometimes with orange on F1 and parts of legs (especially in specimens from Southern localities); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus lacking midline or with faint midline present dorsally, with faint larger punctures, sometimes covered with sparse white setae (females) or covered with dense, white, medially-directed setae (males). Paramedian band separated from or laterally contiguous with pale yellow setae on anterior margin of mesoscutum. Scutellum weakly bigibbous, somewhat extended posteriorly and sloping ventrally; axillar spine triangular (sometimes with apex rounded), not or barely reaching midpoint of scutellum. Mesepisternum with long, erect, simple setae; dorsal half covered with pale, branched setae (sparser or absent on hypoepimeron); ventrally with punctation small and dense, separated by up to a puncture width in some spots, with integument between punctures elevated, tuberculate (females) or covered with dense, white, branched setae, mostly obscuring dense, rough punctures on entire mesepisternum (males). T1 discal patch widely rectangular to ovate; T2 with LLB forming 90 degree angle with ATB. Female: Pseudopygidial area small, triangular, with uniformly silvery setae; S5 straight in profile. Mesosoma and metasoma venter black (S2–S3 with sparse pale yellow apical bands in specimens from Southern localities). Male: Pygidial plate keyhole shaped, with rather weakly defined apical downturned plate and basal transverse ridge; S2–S3 with apical bands of white setae; S4–S5 with apical fringes of brown setae. Comments.—This species is extremely similar to T. brittaini, but T. subalpinus has the paramedian band present on the mesoscutum in both sexes, and the metasomal banding is broader and the mesepisternal punctation is denser than in T. brittaini. See comments under T. brittaini for further discussion. Triepeolus subalpinus is also very similar to the T. paenepectoralis complex of species in overall appearance, due to the presence of long, erect setae on the mesepisternum, the rectangular T1 discal patch, and the

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 117 90 degree angle formed by the transverse and longitudinal bands of pale setae on T2 in these species. How- ever, females of T. subalpinus are easily differentiated from those of the T. paenepectoralis species group by their small, triangular, silvery pseudopygidial area. Distribution.—CANADA: Alberta, Saskatchewan; USA: Arizona, California (Alpine Co., El Dorado Co., Mono Co.), Colorado, Idaho, Kansas, Montana, New Mexico, North Dakota, Utah, Wyoming. Host Records.—Melissodes (Eumelissodes) agilis Cresson? (Hurd et al. 1980: adults collected on same flowers). Floral Records.—Chrysopsis villosa [= Heterotheca villosa (Pursh) Shinners var. villosa], Chrysotham- nus sp., Epilobium brachycarpum K. Presl, Grindelia squarrosa (Pursh) Dunal, Helianthus petiolaris Nutt., Heterotheca subaxillaris (Lam.) Britt. & Rusby. Seasonal Records.—16 June to 10 October. Specimens examined.—103 &, 2 % (BOULDER, DAVIS, LAWRENCE, LOGAN, NEW YORK, OTTAWA, RIV- ERSIDE, SAN FRANCISCO, URBANA, WASHINGTON D.C.).

TRIEPEOLUS SUBLUNATUS Cockerell (Figs. 208, 209)

Triepeolus sublunatus Cockerell 1907b: 62–63 [Holotype: U. S. National Museum of Natural History No. 23289; %, Dripping Spring (Organ Mountains, Dona Ana Co., New Mexico); 10 August].

Description.—Length ca. 10–12.5 mm; ITW 2.0–2.4 mm. Integument black, with orange on basal mandible, labrum, apical clypeus, scape, pedicel, F1, tegula, and part or entire legs (excluding basal coxae and spurs), sometimes on pronotal lobe; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with elevated midline, lacking larger punctures, covered with relatively sparse, long, white setae. Paramedian band tapering at anterior margin, clearly separated from other pale setae on mesoscutum. Scutellum moderately bigibbous; axillar spine triangular, not attaining midpoint of scutellum. Mesepisternum lacking erect, simple setae; dorsal half with dense, pale yellow, branched setae (sparser on hypoepimeron), and small patch of brown, branched setae between pronotal lobe and hypoepimeron; also with pale yellow, branched setae on anterior and ventral surfaces of mesepisternum; ventral half of lateral surface of mesepisternum with sparse, brown, branched setae; with dense, rough punctures (separated by up to one puncture diameter in a few spots), with integument between punctures raised and shining, tuberculate. T1 discal patch wide subovate to triangular; T2 with LLB forming acute angle with ATB; metasomal terga with pattern of bands of yellow setae resembling that of T. verbesinae. Female: Pseudopygidial area subquadrate, with fine, uniformly brown setae, apical margin faintly emarginate; S5 straight in profile. Male: Pygidial plate relatively long, lacking distinct transverse basal ridge and apical downturned plate; S2 with white apical band of setae; S3–S5 with apical fringes of setae (white and slightly less developed on S3, golden to white on S4, brown to golden on S5). Comments.—Based on the pattern of bands of yellow setae on the metasoma, this species resembles T. verbesinae (an unrelated species belonging to the T. verbesinae species group); T. sublunatus is easily distinguished from T. verbesinae by its larger size and the entirely different pseudopygidial area (i.e., not distinctly circular, with stouter, flattened setae on apical margin) and straight S5. Distribution.—USA: Arizona, New Mexico. Seasonal Records.—10 August to 11 September. Specimens examined.—6 &, 4 % (DAVIS, NEW YORK, RIVERSIDE, SAN FRANCISCO, WASHINGTON D.C.).

Triepeolus subnitens Cockerell and Timberlake 1929: 167–169 [Holotype: U. S. National Museum of Natural History No. 54849; &, Riverside, California; Helianthus annuus].

Description.—Length ca. 11–15 mm; ITW 2.2–2.9 mm. Integument black, with red on base of mandible, part or entire labrum, scape, pedicel, T1, and legs (except basally on coxae and spurs); often on part or entire clypeus; rarely on pronotal lobe, tegula, mesoscutum, and mesepisternum (possibly due to preservation?); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus shining, lacking or with very weak midline, with distinct large punctures and minute, weakly impressed punctures (sometimes covered by appressed, white setae in males). Mesoscutum shining; paramedian band clearly separated from other pale setae on mesoscutum (most females) or joined laterally with pale setae on anterior margin of mesoscutum (males and some females). Scutellum weakly bigibbous, flattened and extended posteriorly; axillar spine triangular, apically rounded, not attaining midpoint of scutellum. Mesepisternum lacking erect, simple setae; dorsal half (including hypoepimeron) with dense, pale yellow, branched setae; ventrally with very short, sparse, brown, branched setae (females and some males), or entirely covered with pale, branched setae (some males); punctures fine and not very deeply impressed, separated by up to one puncture diameter (but some nearly contiguous; punctures denser in males). T1 discal patch trapezoidal; BTB distinctly interrupted medially; T2 with LLB forming acute angle with ATB. Female: Pseudopygidial area very short and wide, seemingly formed by downturned integument, setae long and fine; T5 with relatively broad patch of white setae lateral to pseudopygidial area; S5 not or faintly downcurved. Metasomal sterna entirely brown or with lateral white setae on S2–S3 (especially in Southwestern specimens). Male: Pygidial plate relatively wide, keyhole shaped, with transverse basal ridge often hidden beneath dense brown setae and distinct, downturned apical plate; S2–S3 with white apical bands of setae (S3 with white setae surpassing apical margin); S4–S5 with brown, or brown and white apical fringes. Comments.—This species is extremely similar to T. penicilliferus but females can be distinguished by the pseudopygidial area (compare Figs. 169 and 211). Males are harder to differentiate but generally differ in the color of the pronotal lobe (red in T. penicilliferus and brown in T. subnitens), the form of the clypeus (flatter in T. penicilliferus), punctation and pubescence of the mesepisternum (less densely punctate and less setose in T. penicilliferus), and appearance of pygidial plate (narrower, with a more distinct transverse basal ridge in T. penicilliferus). See comments under T. penicilliferus for further discussion. Distribution.—MéXICO: Chihuahua, Durango; USA: Arizona, California (Tehama Co. south through Stanislaus Co. and Inyo Co. to Imperial Co.), Kansas, Nevada, New Mexico, Oklahoma, Texas (as far east as Smith Co.), Utah. Host Records.—Svastra (Epimelissodes) obliqua (Say) (Hurd et al. 1980, adult entering burrow). Floral Records.—Centaurea melitensis L., Coreopsis lanceolata L., Dyssodia aurea [=Thymophylla aurea (Gray) Greene ex Britt. var. aurea], Grindelia squarrosa (Pursh) Dunal, Gutierrezia sarothrae (Pursh) Britt. & Rusby, Helianthus annuus L., Hymenothrix wislizeni Gray, Melilotus alba [= Melilotus officinalis (L.) Lam.], Senecio douglasii [= Senecio flaccidus Less. var. douglasii (DC.) B.L. Turner & T.M. Barkl.], Verbena sp., Verbesina encelioides (Cav.) Benth. & Hook. f. ex Gray. Seasonal Records.—4 May to 12 October. Specimens examined.—112 &, 6 % (ANN ARBOR, AUSTIN, CORVALLIS, DAVIS, GAINESVILLE, LAWRENCE, LOGAN, LOS ANGELES, MEXICO CITY, NEW YORK, SAN FRANCISCO, STARKVILLE, TEMPE, TUC- SON, WASHINGTON D.C.).

Triepeolus tanneri Cockerell 1928: 232–233 [Holotype: U. S. National Museum of Natural History No. 100038; %, Farr West (Weber Co.), Utah].

Description.—Length ca. 11.5–15 mm; ITW 2.4–3.0 mm. Integument black, with red medially on mandible and outer F1; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus lacking or with faint dorsal midline, with faint larger punctures on field of very fine and dense punctures. Paramedian band reaching anterior margin of mesoscutum, separated from or nearly contiguous laterally with pale setae on anterior margin of mesoscutum. Scutellum strongly bigibbous; axillar spine triangular, reaching midpoint of scutellum, apical point slightly incurved. Mesepisternum lacking erect, simple setae; with small patch of pale yellow, branched setae posterior to pronotal lobe, otherwise mostly asetose; with very fine, nearly contiguous punctures, grading ventrally to slightly larger, more widely-spaced punctures (punctures separated by up to almost one puncture diameter in some places). T1 discal patch ovate to quadrate; T2 with LLB reduced or forming obtuse acute angle with ATB. Female: Pseudopygidial area very distinctive, long and narrow with basal patch of fine, slightly paler reflecting setae, with median, elevated, inverted U or V-shaped ridge, and apical circular patch of coarse setae; T5 entirely black; S5 tapered anteriorly and slightly downcurved. Meso- soma and metasoma venter entirely black/brown. Male: Pygidial plate keyhole shaped, with distinct transverse basal ridge and apical downturned plate; ventral metasoma with black setae; S3 apical setae very slightly surpassing apical margin; S4–S5 with true apical fringes of black setae. Comments.—Females of this species are readily distinguished by the entirely unique pseudopygidial area (See Fig. 213 and description, above). Distribution.—USA: Colorado, Kansas, Utah. Floral Records.—Helianthus salicifolius A. Dietr., Solidago sp. Seasonal Records.—26 July to 19 October. Specimens examined.—9 &, 2 % (LAWRENCE, LOGAN, URBANA, WASHINGTON D.C.).

TRIEPEOLUS TEPANECUS (CRESSON) new combination (Figs. 214–216)

Epeolus tepanecus Cresson 1878: 88 [Lectotype: Academy of Natural Sciences No. 2239; %, México]; Cresson 1916: 131 [lectotype designation].

Description.—Length ca. 10–13 mm; ITW 2.0–2.7 mm. Integument black, with reddish brown apically on mandible and outer F1; dorsum of mesosoma and metasoma with relatively narrow bands of yellow setae. Clypeus with strong midline and faint larger punctures (densely covered with white, medially directed setae in males); upper face lacking dense patches of erect setae. Mesoscutum shining; paramedian band narrow, clearly separated from other pale setae on mesoscutum. Scutellum weakly to moderately bigibbous, sloping ventrally; axillar spine triangular, usually reaching scutellum midpoint, apex strongly pointed and somewhat incurved. Mesepisternum lacking erect, simple setae; integument shining, with small, finely impressed punctures separated by one to two puncture diameters; dorsally with white, branched setae (females), or with setae somewhat sparse, and extending anteriorly and ventrally (males). T1 with bands of pale setae mostly on lateral and apical margins, forming <-shape at margin of wide, triangular-shaped discal patch; ATB widely interrupted medially; T2 lacking LLB. Female: Pseudopygidial area semicircular to subquadrate, mostly composed of long, fine, golden-reflecting setae (poorly differentiated from basal setae on T5); S2–S4 with sparse, pale setae apically; S5 straight in profile. Male: Pygidial plate keyhole shaped, with distinct basal

120 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER transverse ridge and apical downturned plate; S2–S3 with white apical bands (S3 with white setae slightly surpassing apical margin); S4–S5 with apical fringes of dark brown setae (sometimes with white laterally on S4). Comments.—This species is very similar to T. laticeps and T. mauropygus; all three species share the darkly shining pseudopygidial area, with a poorly differentiated basal region of finer, denser setae, and the sparse, short, erect, simple setae of the mesepisternum. See Table 2 for further characteristics of each species. Triepeolus tepanecus also resembles T. totonacus but can be differentiated by the pseudopygidial area, which in T. totonacus has a distinctly differentiated, and relatively long, basal region of fine, shining setae; by the mesepisternum, which entirely lacks erect, simple setae in T. totonacus; and by the T1 discal patch, which is widely ovate to trapezoidal (never triangular) in T. totonacus. Distribution.—MéXICO: Chiapas, Colima, Jalisco, Michoacán, Nayarit. Seasonal Records.—4 July to 10 December. Specimens examined.—12 &, 3 % (CHAMELA, LAWRENCE, LOGAN, NEW YORK, PHILADELPHIA, SAN FRANCISCO).

TRIEPEOLUS TEXANUS (Cresson) (Figs. 217, 218)

Epeolus texanus Cresson 1878: 87 [Lectotype: Academy of Natural Sciences No. 2223; %, Texas]; Cresson 1916: 132 [lectotype designation]. Triepeolus wyomingensis Cockerell 1905c: 201–202 [Holotype: U. S. National Museum of Natural History No. 100040; %, Wyoming]. new synonymy Triepeolus eldredi Cockerell 1907a: 52 [Holotype: U. S. National Museum of Natural History No. 100029; %, N. Yakima, Washington; 7 August 1903]. new synonymy Triepeolus helianthi grandior Cockerell 1919b: 300 [Holotype: U. S. National Museum of Natural History No. 100030; %, Florissant (Teller Co.), Colorado; 29 July 1902; Carduus (=Cirsium) acaulescens]. new synonymy Triepeolus rectangularis Cockerell 1921: 9–10 [Holotype: American Museum of Natural History No. 25091; &, Hunts- ville (near Ogden, Weber Co.), Utah; about 41° 17’N 110° 46’W; 26 July 1920]. new synonymy Triepeolus dichropus Cockerell 1921: 11 [Holotype: American Museum of Natural History No. 25094; %, Glenwood Springs (Garfield Co.), Colorado; about 39° 33’N 107° 20’W; 5800 ft; (29) July 1919]. new synonymy Triepeolus pallidiventris Cockerell and Sandhouse 1924: 308 [Holotype: California Academy of Sciences No. 1600; %, Vivian Park (Provo Canyon, Utah Co.), Utah; 7 July 1922]. new synonymy Triepeolus texanus texanus; Hurd 1979: 2096. Triepeolus nr. eldredi; Clement 1984: 300–303, Figs. 1–4 [Biological data].

Description.—Length ca. 10–13 mm; ITW 2.2–2.5 mm. Integument black, with red medially on mandible; brownish orange on F1 and sometimes on parts of legs (especially specimens from more southern localities); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with apical margin surpassing lower tangent of compound eyes by ca. 2 OD, with midline and distinct larger punctures, mostly asetose (females) or covered with dense, long, pale setae (males). Paramedian band clearly separated from other pale setae on mesoscutum, or rarely nearly absent (some females), or joined to diffuse setae on anterior margin of mesoscutum (some females, males). Scutellum weakly to moderately bigibbous, slightly extended posteriorly; axillar spine triangular, not or barely reaching midpoint of scutellum. Mesepisternum with erect, simple setae; dorsally with pale, branched setae on most of upper half except sparser on hypoepimeron, or between hypoepimeron and pronotal lobe (rarely restricted to narrow area near pronotal lobe and scant area near scrobal groove); ventral integument shining, with punctures small and dense (nearly contiguous to separated by one puncture diameter in certain areas), with sparse, short, brown, branched setae (females); or entire mesepisternum covered with dense, pale yellow, branched setae (males). T1 discal patch rectangular to subovate (lacking BTB in two females from Idaho and Washington); T2 with LLB reduced or forming 90 degree angle with ATB. Female: Pseudopygidial area semicircular to subquadrate, with distinct basal shining crescent; S5 very slightly downcurved apically. Male: Pygidial plate keyhole shaped, with distinct transverse

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 121 basal ridge and apical downturned plate; S2–S3 with apical bands of white setae (S3 with white setae slightly surpassing apical margin); S4–S5 with brown apical fringes (sometimes with white setae laterally, especially on S4). Comments.—Triepeolus texanus is similar in appearance to T. paenepectoralis, but can be distinguished from that species by the clypeus, which is elongate and has a relatively strong midline in T. texanus (the clypeus is relatively short, with a faint or absent midline in T. paenepectoralis). Triepeolus texanus is also similar in appearance to T. donatus; however, in T. donatus the bands of setae are white (rather than pale yellow), the T1 discal patch is usually more ovate than rectangular, and the pseudopygidial area’s basal shining crescent is less strongly differentiated from the more apical, coarser setae. The integument of this species is predominantly black in the Northwestern localities, while specimens from Arizona, Texas, Colorado, and Utah have more areas of red. Interestingly, of the specimens of this species collected at flowers, only various species of thistle have been recorded as the floral associate. Distribution.—CANADA: British Columbia; USA: Arizona, Colorado, Idaho, Montana, New Mexico (Guadaloupe Co.), Oregon, Texas (locality not specified), Utah, Washington, Wyoming. Host Records.—Melissodes (Eumelissodes) rustica (Say) [= Melissodes druriella (Kirby)] (Clement, 1973, larvae from nests; Clement, 1984, adults observed entering nests); Nomia melanderi Cockerell (one specimen from nesting site, Benson Ward, Utah). Floral Records.—Cirsium acaulescens (= Cirsium scariosum Nutt.), C. lanceolatum [= Cirsium vulgare (Savi) Ten.], C. texanum Buckl., C. undulatum (Nutt.) Spreng. var. undulatum. Seasonal Records.—27 May to 20 August. Specimens examined.—68 &, 2 % (BOULDER, CORVALLIS, DAVIS, GAINESVILLE, LAWRENCE, LOGAN, NEW YORK, PHILADELPHIA, RIVERSIDE, SAN FRANCISCO, URBANA, WASHINGTON D.C.).

TRIEPEOLUS TOTONACUS (Cresson) (Figs. 219, 220)

Epeolus totonacus Cresson 1878: 87 [Lectotype: Academy of Natural Sciences No. 2222; &, México]; Cresson 1916: 133 [lectotype designation]. Triepeolus totonacus; Cockerell 1905b: 165.

Description.—Length ca. 9.5–11.5 mm; ITW 2.1–2.5 mm. Integument black, with red medially on mandible; orange on F1; dorsum of mesosoma and metasoma with bands of yellow-orange setae. Clypeus with faint to strong midline and larger punctures. Paramedian band clearly separated from other pale setae on mesoscutum. Scutellum weakly to moderately bigibbous; axillar spine triangular, reaching scutellum midpoint. Mesepisternum lacking erect, simple setae; with small patches of dense, white, branched setae below scrobal groove and posterior to pronotal lobe; punctation relatively small, nearly contiguous to separated by one to two puncture diameters (up to three in some specimens); integument between punctures shining, relatively flat. T1 discal patch transversely ovate to trapezoidal, BTB and ATB interrupted medially; T2 with LLB absent or mostly on lateral surface of T2, forming weakly acute angle with ATB. Female: Pseudopygidial area subquadrate, basal half to third covered with fine, shining setae, remaining apical region with coarse, darker setae; S5 straight in profile or very slightly downcurved apically. Male: Unknown. Comments.—This species resembles other species belonging to the T. tepanecus species group (Table 2), but can be differentiated from them by the pseudopygidial area, which has a distinctly differentiated, and relatively long, basal region of fine, shining setae, and by the mesepisternum, which entirely lacks erect, simple setae. It can also be differentiated from some of those species by the T1 discal patch, which is widely ovate to trapezoidal (never triangular).

122 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER A specimen of this species was labeled “PCAM 26” by D. Yanega. Distribution.—MéXICO: Chiapas, Jalisco, Michoacán, Oaxaca, Puebla. Seasonal Records.—21 July to 5 December (majority of specimens collected in October and November). Specimens examined.—13 & (BERKELEY, CHAMELA, LAWRENCE, LOGAN, SAN FRANCISCO).

TRIEPEOLUS TOWNSENDI Cockerell (Figs. 221, 222)

Triepeolus townsendi Cockerell 1907b: 63–64 [Holotype: U. S. National Museum of Natural History No. 100039; %, Rio Ruidoso, White Mountains (Lincoln Co.), New Mexico; 27 July; about 6700 ft; Erigeron macranthus]. Triepeolus concinnus Cockerell 1917b: 300–301 [Holotype: U. S. National Museum of Natural History No. 22894; &, Meadow Valley (= Río Piedras Verdes, 9 km S. Colonia García, Chihuahua, according to Labougle, 1990: 50), Méx- ico]. new synonymy

Description.—Length ca. 8–11 mm; ITW 1.7–2.5 mm. Integument black, with dark reddish brown to orange on apical mandible and usually on outer F1, sometimes on labrum and apical clypeus; orange usually on at least part of legs (excluding coxae and spurs, often brown on at least front and middle trochanters and front femur); brownish orange on tegula; dorsum of mesosoma and metasoma with bands of yellow setae. Clypeus with distinct midline and faint larger punctures, covered with white setae in males. Paramedian band clearly separated from other pale setae on mesoscutum (both sexes). Scutellum moderately to weakly bigibbous; axillar spine triangular, reaching midpoint of scutellum. Mesepisternum apparently lacking or with short, sparse, erect, simple setae; with pale yellow, branched setae on dorsal half and ventral margins of mesepisternum, enclosing circular region of brown, branched setae (smaller brown area in specimens from New Mex- ico); punctures small, nearly contiguous to separated by 0.5 OD, with integument between raised (females), or entire mesepisternum covered with dense, white, branched setae (males). T1 discal patch trapezoidal to subovate; T2 with LLB forming acute angle with ATB. Female: Pseudopygidial area semicircular to subquadrate, with distinct, silvery basal crescent; S5 straight in profile. Male: Pygidial plate keyhole shaped, with distinct transverse basal ridge and apical downturned plate; S2–S3 with apical bands of white setae; S4–S5 with apical fringes of golden setae. Comments.—This species is similar to T. laticaudus, but can be separated from that species by the punctation of the mesepisternum (denser in T. townsendi), and by the relative differentiation of the basal shining setae on the pseudopygidial area (less differentiated in T. townsendi). Triepeolus townsendi is also similar to T. medusa, but T. medusa has longer erect, simple setae on the mesepisternum and the pseudopygidial area has a vague basal shining crescent. Distribution.—MéXICO: Chihuahua, Hidalgo; USA: Arizona, Colorado, New Mexico, Texas, Utah. Floral Records.—Apocynum sp., Cirsium sp., Cryptantha sp. Grindelia sp., Erigeron speciosus (Lindl.) DC. var. macranthus (Nutt.) Cronq., Heliopsis helianthoides (L.) Sweet var. scabra (Dunal) Fern., Ratibida columnifera (Nutt.) Woot. & Standl., Senecio sp., Vicia pulchella Kunth. Seasonal Records.—19 July to 10 September. Specimens examined.—61 &, 1 % (BOULDER, DAVIS, CORVALLIS, GAINESVILLE, LAWRENCE, LOGAN, NEW YORK, RIVERSIDE, SAN FRANCISCO, TEMPE, WASHINGTON D.C.).

TRIEPEOLUS TRISTIS (Smith) (Figs. 223, 224, 261)

Epeolus luctuosus Eversmann 1852: 101–102 [nec Spinola] [Syntype series: Zoological Institute, the Russian Academy of Sciences; Casan (= Kazan) and Orenburg, Russia (see Comments, below)]; Arnold 1885: 286–287, Figs. 1, 2

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 123 [redescription]. (not seen) Epeolus tristis Smith 1854: 258 [replacement name]. Epeolus speciosus Gerstaecker 1869: 158–159 [Holotype: Zoologisches Museum, Humboldt-Universität; %, Deut- schland Arnswalde (Pomerania) (= Choszczno, West Pomeranian Voivodship, Poland]; Bischoff 1930: 2 [synonymy]. Triepeolus tristis; Bischoff 1930: 1–2. Epeolus (Triepeolus) tristis; Warncke 1982:120.

Description.—Length ca. 7.5–11 mm; ITW 1.9–2.5 mm. Integument black, with red medially on mandible; orange on F1; dorsum of mesosoma and metasoma with bands of white setae. Clypeus lacking midline and larger punctures, usually covered with sparse white setae. Paramedian band usually laterally contiguous with diffuse white setae on anterior fourth to third of mesoscutum, rarely distinct in some females. Scutellum strongly to moderately bigibbous; axillar spine elongated and strongly pointed, apically slightly incurved, reaching posterior margin of scutellum. Mesepisternum with sparse, short, suberect, simple setae ventrally; dorsal half densely covered with white, branched setae (sparser on hypoepimeron); ventrally with sparser, black, branched setae; punctures nearly contiguous to separated by one puncture diameter, integument between punctures raised, tuberculate in appearance. T1 discal patch transversely ovate; T2 with LLB forming 90 degree angle with ATB; all metasomal terga with ATBs interrupted medially. Female: Pseudopygidial area quadrate, poorly differentiated from basal setae of T5, but with median apical setae slightly elongate, silvery; S5 straight in profile. Male: Pygidial plate keyhole shaped, with distinct apical downturned plate, but somewhat indistinct basal transverse ridge; S3–S5 with rather short apical fringes of setae brown or brown intermixed with white. Comments.—This is the only known species of Triepeolus in Europe. The following notes were supplied by Y. Pesenko (in lit., 2005), regarding the syntypes of Epeolus luctuosus in the Zoological Institute, Russian Academy of Sciences: “Epeolus luctuosus Eversmann, 1852: 101 (nec Spinola, 1851) was described from an unknown number of males and females ‘Hab. in provinciis Casanensi et Orenburgensi’. In that time, both provinces, Kazan and Orenburg, occupied a much larger territory than at the present, covering nearly all the southeast of European Russia, southern Urals, and western Kazakhstan (named "Kirgis" at that time). The lectotype of the species was not designated. In the collection of our institute, there are three syntypes of the species: 1 male labeled "Kas." [Kazan], "luctuosus" (both by Eversmann's hand); 1 male labeled "Kirgis"; 1 female labeled "Spassk" [at present, Pugaschev in Saratov province]. I have examined these syntypes, they really belong to E. tristis in the current understanding.” Distribution.—AUSTRIA: Burgenland, Tirol; ITALY: Trentino-South Tyrol; POLAND: West Pomerania; RUSSIA: Bashkir ASSR (= Bashkortostan), Tatarstan, Saratov; SLOVAKIA: Trebišov; SLOVENIA: Nitra; TUR- KEY: Erzurum. Host Records.—Tetralonia (Tetralonia) malvae Rossi? (Bischoff, 1930, no supporting evidence given); Tetraloniella (Tetraloniella) macroglossa Illiger (= Tetralonia (Tetralonia) malvae Rossi) (Westrich, 1989, no supporting evidence given); Tetraloniella (Tetraloniella) nana Morawitz (Gogala, 1999, no supporting evidence given). Floral Records.—Centaurea axillaris Willd. Seasonal Records.—15 July to 8 August. Specimens examined.—19 &, 10 % (LAWRENCE, LAWRENCE-BAKER; WASHINGTON D.C.). I have additionally seen 2 female and 6 male specimens in the collection of Hikmet Özbek, Atatürk University, Erzurum, Turkey.

TRIEPEOLUS VENTRALIS (Meade-Waldo) (Figs. 225, 226)

124 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Epeolus ventralis Meade-Waldo 1913: 96–97 [Holotype: The Natural History Museum, London 17B.502; %, Hsikou, near Tientsin (= Tianjin), China; 17 June 1906]; Yasumatsu 1933: 1, Figs. e, f, h, Plate 1 [distribution within Japan, illustrations of female S6, male pygidial plate, and dorsal habitus]; Yasumatsu 1938: 223 [placed within Triepeolus- group]. Epeolus tsushimensis (Cockerell); Bischoff 1930: 2–3 (probable misidentification). Triepeolus ventralis; Maeta et al. 1987: 26 [egg index]; Rightmyer 2004b: 256–262, Figs. 19–41 [redescription, photographs of male holotype, illustrations of male and female]. Triepeolus signatus Hedicke 1940: 345–347 [&; Kintschou (= Jinzhou), Liauhsi (= Liaoning) Province, China; July]. (not seen) new synonymy

Description.—Length ca. 8–13 mm; ITW 2.0–2.9 mm. Integument black, with red medially on mandible; brownish orange on F1; dorsum of mesosoma and metasoma with bands of white setae. Clypeus with faint midline (often stronger dorsally), with distinguishable larger punctures, often covered with sparse (females) or dense (males) white setae. Paramedian band clearly separated from other pale setae on mesoscutum, or barely contiguous with sparse anterior setae (females) or surrounded by diffuse anterior setae (males). Scutellum moderately bigibbous; axillar spine triangular, reaching midpoint of scutellum. Mesepisternum with sparse, short, suberect, simple setae ventrally; dorsal half covered with white, branched setae (sparser on hypoepimeron), venter with sparser brown, branched setae (females), or entirely covered with dense, white, branched setae (males); punctures nearly contiguous to separated by a puncture diameter in a few spots; integument between punctures raised, tuberculate. T1 discal patch transversely ovate, ATB widely interrupted medially, BTB not or barely interrupted; T2 with LLB absent. Female: Pseudopygidial area quadrate, poorly differentiated from basal setae of T5, but with median apical setae slightly elongate, silvery; S5 straight in profile. Male: Pygidial plate keyhole shaped, with strongly differentiated apical downturned plate and basal transverse ridge; S3–S5 with apical fringes of setae rather short, white (S3) grading to brown (S5). Comments.—This is the only known species of Triepeolus in eastern Asia. Although I have not seen the holotype of T. signatus, based on the original description and type locality, the name is almost certainly a junior synonym of T. ventralis. Distribution.—CHINA: Liaoning, Tianjin, [and, according to Wu Yan-ru, in lit., 2001, Guangxi, Zhe- jiang]; JAPAN: Chūbu, Kyūshū, Shikoku; RUSSIA: Jewish Autonomous Oblast, Khabarovsk Krai, Primorsky Krai (or Maritime Province). Host Records.—Tetraloniella (Tetraloniella) mitsukurii Cockerell (Maeta et al., 1996, reared from host cell)]. Seasonal Records.—17 June to 9 October. Specimens examined.—52&, 37 % (ANSFELDEN, FUKUOKA, LAWRENCE, LAWRENCE-BAKER, LONDON, TORONTO, WASHINGTON D.C.).

TRIEPEOLUS VERNUS new species (Figs. 227, 228)

Diagnosis.—This species can be distinguished by the combination of the relatively short clypeus; the pale, branched, appressed setae on the dorsal half to third of the mesepisternum and the absence of erect, simple setae on that sclerite; and the medially interrupted ATB on T1. Triepeolus vernus strongly resembles T. diffusus, but the mesoscutum is not covered with diffuse pale setae in T. vernus (as it often is in T. diffusus), and T. vernus is found in March and April, while specimens of T. diffusus from within T. vernus’ range (Texas) are apparently found in mid-July. Triepeolus vernus also resembles T. norae, but the axillar spines are not as long in T. vernus as in T. norae, and the basal silvery setae of the pseudopygidial area are less strongly differentiated from the coarser setae of the basal region in T. vernus. Triepeolus norae flies from April to October. Triepeolus vernus keys out with T. townsendi, in part of the first key above, but in T. townsendi the mesepister-

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 125 num has a medioventral spot of brown, branched setae, while in T. vernus the mesepisternum is sparsely covered with only pale yellow setae. Description of holotype female.—Length ca. 7.5 mm (7–10 mm); ITW 1.6 mm (1.5–2.0 mm). Integu- ment black, with red on basal half of mandible, entire (or part of) labrum, and apical margin (or none) of clypeus; orange on scape, pedicel, F1, pronotal lobe, tegula, and legs including spur of midleg, excluding very basal part of coxae; spurs only slightly darker than orange legs on hindleg; dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus with elevated midline and distinct larger punctures, sparsely covered with white setae (sometimes lacking sparse, white setae). Paramedian band separated from (or barely contiguous with) lateral setae. Scutellum weakly bigibbous; axillar spine triangular, almost reaching (or reaching) midpoint of scutellum. Mesepisternum lacking erect, simple setae; dorsally with dense, pale yellow, branched setae, grading to slightly sparser, pale, branched setae ventrally (sometimes with ventral setae worn partially away); integument with punctation very small, almost contiguous. T1 discal patch trapezoidal (to subovate); T2 with LLB forming acute (to weakly acute) angle with ATB. Pseudopygidial area subquadrate, basally with silvery, somewhat pilose crescent; S5 straight in profile. (Male: Unknown.) Distribution.—USA: Texas [Dimmit Co., Hidalgo Co., Maverick Co., Uvalde Co., Val Verde Co., Zap- ata Co.]. Floral Records.—Aphanostephus sp., Helenium microcephalum DC., Opuntia sp., Parkinsonia sp. Seasonal Records.—29 March to 27 April. Holotype.—“Lopeno, Tex. [Texas], IV [April]-16-1952, Michener, Beamers, Wille, LaBerge // Taken on Helenium microcephalum // M. G. Rightmyer Triepeolus Database Specimen No. 1366 // HOLOTYPE & Trie- peolus vernus Rightmyer 2007” (LAWRENCE). Paratypes.—USA: TEXAS: Dimmit Co., Chaparral Wildlife Management Area, 12–13 April 2001, A. Hook, J. Neff (4&, AUSTIN); Hidalgo Co., Bentsen-Rio Grande State Park, 29 March 1991, J. L. Neff (1&, AUSTIN); Hidalgo, 16 April 1952, Michener, Beamers, Wille, LaBerge (1&, LAWRENCE); 15 mi NW Mission, 30 March 1946, C. D. Michener, Opuntia sp. (1&, NEW YORK); Maverick Co., 1 mi E Eagle Pass, 9 April 1999, J. L. Neff (1&, AUSTIN); (Uvalde Co.), Uvalde, 14 April 1952, Michener, Beamers, Wille, LaBerge, Parkinsonia sp. (1&, LAWRENCE); Val Verde Co., Dolan Falls, 27 April 2001, J. L. Neff (5&, AUSTIN); 28 April 2001, J. L. Neff (2&, AUSTIN); (Zapata Co.), Lopeno, 16 April 1952, Michener, Beamers, Wille, LaB- erge, (1&, LAWRENCE); Helenium microcephalum (2&, LAWRENCE). Etymology.—This name is in reference to the spring flight season of this species. From the Latin, “vernus” (of springtime).

TRIEPEOLUS VICINUS (Cresson) (Figs. 229–231)

Epeolus vicinus Cresson 1865: 185 [Lectotype: Academy of Natural Sciences No. 2217; %, Cuba]; Cresson 1916: 133 [lectotype designation]. Triepeolus vicinus; Cockerell 1919a: 179; Genaro 1999: 216, Figs. 1d, 2a, 3b [redescription, illustrations of dorsal habitus, pseudopygidial area, male S7–8].

Description.—Length ca. 9–12 mm; ITW 2.0–2.5 mm. Integument black, with red sometimes on pronotal lobe and apical tip of axillar spine; red to orange on basal mandible, labrum, and clypeus; orange on scape, pedicel, F1, tegula, and legs (excluding basal coxae); dorsum of mesosoma and metasoma with bands of yellow-orange setae. Clypeus with elevated midline and faint larger punctures (obscured by rough, smaller punctation on clypeus). Paramedian band relatively long, clearly separated from other pale setae on mesoscutum. Scutellum strongly bigibbous; axillar spine pointed, reaching posterior margin of scutellum, apical point incurved. Mesepisternum lacking erect, simple setae; with dense, yellow, branched setae on margins, enclos-

126 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER ing median circular region of black, shorter, branched setae. T1 discal patch transversely ovate, rectangular, or trapezoidal, medially widened due to medial interruption of BTB and ATB; T2 with LLB mostly on lateral surface of T2, forming weakly acute angle with ATB; all metasomal tergal bands (except T6 of male) with ATBs medially interrupted. Female: Pseudopygidial area subquadrate, with fairly uniform, coarse, golden setae, basally with vague crescent of finer setae; S5 straight in profile. Male: Pygidial plate keyhole shaped, with distinct transverse basal ridge and apical downturned plate; S2–S3 with apical bands of white setae; S4– S5 with brown, to brown with white basally, apical fringes of setae. Comments.—As in T. cuabitensis, this is a Caribbean species that lacks a median notch on the posterior margin of the head (this notch found in the Caribbean species T. nisibonensis). Triepeolus vicinus can be distinguished from T. cuabitensis by its somewhat larger size and by the pseudopygidial area, which is subtriangular with a distinguishable basal crescent of shining setae (pseudopygidial area entirely brown, vaguely distinct from rest of T5 in T. cuabitensis). Distribution.—CUBA: Havana, Guantánamo (Sagua Baracoa), Pinar del Río (Viñales). Seasonal Records.—October (day unspecified). Specimens examined.—3 &, 2 % (ITHACA, LAWRENCE, PHILADELPHIA).

TRIEPEOLUS VICTORI Genaro (Figs. 232, 233)

Triepeolus victori Genaro 1998: 92–94, Figs. 1–3 [Holotype: Florida State Collection of Arthropods; &, near Filipinas Larimar Mine, Prov. Barahona, Dominican Republic; 26 June–7 July 1992].

Description.—Length ca. 15–17 mm; ITW 3.3–3.6 mm. Integument black, with orange to red on basal mandible, labrum, clypeus, scape, pedicel, F1, pronotal lobe, tegula, and legs including spurs (excluding basal halves of coxae), often on interantennal area, axillar spine, venter of metasoma, and posterior of T5; dorsum of mesosoma and metasoma with bands of yellow-orange setae. Clypeus with weak midline and larger punctures, sometimes covered with pale setae (especially in males). Paramedian band relatively long and wide, curving slightly outward at anterior margin (nearly contiguous with pale setae on anterior margin of mesoscutum in males). Scutellum strongly bigibbous; axillar spine well surpassing midpoint of scutellum, sharply pointed, with apical point directed inward. Mesepisternum with erect, simple setae; with yellow, branched setae on margins, surrounding median circular region of black, branched setae; punctures small, nearly contiguous. T1 lacking BTB, with wide, nearly continuous ATB (medially interrupted by minute, longitudinal black line), forming black basal semicircular region; T2 with LLB absent. Female: Pseudopygidial area ovate, poorly differentiated from rest of T5, composed of fine, golden setae; S5 straight in profile. Male: Pygidial plate rectangular, with transverse basal ridge and apical downturned plate; S2–S3 with apical bands of pale yellow setae (S3 with pale setae slightly surpassing apical margin); S4–S5 with apical fringes of golden setae. Comments.—Triepeolus victori is one of two robust species of Triepeolus in the Caribbean, the other one being T. wilsoni. The two species can be differentiated by the pattern of yellow bands on the metasoma, which are conspicuously medially interrupted in T. wilsoni and nearly or completely contiguous in T. victori. In addition, the T1 of T. victori lacks a basal transverse band of yellow setae, while this band is present in T. wilsoni. Distribution.—DOMINICAN REPUBLIC: Barahona, El Seibo. Seasonal Records.—26 June to 7 July. Specimens examined.—2 &, 2 % (GAINESVILLE, LOS ANGELES, NEW YORK).

Diagnosis.—This species can be distinguished by the combination of the strongly rectangular, nearly plus- sign shaped T1 discal patch; the oblique angle formed by the LLB and ATB of T2; and the uniform or nearly uniform, dark setae of the ovate pseudopygidial area. Triepeolus warriti is similar to T. cruciformis in the shape of the T1 discal patch and overall size, but differs in the punctation of the mesepisternum (with small punctures relatively evenly spaced ca. 0.5–1 puncture width apart in T. cruciformis; irregularly punctate, with tuberculate regions of integument in between in T. warriti). Triepeolus warriti additionally can be separated from T. cruciformis and T. dilutus (to which it also resembles) due to aspects of the pseudopygidial area (T. cruciformis and T. dilutus have a distinct basal region of silvery setae on the pseudopygidial area) and S5 (strongly downturned in T. cruciformis and T. dilutus; straight along length or only modestly downturned in T. warriti). Triepeolus warriti keys out with T. phaeopygus in the key above, but the pseudopygidial area of T. warriti is rounded (subquadrate in T. phaeopygus), and the ATB of T2 is interrupted medially (sometimes only barely) in T. warriti (medially continuous in T. phaeopygus). Triepeolus warriti is polymorphic in the abundance of pale setae on the mesepisternum and metasomal sterna, and in the amount that S5 is apically downturned. The latter character does not have a detectable geographic pattern associated with it. In general, specimens from Texas tend to have less pale setae than those from other localities; however, specimens with more restricted areas of pale setae can be found throughout the range of this species. Description of holotype female.—Length ca. 9 mm (7–10 mm); ITW 2.1 mm (1.5–2.1 mm). Integument black, with red medially on mandible; orange on labrum, apical margin of clypeus, scape, pedicel, F1, F2, tegula, and legs excluding basal halves of coxae and spurs (and sometimes parts of front leg) (sometimes black on labrum, part of or entire clypeus, scape, pedicel, F2, and tegula); dorsum of mesosoma and metasoma with bands of pale yellow setae. Clypeus lacking midline, with faint larger punctures. Paramedian band separated from (or contiguous with) lateral setae. Scutellum weakly bigibbous; axilla triangular, almost reaching midpoint of scutellum. Mesepisternum lacking erect, simple setae; dorsal half (at least) with distinct region of pale yellow, branched setae; these pale branched setae extending down anterior surface of mesepisternum (usually); ventrally with short, black, branched setae (or with sparse pale setae); punctation relatively small, dense, and irregular, integument between raised and somewhat tuberculate, punctures separated by up to 1.5 puncture diameters. T1 discal patch strongly rectangular, nearly plus-sign shaped when considered with median interruption of BTB and ATB; T2 with LLB forming obtuse angle with ATB. Pseudopygidial area widely ovate, setae uniformly golden, fine, relatively long (sometimes with vague, small basal region of more silvery-reflecting setae); S5 straight in profile (or modestly downcurved apically); S2–S4 with medially continuous apical bands of white setae (ranging to only S4 with small lateral patch white setae in some specimens from AZ). (Male: Unknown.) Distribution.—MéXICO: Chihuahua, Durango, Zacatecas; USA: Arizona, New Mexico, Texas. Floral Records.—Amphiachyris dracunculoides (DC.) Nutt., Baccharis glutinosa [= Baccharis salicifolia (Ruiz & Pavón) Pers.], Sphaeralcea hastulata Gray. Seasonal Records.—17 August to 24 October. Holotype.—“ARIZ. [Arizona]: Portal, 4800 ft., 16 Sept. [September] 1965 // C. W. Sabrosky collector // M. G. Rightmyer Triepeolus Database Specimen No. 6811 // HOLOTYPE & Triepeolus warriti Rightmyer 2007” (WASHINGTON D.C.). Paratypes.—USA: ARIZONA: Cochise Co., 5 mi SW Apache, 17 August 1961, M. A. Cazier, Baccha- ris glutinosa (4&, NEW YORK); 5.6 mi SW Apache, 14 September 1999, Rozen, LeBuhn, Rightmyer (1&, NEW YORK); 14 mi SW Apache, 20 August 1996, A. Pence, J. G. & B. L. Rozen (1&, NEW YORK); 18 mi SW

128 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Apache, 28 August 1994, J. G. Rozen, J. S. Ascher (1&, NEW YORK); Gila Co., 5 km SE Pine, 11 September 1986, F. Parker, T. Griswold (1&, LOGAN); Payson, 7 September 1953, Timberlake (1&, WASHINGTON D.C.); Yavapai Co., 31 km E Camp Verde, 12 September 1995, Griswold, Tepedino (1&, LOGAN); NEW MEXICO: Hidalgo Co., 26 mi S Animas, 20 August 1986, R. J. McGinley, B. M. Norden (1&, WASHINGTON D.C.); 33 mi S. Animas, 29 August 1982, J. G. & B. L. Rozen (1&, NEW YORK); Rodeo, 2 September 1973, M. Favreau (2&, NEW YORK). Additional specimens.—MÉXICO: CHIHUAHUA: 6 km E Ciudad Jiménez, 21 August 1991, R. L. Minckley, Sphaeralcea hastulata (1&, LAWRENCE); 10 mi N Jiménez, 21 September 1970, G. E. & R. M. Bohart (2&, LOGAN); DURANGO: 1 mi N Registro, 17 September 1950, R. F. Smith (1&, NEW YORK); ZACATECAS: 5 mi N Tobasco, 18 September 1970, G. E. & R. M. Bohart (2&, LOGAN); USA: TEXAS: (Bexar Co.), ca. 10 mi NE San Antonio, 24 October 1968, A., A. Gillogly (1&, RIVERSIDE); San Antonio, 24 October 1923, F. M. Jones (2&, NEW YORK); Culberson Co., 32 mi E Van Horn, 27 September 1999, J. L. Neff, Amphiachryis dranunculoides (1&, AUSTIN); Davis Co., 20 mi N Ft. Davis, 26 September 1950, W. Gertsch, M. Cazier, 4000 ft. (1&, NEW YORK); (Tom Green Co.), 30 mi S San Angelo, 10 October 1968, Hud- dleston, Ward (1&, LOS ANGELES). Etymology.—This species is named in honor of Natapot Warrit for his support and kindness, especially during the last year of my dissertation work.

TRIEPEOLUS WILSONI (Cresson) (Figs. 237, 238)

Epeolus wilsoni Cresson 1865: 183–184 [Lectotype: Academy of Natural Sciences No. 2216; %, Cuba]; Cresson 1916: 134 [lectotype designation]. Triepeolus wilsoni; Cockerell 1919a: 179–180; Genaro 1998: Figs. 1b, 2a, b [illustrations of dorsal habitus, male S7–8]; Genaro 1999: 216, Figs. 1a, 3a [redescription, illustrations of dorsal habitus, male S7–8]. Triepeolus buscki Cockerell 1919a: 179–180 [Holotype: U. S. National Museum of Natural History No. 20712; &, Bara- coa (Guantánamo Prov.), Cuba; August 1902]; Genaro 1999: 216 [synonymy].

Description.—Length ca. 15–16 mm; ITW 3.13.3 mm. Integument black to reddish brown, with orange on basal mandible, labrum, clypeus, interantennal area, frons near compound eye, entire antenna or basal to F2, and parts of legs (especially distal to femora); dorsum of mesosoma and metasoma with bands of yellow- orange setae. Clypeus with midline on dorsal half and vague larger punctures (covered with pale setae in males). Paramedian band curving outward at anterior margin and clearly separated from other pale setae on mesoscutum (females; nearly contiguous with yellow setae on anterior margin of mesoscutum in males). Scutellum strongly bigibbous; axillar spine well surpassing midpoint of scutellum, very sharply pointed, curving inward at apical point. Mesepisternum with erect, simple setae; with yellow, branched setae at margins surrounding darker circular area of short, brown, branched setae (pale setae absent on hypoepimeron), punctures dense, with integument between raised, somewhat tuberculate. T1 discal patch transversely ovate to triangular; BTB and ATB interrupted medially; T2 with LLB absent; metasomal terga posterior of T1 with ATBs widely interrupted medially. Male: Pygidial plate keyhole shaped, with distinct transverse basal ridge and apical downturned plate; S2–S3 with apical bands of white setae (slightly surpassing apical margin on S3); S4–S5 with apical fringes of golden setae. Comments.—This species is similar to T. victori. The two species can be differentiated by the pattern of yellow bands on the metasoma, which are conspicuously medially interrupted in T. wilsoni and nearly or completely contiguous in T. victori. In addition, the T1 of T. victori lacks a basal transverse band of yellow setae, while this band is present in T. wilsoni. Distribution.—CUBA: Pinar del Rio, San Vicente. Seasonal Records.—June (day unspecified) to August (day unspecified).

TRIEPEOLUS ZACATECUS (Cresson) new combination (Figs. 239, 240)

Epeolus zacatecus Cresson 1878: 85 [Lectotype: Academy of Natural Sciences 2240; &, México]; Cresson 1916: 134 [lectotype designation].

Description.—Length ca. 14–17 mm; ITW 3.2–3.5 mm ITW, rarely as small as 11 mm in length, 2.3 mm. Integument black, with reddish orange medially on mandible and outer portion of F1; dorsum of mesosoma and metasoma with bands of yellow-orange to yellow setae (paler in males). Clypeus with elevated midline and faint larger punctures (females) or densely covered with white setae (males). Paramedian band wide, but clearly separated from other pale setae on mesoscutum (most females) or laterally contiguous with yellow setae on anterior mesoscutum (some females, males). Scutellum strongly to moderately bigibbous; axillar spine surpassing midpoint of scutellum, sharply-pointed, anterior point curving inward; inner margin outlined with white setae. Mesepisternum with sparse, erect, simple setae; dorsal half covered with dense, pale yellow, branched setae, ventrally with sparser, black, branched setae (females) or densely covered with white, branched setae (males); punctures small, deeply impressed so that integument between appears tuberculate, separated by up to one puncture diameter. T1 discal patch ovate, triangular, or trapezoidal; T2 with LLB forming 90 degree angle with ATB, or enlarged to form semicircular basal area of dark brown setae. Female: Pseudopygidial area ovate, formed of nearly uniformly long, darkly shining setae; S5 straight in profile. Male: Pygidial plate relatively wide, with distinct transverse basal ridge (sometimes covered by dark setae) and apical downturned plate; S3–S5 with apical fringes of setae, white and sometimes reduced on S3, brown to pale golden on S4–S5. Comments.—This species is very similar to T. grandis; the two species are only separable (usually) by the orange legs, paler yellow bands of setae, narrower bands of yellow setae on T1 and T2, and shorter axillar spines in T. grandis. (See comments under T. grandis for further discussion.) This species is superficially similar to T. kathrynae (in the T. simplex species group), due to the very long axillar spines, the yellow coloration, and the robust body form; however females can easily be separated by the entirely different pseudopygidial areas (apically concave in T. kathrynae). This species was labeled “PCAM 34” by T. Griswold. Distribution.—COSTA RICA: Guanacaste; MéXICO: Chiapas, Colima, Jalisco, Michoacán, Oaxaca, San Luis Potosí, Sinaloa, Sonora, Tamaulipas, Veracruz. Floral Records.—Croton sp. Seasonal Records.—7 June to 7 December. Specimens examined.—42 &, 7 % (BERKELEY, CHAMELA, DAVIS, GAINESVILLE, HEREDIA, LAWRENCE, LOGAN, NEW YORK, PHILADELPHIA, SAN FRANCISCO, WASHINGTON D.C.).

Triepeolus simplex species group

Diagnosis.—This species group is known by the combination of the following characters: the pseudopygidial area is concave or sometimes nearly straight on the apical margin (Fig. 2b), sometimes with stout setae laterally near the apical margin; and the female S5 is straight in profile, with bristle-like setae on the apical margin (Fig. 2c). In addition, the following characters are typical, but not necessarily diagnostic for the group: the clypeus usually lacks a midline, and the axillar spine is often pointed, reaching the midpoint of the scutellum.

130 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Included species.—T. alvarengai Moure, T. kathrynae Rozen, T. mexicanus (Cresson), T. obliteratus (Graenicher), T. rhododontus Cockerell, T. roni Genaro, T. saturninus Cockerell and Sandhouse, T. segregatus (Cockerell), T. simplex Robertson. Comments.—This group is primarily found in North America, from Saskatchewan in Canada, throughout most of the United States, and south through Mexico. There are seven described species in North America, with approximately seven or eight additional species yet to be named from the western United States. Trie- peolus roni is found in the Caribbean, T. mexicanus ranges into Central America, and T. alvarengai is the only currently-known species from South America (Brazil).

Triepeolus verbesinae species group

Diagnosis.—This species group is known by the unique form of the pseudopygidial area, which is distinctly circular, with setae on the apical margin that are denser and more flattened than those found medially on the area (Fig. 2a). In addition, the following characters are typical, but not necessarily diagnostic for the group: the clypeus usually lacks a midline; the scutellum usually has a pronounced posterior, transverse ridge; and the paramedian band often tapers anteriorly, meeting the anterior margin of the mesoscutum. In North Amer- ica (but not in South America), this group is usually additionally characterized by the female S5, which is strongly downturned; the axillae, which are often rounded apically and do not attain the midpoint of the scutellum in length; and the LLB of T2, which forms a strongly acute angle with the ATB. A distinctive species of the North American T. verbesinae species-group is T. verbesinae, which has the bands of pale setae medially interrupted on all terga. In South America, many species of this group similarly have medially interrupted bands of pale setae on most or all of the metasomal terga. Included species.—T. aguilari Moure, T. atoconganus Moure, T. bilineatus Cockerell, T. buchwaldi (Friese), T. callopus Cockerell, T. flavipennsis (Friese), T. grindeliae Cockerell, T. inyoensis Cockerell & Sandhouse, T. osiriformis (Schrottky), T. rufotegularis (Ashmead), T. verbesinae (Cockerell), T. schwarzi Cockerell, T. timberlakei Cockerell. Comments.—This species group constitutes a major component of the Triepeolus fauna found in South America, with six described species and an additional undescribed species from Venezuela. Of the South American species, T. osiriformis and T. rufotegularis are also known from the Caribbean. Triepeolus bilineatus ranges from Guatemala and Honduras to Texas, and an additional six described species are known from North America. There are approximately six to eight additional species yet to be described from North Amer- ica, primarily from the western United States.

Unplaced species names

The following species names were not associated with any female specimens available to me, and were not definitively placeable within the T. verbesinae or T. simplex species groups. In many cases, these names are based on male holotypes or the type material was missing.

TRIEPEOLUS CUNEATUS Cockerell Triepeolus cuneatus Cockerell 1917b: 300 [Holotype: U. S. National Museum of Natural History No. 22893; %, Meadow Valley (= Rio Piedras Verdes, 9 km S. of S. Colonia García, Chihuahua, according to Labougle, 1990: 50), México].

TRIEPEOLUS HOPKINSI Cockerell Triepeolus hopkinsi Cockerell 1905b: 184 [%; Grand Canyon of the Colorado, Arizona; 3 August 1904] [Type lost?].

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 131 TRIEPEOLUS ISOCOMAE Cockerell Triepeolus isocomae Cockerell 1904: 38 [Holotype: U. S. National Museum of Natural History No. 9706; %, Albuquer- que New Mexico; 16 September; Bigelovia (=Isocoma) wrightii].

TRIEPEOLUS? MERCATUS (Fabricius) Epeolus mercatus Fabricius 1804: 389 [Type lost?]; Cresson 1878: 88 [redescription]. Comments.—Epeolus mercatus cannot be confidently assigned to either Epeolus or Triepeolus, although the original description suggests a species similar to T. helianthi. There does not appear to be any consensus among previous workers as to the identity of this species, as evidenced by numerous, different species identified as T. mercatus in the collections examined for this study. See further comments under T. remigatus.

TRIEPEOLUS NEMORALIS (Holmberg) Doeringiella nemoralis Holmberg 1886: 278, 280 [&, Chaco, Formosa (Argentina), March]. (type not seen; location unknown?) Triepeolus nemoralis; Roig-Alsina 1989: 578. Comments.—According to the original description, based on a female specimen from northeastern Argentina, this species is black with a layer of gold hairs; with red on the first three antennal segments, last two metasomal segments, and legs; the T1 has the “lateral bands” narrowed medially; T2–T4 have gold bands reaching their margins; T5 is entirely golden; the venter has pale yellow setae at the margins of the segments; all the “sutures” of the mesosoma are covered with dirty white, short, appressed setae; and the “mesonotum” (=mesoscutum) has spots (=paramedian bands) tapering anteriorly.

TRIEPEOLUS POMONALIS Cockerell Tniepeolus (sic) pomonalis Cockerell 1916: 392–393 [Holotype: (supposedly Pomona College No. 160); %, Claremont, California] [Type lost?]. Comments.—The holotype of T. pomonalis is listed in the original publication as belonging to Pomona College, in Claremont California, but apparently is not presently there (Jonathan Wright, in lit., 2005), nor is it in any of the institutions mentioned in the Acknowledgements and Materials sections herein. Unfortunately, the identity of this species is not clear based solely on the original description.

TRIEPEOLUS SCELESTUS TUBERCULARIS Brues Epeolus scelestus tubercularis Brues 1903: 82 [&; Austin, Texas] [Type lost?]. Comments.—It is likely that T. scelestus tubercularis is synonymous with T. scelestus, but this cannot be confidently assumed based on the description alone.

TRIEPEOLUS SEQUIOR Cockerell Triepeolus sequior Cockerell 1921: 8–9 [Holotype: American Museum of Natural History No. 25040; %, Ridgeway Col- orado; 15 July 1919].

Acknowledgements

I thank John S. Ascher, James S. Ashe, Bryan N. Danforth, Michael S. Engel, Terry L. Griswold, Charles D. Michener, Laurence Packer, Jerome G. Rozen, Jr., and Natapot Warrit for advice and discussions that have greatly improved this work. Additionally, I thank Charles D. Michener, Jerome G. Rozen, Jr., Sam Droege, Terry L. Griswold, Rebekah Nelson, Marie Wendel, and especially Laurence Packer for offering feedback on various versions of the keys to Triepeolus species. I thank Michael S. Engel and James S. Ashe for making their Microptics camera systems available for my work, and Matthew L. Buffington for use of his EntoVision camera system. Robert W. Brooks, Michael S. Engel, Charles D. Michener, and Zachary H. Falin are gratefully acknowledged for facilitating loans from many of the institutions listed in the Materials section. I thank

132 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER John S. Asher for providing access to primary literature housed at the American Museum and for bringing several host associations to my attention. Elaine R. S. Hodges is gratefully acknowledged for permission to borrow her masterful illustrations of Triepeolus, which she created for Paul Hurd during his tenure at the Smithsonian Institution (Frontispiece and Figs. 241–263). For providing these illustrations, I thank Scott Miller and George Venable, also of the Smithsonian Institution. Matthew L. Buffington, Darci Falin, and Sara L. Taliaferro generously provided instruction and resources for the completion of the photographs and figures found herein. Favizia F. de Oliveira and Gabriel A. R. Melo are thanked for providing photographs of specimens found in Curitiba, Brazil. Caleb Morse is thanked for his expert advice regarding plant taxonomy and nomenclature. I am tremendously grateful to David G. Furth and Ted R. Schultz for providing me with space and resources at the Smithsonian for the completion of this study. I am greatly indebted to each of the individuals who loaned specimens used in this study; the list of those individuals can be found in the Materials section. During my search for missing holotypes, a number of people kindly provided information about specimens found in their respective institutions. For this service I thank C. van Achterberg (Nationaal Natuurhistorisch Museum, Leiden, The Netherlands), Hannes Baur (Naturhistorische Museum der Burgergemeinde, Bern, Switzerland), Christy Bills (University of Utah Museum of Natural History, Salt Lake City), C. Roberto F. Brandão and Beatriz Coelho (Museu de Zoologia da Universidade de São Paulo, Brazil), Daniel Burckhardt (Naturhistorisches Museum, Basel, Switzerland), Michael S. Caterino (Santa Barbara Museum of Natural History, California), Paula Cushing (Denver Museum of Nature and Science, Colorado), Roy Danielsson (Zoologiska museet, Lunds Universitet, Sweden), Ismael Hinojosa-Díaz (Universidad Nacional Autónoma de México, Mexico City), Chris Durden (Texas Memorial Museum, Austin), Willem Hogenes (Instituut voor Taxonomische Zoölogie, Amsterdam, The Netherlands), Joe Keiper and Thomas Pucci (Cleveland Museum of Natural History, Ohio), Jens-Peter Kopelke (Fors- chungsinstitut und Naturmuseum Senckenberg, Frankfurt, Germany), Steve Matter (Cincinnati Museum of Natural History, Ohio), Donald McFarlane and Jonathan Wright (Joint Sciences Department and Pomona Col- lege, of the Claremont Colleges, California), Yuri A. Pesenko (Zoological Institute, the Russian Academy of Sciences, St. Petersburg), Raymond J. Pupedis (Peabody Museum of Natural History, Yale University, Con- necticut), Leslie L. Skibinski (Delaware Museum of Natural History, Wilmington), Lars Vilhelmsen (Zoolo- gisk Museum, Københavns Universitet, Denmark), Claire Villemont (Musée National d'Histoire Naturelle, Paris, France), Kate Wellspring (Amherst College, Pratt Museum of Natural History, Massachusetts), Wu Yan-ru (Academia Sinica, Beijing, China), and Herbert Zettel (Naturhistorisches Museum, Wien, Austria). I thank the following people and organizations for funding or otherwise supporting my Triepeolus research: the American Museum of Natural History; the Philanthropic Educational Organization (Chapter MH, Missouri); the National Science Foundation; the United States Department of Agriculture; the United States National Museum of Natural History; the University of Kansas Graduate School, Natural History Museum and Biodiversity Research Center, and Entomology Program Endowment; Charles and Jennie Right- myer; and Clayton and Margaret Depue. This paper is based upon work supported by a National Science Foundation Graduate Research Fellowship.

References

Arnold, N. (1885) Apum Mohileviensium species parum cognitae vel imperfecte descriptae. Horae Societatis Entomo- logicae Rossicae, sermonibus in Rossia usitatis editae, 19, 282–287. Ayala, R. (1988) Abejas silvestres (Hymenoptera: Apoidea) de Chamela, Jalisco, Mexico. Folia Entomológica Mexi- cana, 77, 395–493. Ayala, R. (1999) Revision de las abejas sin aguijon de Mexico (Hymenoptera: Apidae: Meliponini). Folia Ento- mológica Mexicana, 106, 1–123. Bischoff, H. (1930) Beitrag zur Kenntnis paläarktischer Arten der Gattung Epeolus (Hym. Apid.). Deutsche Entomolo- gische Zeitschrift, 1930, 1–15.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 133 Bohart, G.E. (1966) Notes on Triepeolus remigatus (Fabricius), a “cuckoo bee” parasite of the squash bee, Xenoglossa strenua (Cresson) (Hymenoptera: Apoidea). Pan-Pacific Entomologist, 42, 255–262. Bohart, G.E. (1970) The Evolution of Parasitism Among Bees. 41st Faculty Honor Lecture. Utah State University, Logan, Utah, ii + 30 pp. Brues, C.T. (1903) Studies of Texan bees. Part 1. Entomological News, 14, 79–85. Brumley, R.L. (1965) A Revision of the Bee Genus Epeolus Latreille of Western America North of Mexico. M.S. thesis. Utah State University, Logan, Utah, iii + 92 pp. Cameron, P. (1907) Description of a new species of Epeolus from Mexico. (Hym.). Zeitschrift für Systematische Hymenopterologie und Dipterologie, 7, 136. Cane, J.H. (1995) Notes on the nesting biology of Svastra atripes atrimitra (LaBerge) (Hymenoptera: Apidae). Journal of the Kansas Entomological Society, 68, 238–240. Clement, S.L. (1973) The nesting biology of Melissodes (Eumelissodes) rustica (Say), with a description of the larva (Hymenoptera: Anthophoridae). Journal of the Kansas Entomological Society, 46, 516–525. Clement, S.L. (1984) Observations on the behavior of Triepeolus nr. eldredi Cockerell (Hymenoptera: Anthophoridae). Pan-Pacific Entomologist, 60, 300–303. Cockerell, T.D.A. (1898) Tables for the determination of New Mexico bees. Bulletin of the University of New Mexico, 1, 41–73. Cockerell, T.D.A. (1900) Notes on New Mexico bees. Canadian Entomologist, 32, 361–364. Cockerell, T.D.A. (1903) Some North American bees: Osmia and Triepeolus. Entomological News, 14, 331–333. Cockerell, T.D.A. (1904) Some parasitic bees. Annals and Magazine of Natural History (7), 13, 33–42. Cockerell, T.D.A. (1905a) Notes on bees in the British Museum. Transactions of the American Entomological Society, 31, 309–364. Cockerell, T.D.A. (1905b) Une nouvelle abeille du Mexique. Bulletin du Muséum d’Histoire Naturelle, 1905, 165. Cockerell, T.D.A. (1905c) The bees of Southern California. Bulletin of the Southern California Academy of Sciences, 4, 99–106. Cockerell, T.D.A. (1906) The bees of New Mexico. Transactions of the American Entomological Society, 32, 289–314. Cockerell, T.D.A. (1907a) Two new bees of the genus Triepeolus. Canadian Entomologist, 39, 51–52. Cockerell, T.D.A. (1907b) Descriptions and records of bees.—XV. Annals and Magazine of Natural History (7), 20, 59–68. Cockerell, T.D.A. (1907c) New American bees.—III. Entomologist, 40, 135–138. Cockerell, T.D.A. (1910a) New American bees.—IX. Entomologist, 43, 90–92. Cockerell, T.D.A. (1910b) Some bees from Eldora, Colorado. Psyche, 17, 244–247. Cockerell, T.D.A. (1911) Descriptions and records of bees.—XXXIX. Annals and Magazine of Natural History (8), 8, 660–673. Cockerell, T.D.A. (1916) Some California bees. Canadian Entomologist, 48, 391–393. Cockerell, T.D.A. (1917a) Descriptions and records of bees.—LXXV. Annals and Magazine of Natural History (8), 19, 473–481. Cockerell, T.D.A. (1917b) Descriptions and records of bees.—LXXVII. Annals and Magazine of Natural History (8), 20, 298–304. Cockerell, T.D.A. (1919a) Bees in the collection of the United States National Museum.—3. Proceedings of the United States National Museum, 55, 167–221. Cockerell, T.D.A. (1919b) The bees of Peaceful Valley, Colorado. Journal of the New York Entomological Society, 27, 298–300. Cockerell, T.D.A. (1921) The epeoline bees of the American Museum Rocky Mountain expeditions. American Museum Novitates, 23, 1–16. Cockerell, T.D.A. (1925a) Descriptions and records of bees.—CVII. Annals and Magazine of Natural History (9), 16, 621–629. Cockerell, T.D.A. (1925b) Descriptions and records of bees.—CIV. Annals and Magazine of Natural History (9), 15, 489–496. Cockerell, T.D.A. (1928) Some bees from Utah. Psyche, 35, 232–234. Cockerell, T.D.A. (1931) Two genera of bees new to the recorded fauna of Nova Scotia. Canadian Entomologist, 43, 279. Cockerell, T.D.A. (1937) The bees of Alberta.—III. Canadian Entomologist, 59, 86–89. Cockerell, T.D.A. (1939) The bees of the Southern California Islands. Proceedings of the California Academy of Sci- ences, 23, 427–436. Cockerell, T.D.A. (1949) Bees from Central America, principally Honduras. Proceedings of the United States National Museum, 98, 429–490. Cockerell, T.D.A. & Sandhouse, G. (1924) Parasitic bees (Epeolinae and Melectinae) in the collection of the California Academy of Sciences. Proceedings of the California Academy of Sciences, 13, 305–324.

134 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Cockerell, T.D.A. & Timberlake, P.H. (1929) Two new bees of the genus Triepeolus. Pan-Pacific Entomologist, 5, 167–169. Commonwealth Institute of Entomology. (1962) Section 13. Insecta. In: Vevers, H. G. (Ed), The Zoological Record. Zoological Society of London, London, England, pp. 1–440. Cresson, E.T. (1865) Hymenoptera of Cuba. Proceedings of the Entomological Society of Philadelphia, 4, 1–200. Cresson, E.T. (1878) Descriptions of new North American Hymenoptera in the collection of the American Entomologi- cal Society. Transactions of the American Entomological Society, 7, 61–136. Cresson, E.T. (1916) The Cresson types of Hymenoptera. Memoires of the American Entomological Society, 1, 1–141. Custer, C.P. (1928) On the nesting habits of Melissodes Latr. (Hymenop.). Canadian Entomologist, 50, 28–31. Dalla Torre, C.G.de. (1896) Catalogus Hymenopterorum, Vol. 10, Apidae (Anthophila). Engelmann, Leipzig, Germany, viii + 643 pp. Engel, M.S. (2001) A monograph of the Baltic amber bees and evolution of the Apoidea (Hymenoptera). Bulletin of the American Museum of Natural History, 259, 1–192. Eversmann, E. (1852) Fauna hymenopterologica Volgo-Uralensis. Bulletin de la Société impériale des Naturalistes de Moscou, 25, 1–137. Fabricius, J.C. (1804) Systema Piezatorum, secundum Ordines, Genera, Species adiectis Synonymis, Locis, Observa- tionibus, Descriptionibus. Carolum Reichard, Brunsvigae [Brunswick], Germany, xiv + [15]–[440] + [1]–30 pp. Fox, W.J. (1891) On a collection of Hymenoptera made in Jamaica during April, 1891. Transactions of the American Entomological Society, 18, 337–348. Friese, H. (1908) Die Apidae (Blumenwespen) von Argentina nach den Reisenergebnissen der Herren A. C. Jensen- Haarup und P. Jörgensen in den Jahren 1904–1907. Flora og Fauna, 1908, 1–111. Friese, H. [1917(1916)]. Zur Bienenfauna von Costa Rica. (Hym.). Stettiner Entomologische Zeitung, 77, 287–339. Friese, H. (1925) Neu neotropische Bienenarten, zugleich II. Nachtrag zur Bienenfauna von Costa Rica. Stettiner Ento- mologische Zeitung, 86, 1–41. Genaro, J.A. (1998) Especie nueva de Triepeolus para la República Dominicana (Hymenoptera: Apidae). Caribbean Journal of Science, 34, 92–94. Genaro, J.A. (1999) Revisión del género Triepeolus en Cuba (Hymenoptera: Apidae), con descripción de dos especies nuevas. Caribbean Journal of Science, 35, 215–220. Genaro, J.A. (2001) Especies nuevas de abejas de Cuba y La Española (Hymenoptera: Colletidae, Megachilidae, Api- dae). Revista de Biología Tropical, 49, 1027–1035. Gerstaecker, A. (1869) Beiträge zur näheren Kenntniss einiger Bienen-Gattungen. Entomologische Zeitung heraus- gegeben von dem entomologischen Vereine zu Stettin, 30, 139–184. Gogala, A. (1999) Bee fauna of Slovenia: Checklist of species (Hymenoptera: Apoidea). Scopolia, 42, 1–79. Graenicher, S. (1905) Some observations on the life history and habits of parasitic bees. Bulletin of the Wisconsin Nat- ural History Society, 3, 153–167. Graenicher, S. (1928) New bees from the Miami region of Florida (Hymen.: Andrenidae, Megachilidae). Entomologi- cal News, 39, 279–284. Hedicke, H. (1940) Ueber palaarktische Apiden. (Hym.) II. Sitzungsberichte der Gesellschaft Naturforschender Fre- unde zu Berlin, 1939, 335–350. Hicks, C.H. (1926) Nesting habits and parasites of certain bees of Boulder County, Colorado. University of Colorado Studies, 26, 217–252. Holmberg, E.L. (1886) Sobre ápidos Nómadas de la República Argentina. Anales de la Sociedad Científica Argentina, 22, 272–286. Hurd, P.D., Jr. (1979) Superfamily Apoidea. In: Krombein, K. V., Hurd, P. D., Jr., Smith, D. R., & Burks, B. D. (Eds), Catalog of Hymenoptera in America North of Mexico, Vol. 2. Smithsonian Institution Press, Washington, D. C., pp. 1741–2209. Hurd, P.D., Jr. & Linsley, E.G. (1959) Observations on the nest-site behavior of Melissodes composita Tucker and its parasites, with notes on communal use of nest entrances (Hymenoptera: Apoidea). Entomological News, 70, 141– 146. Hurd, P.D., Jr., LaBerge, W.E. & Linsley, E.G. (1980) Principal sunflower bees of North America with an emphasis on the Southwestern United States (Hymenoptera: Apoidea). Smithsonian Contributions to Zoology, 310, i–iv, 1–158. Labougle, J.M. (1990) Bombus of México and Central America (Hymenoptera, Apidae). University of Kansas Science Bulletin, 54, 35–73. Lepeletier de Saint-Fargeau, A.L.M. & Serville, A. (1825) [Article] In: Diderot, M., et al., [Latreille, P. A. (Ed)], Ency- clopédie Méthodique. Histoire Naturelle. Entomologie, ou Histoire Naturelle des Crustaces, des Arachnides et des Insectes, Vol. 10. Paris, France, 1–344. Linsley, E.G. (1939) Some new genera and species of epeoline and nomadine bees (Hymenoptera, Nomadidae). Pan- Pacific Entomologist, 15, 1–11. Linsley, E.G. (1962) Sleeping aggregations of the Aculeate Hymenoptera—II. Annals of the Entomological Society of

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 135 America, 55, 148–164. Linsley, E.G. & MacSwain, J.W. (1942) The parasites, predators, and inquiline associates of Anthophora linsleyi. Amer- ican Midland Naturalist, 27, 402–417. Linsley, E.G. & Michener, C.D. (1939) A revision of the North American Nomadidae (Hymenoptera). Transactions of the American Entomological Society, 65, 265–305, pls xv–xviii. Lovell, J.H. & Cockerell, T.D.A. (1905) The nomadine and epeoline bees of southern Maine. Psyche, 12, 39–42. Maeta, Y., Kubota, N. & Sakagami, S.F. (1987) Nomada japonica as a thelytokous cleptoparasitic bee, with notes on egg size and egg complement in some cleptoparasitic bees. Kontyû, 55, 21–31. Maeta, Y., Gôukon, K., Sugiura, N. & Miyanaga, R. (1996) Host records of cleptoparasitic bees in Japan (Hymenoptera, Apoidea). Japanese Journal of Entomology, 64, 830–842. Meade-Waldo, G. (1913) Notes on the Apidae (Hymenoptera) in the collection of the British Museum, with descriptions of new species. Annals and Magazine of Natural History (8), 12, 92–103. Michener, C.D. (1944) Comparative external morphology, phylogeny, and a classification of the bees (Hymenoptera). Bulletin of the American Museum of Natural History, 82, 151–326. Michener, C.D. (2000) The Bees of the World. Johns Hopkins University Press, Baltimore, Maryland, xiv + [1] + 913 pp. Minckley, R.L., Wcislo, W.T., Yanega, D. & Buchmann, S.L. (1994) Behavior and phenology of a specialist bee (Dieu- nomia) and sunflower (Helianthus) pollen availability. Ecology, 75, 1406–1419. Mitchell, T.B. (1962) Bees of the Eastern United States, Volume II. North Carolina Agricultural Experiment Station Technical Bulletin, 152, 1–557. Moure, J.S. (1954) Notas sôbre Epeolini sul-americanos (Hymenopt.-Apoidea). Dusenia, 5, 259–286. Moure, J.S. (1955) Notas sôbre Epeolini sulamericanos (Hymenopt.-Apoidea). Dusenia, 6, 115–138. Parker, F.D., Tepedino, V.J. & Bohart, G.E. (1981) Notes on the biology of a common sunflower bee, Melissodes (Eumelissodes) agilis Cresson. Journal of the New York Entomological Society, 89, 43–52. Provancher, L. (1895) Les dernières descriptions de l’Abbé Provancher. Ordre des Hyménoptères. Fam. XXI— Andrenidae. Naturaliste Canadien, 22, 189–191. Raw, A. (1984) The nesting biology of nine species of Jamaican bees (Hymenoptera). Revista brasileira de Entomolo- gia, 28, 497–506. Rightmyer, M.G. (2004a) Phylogeny and classification of the parasitic bee tribe Epeolini (Hymenoptera: Apidae, Noma- dinae). Scientific Papers, Natural History Museum, The University of Kansas, 33, 1–51. Rightmyer, M.G. (2004b) Redescription of two East Asian species of the tribe Epeolini (Hymenoptera: Apidae; Noma- dinae). Entomological Science, 7, 251–262. Robertson, C. (1897) North American bees—Descriptions and synonyms. Transactions of the Academy of Science of St. Louis, 7, 315–356. Robertson, C. (1898) New or little known North American bees. Transactions of the Academy of Science of St. Louis, 7, 43–54. Robertson, C. (1899) Flower visits of oligotropic bees. Botanical Gazette, 28, 215. Robertson, C. (1901) Some new or little known bees. Canadian Entomologist, 33, 229–231. Robertson, C. (1902) Some new or little known bees. Entomological News, 13, 79–81. Robertson, C. (1903) Synopsis of the Epeolinae. Canadian Entomologist, 35, 284–288. Robertson, C. (1928) Flowers and Insects. Lists of Visitors of Four Hundred and Fifty-Three Flowers. The Science Press Printing Company, Lancaster, Pennsylvania, 221 pp. Roig-Alsina, A. (1989) A revision of the bee genus Doeringiella (Hymenoptera, Anthophoridae, Nomadinae). Univer- sity of Kansas Science Bulletin, 53, 576–621. Rozen, J.G., Jr. (1964) The biology of Svastra obliqua obliqua (Say), with a taxonomic description of its larvae (Apoidea, Anthophoridae). American Museum Novitates, 2170, 1–13. Rozen, J.G., Jr. (1966) The larvae of the Anthophoridae (Hymenoptera, Apoidea) Part 2. The Nomadinae. American Museum Novitates, 2244, 1–38. Rozen, J.G., Jr. (1983) Nesting biology of the bee Svastra sabinensis (Hymenoptera, Anthophoridae). Journal of the New York Entomological Society, 91, 264–268. Rozen, J.G., Jr. (1984) Nesting biology of diphaglossine bees (Hymenoptera, Colletidae). American Museum Novitates, 2786, 1–33. Rozen, J.G., Jr. (1989a) Two new species and the redescription of another species of the cleptoparasitic genus Triepeolus with notes on their immature stages (Anthophoridae: Nomadinae). American Museum Novitates, 2956, 1–18. Rozen, J.G., Jr. (1989b) Morphology and systematic significance of first instars of the cleptoparasitic bee tribe Epeolini (Anthophoridae: Nomadinae). American Museum Novitates, 2957, 1–19. Rozen, J.G., Jr. (2003) Eggs, ovariole numbers, and modes of parasitism of cleptoparasitic bees, with emphasis on Neo- tropical species (Hymenoptera: Apoidea). American Museum Novitates, 3413, 1–36. Rust, R.W. (1984) Synonymy in California Channel Island Epeolini bees (Hymenoptera: Anthophoridae). Pan-Pacific

136 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER Entomologist, 60, 119–121. Say, T. (1823) A description of some new species of hymenopterous insects. Western Quarterly Reporter of Medical, Surgical, and Natural Sciences, 2, 71–82. Say, T. (1825) Appendix. Part I.—Natural History. 1. Zoology. In: Keating, W. H., Narrative of an Expedition to the Source of the St. Peter’s River, Lake Winnepeek, Lake of the Woods, &c. Performed in the Year 1823, by order of the Hon. J. C. Calhoun, Secretary of War, Under the Command of Stephen H. Long, U. S. T. E., Volume 2 . Cox and Bay- lis, London, England, pp. 1–104. Schrottky, C. (1910) Two new Nomadidae (Hymenoptera) from South America. Journal of the New York Entomologi- cal Society, 18, 208–210. Schrottky, C. (1913) La distributión geográfica de los himenópteros Argentinos. Annales de la Sociedad Cientifíca Argentina, 75, 115–286. Smith, F. (1854) Catalogue of Hymenopterous Insects in the Collection of the British Museum, Part 2. British Museum, London, England, pp. 199–465, pls. vii–xii. Smith, F. (1879) Descriptions of New Species of Hymenoptera in the Collection of the British Museum. British Museum, London, England, xxi + 240 pp. Spinola, M. (1851) Himenopteros. In: Gay, C., Historia Fisica y Politica de Chile Segun Documentos Adquiridos en esta Republica Durante Doce Años de Residencia en Ella y Publicada Bajo los Auspicios del Supremo Gobierno. Zoologia, Vol. 6. Casa del autor, Paris, France, pp. 153–569. Stevens, O.A. (1919) The panurgine bees of North Dakota and a new Epeolus. (Hym.). Canadian Entomologist, 51, 205–210. Torchio, P.F. (1986) Late embryogenesis and egg eclosion in Triepeolus and Anthophora with a prospectus of nomadine classification (Hymenoptera: Anthophoridae). Annals of the Entomological Society of America, 79, 588–596. Torchio, P.F. & Burdick, D.J. (1988) Comparative notes on the biology and development of Epeolus compactus Cres- son, a cleptoparasite of Colletes kincaidii Cockerell (Hymenoptera: Anthophoridae, Colletidae). Annals of the Ento- mological Society of America, 81, 626–636. Viereck, H.L. (1905) Synopsis of bees of Oregon, Washington, British Columbia and Vancouver.—IV. Canadian Ento- mologist, 37, 277–287. Warncke, K. (1982) Zur Systematik der Bienen—Die Unterfamilie Nomadinae (Hymenoptera, Apidae). Entomofauna. Zeitschrift für Entomologie, 3, 97–126. Wcislo, W.T. (1993) Communal nesting in a North American pearly-banded bee, Nomia tetrazonata, with notes on nesting behavior of Dieunomia heteropoda (Hymenoptera: Halictidae: Nomiinae). Annals of the Entomological Society of America, 86, 813–821. Wcislo, W.T., Minckley, R.L., Leschen, R.A.B., & Reyes, S. (1994) Rates of parasitism by natural enemies of a solitary bee, Dieunomia triangulifera (Hymenoptera, Coleoptera and Diptera) in relation to phenologies. Sociobiology, 23, 265–273. Webb, D.W. (1980) Primary insect types in the Illinois Natural History Survey Collection, exclusive of the Collembola and Thysanoptera. Illinois Natural History Survey Bulletin, 32, 51–191. Westrich, P. (1989) Die Wildbienen Baden-Württembergs: Allgemeiner Teil. Eugene Ulmer, Stuttgart, Germany, 431 pp. Wuellner, C.T. & Hixon, M.S. (1999) Behavior of a cleptoparasitic bee, Triepeolus distinctus (Hymenoptera: Nomadi- nae), before departing from the nest of its host, Dieunomia triangulifera (Hymenoptera: Halictidae). University of Kansas Natural History Museum Special Publication, 24, 143–150. Yasumatsu, K. (1933) Die Schmuckbienen (Epeolus) Japans (Hymenoptera, Melectidae). Transactions of the Kansai Entomological Society, 4, 1–6, 3 pls. Yasumatsu, K. (1938) Schmuckbienen (Epeolus) der mandschurischen Subregion (Hymenoptera, Apoidea). Transac- tions of the Sapporo Natural History Society, 15, 223–226. Zimsen, E. (1964) The Type Material of I. C. Fabricius. Munksgaard, Copenhagen, Denmark, 445 pp.

Taxonomic histories of species originally described in Triepeolus or subsequently placed in Triepeolus, now confirmed to belong to other genera.

EPEOLUS BANKSI (Cockerell) Triepeolus banksi Cockerell 1907c: 135–136. Epeolus banksi; Mitchell 1962: 442.

EPEOLUS FLAVOFASCIATUS Smith Epeolus flavofasciatus Smith 1879: 103. Triepeolus flavofasciatus; Cockerell 1904: 36. Triepeolus agaricifer Cockerell 1907b: 60. new synonymy Comments.—Brumley (1965: 30–33) redescribed this species under a manuscript name.

EPEOLUS MINIMUS (ROBERTSON) Triepeolus minimus Robertson 1902: 81. Argyroselenis minima; Robertson 1903: 284. Epeolus minimus; Brumley 1965: 56–60.

EPEOLUS SCUTELLARIS Say Epeolus scutellaris Say 1825: 85–86. Triepeolus scutellaris; Lovell and Cockerell 1905: 42.

DOERINGIELLA GAYI (Spinola) Epeolus gayi Spinola 1851: 188–189. Triepeolus fazi Cockerell 1925b: 493–494; Roig-Alsina 1989: 601 [synonymy]. Triepeolus gayi; Cockerell 1925b:494. Doeringiella gayi; Moure 1954: 267–268.

DOERINGIELLA HOLMBERGI (Schrottky) Epeolus variegatus Holmberg 1886: 279 [nec Linnaeus]. Epeolus holmbergi Schrottky 1913: 265 [replacement name]. Triepeolus pruinosus Cockerell 1917a: 478–479; Moure 1955: 126 [synonymy]. Doeringiella holmbergi; Moure 1954: 267.

138 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER FIGURES 3 and 4. Triepeolus antiguensis, female, from San José, Costa Rica. 3. Dorsal habitus. 4. Pseudopygidial area. Figs. 5 and 6. Triepeolus antiochensis, female, from California. 5. Dorsal habitus. 6. Pseudopygidial area. Figs. 7 and 8. Triepeolus argentimus, female, from California. 7. Dorsal habitus. 8. Pseudopygidial area. Figs. 9 and 10. Triepeolus argus, females, from North Dakota (Fig. 9) and Utah (Fig. 10). 9. Dorsal habitus. 10. Pseudopygidial area.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 139 FIGURES. 11 and 12. Triepeolus argyreus, female, from Washington. 11. Dorsal habitus. 12. Pseudopygidial area. Figs. 13 and 14. Triepeolus atripes, females, from Illinois (Fig. 13) and Missouri (Fig. 14). 13. Dorsal habitus. 14. Pseudopygidial area. Figs. 15–17. Triepeolus aztecus, females, from Alajuela (Figs. 15 and 16) and San José (Fig. 17), Costa Rica. 15. Dorsal habitus. 16. Close-up of first and second terga. 17. Pseudopygidial area. Fig. 18. Triepeolus balteatus, female, from Colorado. 18. Dorsal habitus.

140 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER FIGURES. 19 and 20. Triepeolus balteatus, female, from Colorado. 19. Pseudopygidial area. 20. Clasper-like intercoxal area. Figs. 21 and 22. Triepeolus bimorulus, female, from New Mexico. 21. Dorsal habitus. 22. Pseudopygidial area. Figs. 23 and 24. Triepeolus brittaini, female, from Nova Scotia, Canada. 23. Dorsal habitus. 24. Pseudopygidial area. Figs. 25 and 26. Triepeolus californicus, female, from California. 25. Dorsal habitus. 26. Pseudopygidial area.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 141 FIGURES. 27 and 28. Triepeolus cameroni, female, from Veracruz, México. 27. Dorsal habitus. 28. Pseudopygidial area. Figs. 29 and 30. Triepeolus charlesi, females, from Texas. 29. Dorsal habitus. 30. Pseudopygidial area. Figs. 31 and 32. Triepeolus circumculus, females, from New Mexico (Fig. 31) and Arizona (Fig. 32). 31. Dorsal habitus. 32. Pseudopygidial area. Figs. 33 and 34. Triepeolus claytoni, females, from Durango (Fig. 33) and Chihuahua (Fig. 34), México. 33. Dorsal habitus. 34. Pseudopygidial area.

142 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER FIGURES. 35–38. Triepeolus concavus, female, from California. 35. Posterior view of pseudopygidial area. 36. Lateral view of pseudopygidial area. 37. Mesosoma. 38. First and second terga. Figs. 39 and 40. Triepeolus cressonii, females, from Kansas. 39. Dorsal habitus. 40. Pseudopygidial area. Figs. 41 and 42. Triepeolus cruciformis, female, from Pue- bla, México. 41. Dorsal habitus. 42. Pseudopygidial area.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 143 FIGURES. 43 and 44. Triepeolus cuabitensis, female, from Santiago de Cuba, Cuba. 43. Dorsal habitus. 44. Pseudopygidial area. Figs. 45 and 46. Triepeolus dacotensis, females, from Alberta, Canada. 45. Dorsal habitus. 46. Pseudopygidial area. Figs. 47–50. Triepeolus denverensis, females, from Arizona (Figs. 47 and 50) and Utah (Figs. 48 and 49). 47–49. Dorsal habitus photographs, showing range in amount of pale setae. 50. Pseudopygidial area.

144 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER FIGURES. 51 and 52. Triepeolus diffusus, females, from Colorado (Fig. 51) and Utah (Fig. 52). 51. Dorsal habitus. 52. Pseudopygidial area. Figs. 53 and 54. Triepeolus dilutus, female, from Texas. 53. Dorsal habitus. 54. Pseudopygidial area. Figs. 55–57. Triepeolus distinctus, females, from Illinois (Fig. 55) and Texas (Figs. 56 and 57). 55. Dorsal habitus. 56. Pseudopygidial area. 57. Face. Fig. 58. Triepeolus donatus, female, from Ohio. Face, showing relative positions of apical margin of clypeus and lower tangent of compound eye.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 145 FIGURES. 59 and 60. Triepeolus diversipes, females, from Utah (Fig. 59) and New Mexico (Fig. 60). 59. Dorsal habitus. 60. Pseudopygidial area. Figs. 61 and 62. Triepeolus donatus, male from Kansas (Fig. 61) and female from Ohio (Fig. 62). 61. Dorsal habitus. 62. Pseudopygidial area. Figs. 63–65. Triepeolus edwardi, females, from California. 63. Dorsal habitus. 64. Dorsal habitus. 65. Pseudopygidial area. Fig. 66. Triepeolus eldoradensis, female, from Utah. Dor- sal habitus.

146 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER FIGURE 67. Triepeolus eldoradensis, female, from Utah. Pseudopygidial area. Figs. 68 and 69. Triepeolus engeli, female, from Texas. 68. Dorsal habitus. 69. Pseudopygidial area. Figs. 70 and 71. Triepeolus epeolurus, female, from Jalisco, México. 70. Dorsal habitus. 71. Pseudopygidial area. Figs. 72 and 73. Triepeolus exilicurvus, female, from Coahuila, México. 72. Dorsal habitus. 73. Pseudopygidial area. Fig. 74. Triepeolus flavigradus, female, from Puebla, México. Dorsal habitus.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 147 FIGURE 75. Triepeolus flavigradus, female, from Puebla, México. Pseudopygidial area. Figs. 76 and 77. Triepeolus fraserae, females, from Utah (Fig. 76) and Arizona (Fig. 77). 76. Dorsal habitus. 77. Pseudopygidial area. Figs. 78 and 79. Triepeolus fulgidus, females, from New Mexico. 78. Dorsal habitus. 79. Pseudopygidial area. Figs. 80 and 81. Trie- peolus georgicus, female, from Florida. 80. Dorsal habitus. 81. Pseudopygidial area. Fig. 82. Triepeolus grandis, female, from Arizona. Dorsal habitus.

148 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER FIGURE 83. Triepeolus grandis, female, from New Mexico. Pseudopygidial area. Figs. 84 and 85. Triepeolus griswoldi, females, from Utah. 84. Dorsal habitus. 85. Pseudopygidial area. Figs. 86 and 87. Triepeolus helianthi, females, from California (Fig. 86) and Colorado (Fig. 87). 86. Dorsal habitus. 87. Pseudopygidial area. Figs. 88–90. Triepeolus heterurus, females, from California (Figs. 88 and 89) and Baja California, México (Fig. 90). 88. Dorsal habitus. 89. Pseudopygidial area. 90. Dorsal habitus.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 149 FIGURE 91. Triepeolus heterurus, female, Baja California, México. Pseudopygidial area. Figs. 92 and 93. Triepeolus interruptus, female, from Michoacán, México. 92. Dorsal habitus. 93. Pseudopygidial area. Figs. 94 and 95. Triepeolus intrepidus, female, from Arizona. 94. Dorsal habitus. 95. Pseudopygidial area. Figs. 96 and 97. Triepeolus isohedrus, female, from New Mexico. 96. Dorsal habitus. 97. Pseudopygidial area. Fig. 98. Triepeolus jennieae, female, from Hidalgo, México. Dorsal habitus.

150 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER FIGURE 99. Triepeolus jennieae, female, from Hidalgo, México. Pseudopygidial area. Figs. 100 and 101. Triepeolus joliae, females, from Nevada (Fig. 100) and Arizona (Fig. 101). 100. Dorsal habitus. 101. Pseudopygidial area. Figs. 102 and 103. Triepeolus lateralis, female, from Puebla, México. 102. Dorsal habitus. 103. Pseudopygidial area. Figs. 104 and 105. Triepeolus laticaudus, female, from Texas. 104. Dorsal habitus. 105. Pseudopygidial area. Fig. 106. Trie- peolus laticeps, female, from México, México. Dorsal habitus.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 151 FIGURE 107. Triepeolus laticeps, female, from México, México. Pseudopygidial area. Figs. 108 and 109. Triepeolus loomisorum, females, from Arizona (Fig. 108) and New Mexico (Fig. 109). 108. Dorsal habitus. 109. Pseudopygidial area. Figs. 110–113. Triepeolus lunatus, female, from Kansas. 110. Dorsal habitus. 111. Dorsal habitus. 112. Face. 113. Pseudopygidial area. Fig. 114. Triepeolus margaretae, female, from Texas. Dorsal habitus.

152 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER FIGURE 115. Triepeolus margaretae, female, from Texas. Pseudopygidial area. Figs. 116 and 117. Triepeolus martini, females, from Colorado. 116. Dorsal habitus. 117. Pseudopygidial area. Figs. 118 and 119. Triepeolus mauropygus, females, from Arizona (Fig. 118) and Sinaloa, México (Fig. 119). 118. Dorsal habitus. 119. Pseudopygidial area. Figs. 120 and 121. Triepeolus medusa, female, from Jalisco, México. 120. Dorsal habitus. 121. Pseudopygidial area. Fig. 122. Triepeolus melanarius, female, from California. Dorsal habitus.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 153 FIGURE 123. Triepeolus melanarius, female, from California. Pseudopygidial area. Figs. 124–126. Triepeolus micheneri, female, from Jalisco (Figs. 124 and 126) and Morelos (Fig. 125), México. 124. Dorsal habitus. 125. Pseudopygidial area. 126. Mesepisternum, lateral view. Figs. 127 and 128. Triepeolus michiganensis, females, from New Hampshire (Fig. 127) and New York (Fig. 128). 127. Dorsal habitus. 128. Pseudopygidial area. Figs. 129 and 130. Triepeolus micropygius, female from Arizona. 129. Dorsal habitus. 130. Pseudopygidial area.

154 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER FIGURE 131. Triepeolus mitchelli, holotype male, from North Carolina. Figs. 132 and 133. Triepeolus mojavensis, females, from California. 132. Dorsal habitus. 133. Pseudopygidial area. Figs. 134 and 135. Triepeolus monardae, female, from North Carolina. 134. Dorsal habitus. 135. Pseudopygidial area. Figs. 136 and 137. Triepeolus nayaritensis, female, from Nayarit, México. 136. Dorsal habitus. 137. Pseudopygidial area. Fig. 138. Triepeolus nevadensis, female, from Durango, México. Dorsal habitus.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 155 FIGURES 139 and 140. Triepeolus nevadensis, females, from Durango, México. 139. Mesosoma. 140. Pseudopygidial area. Figs. 141–144. Triepeolus nigrihirtus, female, from Texas. 141. Dorsal habitus. 142. Pseudopygidial area. 143. Face. 144. Head and mesosoma, oblique view showing preoccipital carina on posterior margin of head behind vertex. Figs. 145 and 146. Triepeolus nisibonensis, male, from La Altagracia, Dominican Republic. 145. Dorsal habitus. 146. Posterior margin of vertex, showing median emargination.

156 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER FIGURES 147 and 148. Triepeolus norae, females, from Arizona (Fig. 147) and New Mexico (Fig. 148). 147. Dorsal habitus. 148. Pseudopygidial area. Figs. 149 and 150. Triepeolus occidentalis, females, from North Dakota (Fig. 149) and Colorado (Fig. 150). 149. Dorsal habitus. 150. Pseudopygidial area. Figs. 151–154. Triepeolus paenepectoralis, females, from Washington (Fig. 151), Oregon (Fig. 152), and California (Figs. 153 and 154). 151. Dorsal habitus. 152. Dorsal habitus. 153. Dorsal habitus. 154. Pseudopygidial area.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 157 FIGURES 155 and 156. Triepeolus parkeri, female, from Guanacaste, Costa Rica. 155. Dorsal habitus. 156. Pseudopy- gidial area. Figs. 157 and 158. Triepeolus partitus, females, from Arizona. 157. Dorsal habitus. 158. Pseudopygidial area. Figs. 159 and 160. Triepeolus parvidiversipes, females, from Arizona. 159. Dorsal habitus. 160. Pseudopygidial area. Figs. 161 and 162. Triepeolus parvus, females, from New Mexico. 161. Dorsal habitus. 162. Pseudopygidial area.

158 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER FIGURES 163 and 164. Triepeolus pectoralis, female, from North Carolina. 163. Dorsal habitus. 164. Pseudopygidial area. Figs. 165 and 166. Triepeolus pectoralis, female mesepisternum, from North Carolina. 165. Lateral view. 166. Dorsal view showing long, erect, simple setae. Figs. 167–169. Triepeolus penicilliferus, females, from Arizona (Figs. 167 and 169) and California (Fig. 168). 167. Dorsal habitus. 168. Dorsal habitus. 169. Pseudopygidial area. Fig. 170. Triepeolus perpictus, female, from Nevada. Dorsal habitus.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 159 FIGURE 171. Triepeolus perpictus, females, from Utah. Pseudopygidial area. Figs. 172 and 173. Triepeolus phaeopygus, females, from Arizona. 172. Dorsal habitus. 173. Pseudopygidial area. Figs. 174 and 175. Triepeolus punctoclypeus, female, from California. 174. Dorsal habitus. 175. Pseudopygidial area. Figs. 176 and 177. Triepeolus quadratus, female, from California. 176. Dorsal habitus. 177. Pseudopygidial area. Fig. 178. Triepeolus quadrifasciatus atlanticus, female, from Alabama. Dorsal habitus.

160 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER FIGURES 179–181. Triepeolus quadrifasciatus atlanticus, female, from North Carolina. 179. Pseudopygidial area. 180. Face. 181. Mesoscutum. Figs. 182–184. Triepeolus quadrifasciatus quadrifasciatus, female, from Texas. 182. Face. 183. Dorsal habitus. 184. Pseudopygidial area. Figs. 185 and 186. Triepeolus remigatus, female, from Arizona. 185. Dorsal habitus. 186. Mesosoma.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 161 FIGURE 187. Triepeolus remigatus, female, from Arizona. Pseudopygidial area. Figs. 188–190. Triepeolus robustus, female, from New Mexico. 188. Dorsal habitus. 189. Pseudopygidial area. 190. Face, oblique view showing long, erect setae surrounding clypeus. Figs. 191 and 192. Triepeolus rohweri, female, from Colorado. 191. Dorsal habitus. 192. Pseudopygidial area. Figs. 193 and 194. Triepeolus rufithorax, females, from Florida. 193. Dorsal habitus. 194. Dorsal habitus.

162 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER FIGURE 195. Triepeolus rufithorax, female, from Florida. Pseudopygidial area. Figs. 196 and 197. Triepeolus rufoclypeus, females, from Tamaulipas, México (Fig. 196) and Texas (Fig. 197). 196. Dorsal habitus. 197. Pseudopygidial area. Figs. 198–200. Triepeolus rugosus, females, from Florida (Figs. 198 and 199) and New Jersey (Fig. 200). 198. Dorsal habitus. 199. Mesepisternum, lateral view. 200. Pseudopygidial area. Figs. 201 and 202. Triepeolus scelestus, females, from Kansas. 201. Dorsal habitus. 202. Pseudopygidial area.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 163 FIGURES2 203–205. Triepeolus simulatus, female, from California. 203. Dorsal habitus. 204. Pseudopygidial area. 205. Mesosoma, showing pattern of setae on scutellum. Figs. 206 and 207. Triepeolus subalpinus, female, from Wyo- ming. 206. Dorsal habitus. 207. Pseudopygidial area. Figs. 208 and 209. Triepeolus sublunatus, females, from Arizona. 208. Dorsal habitus. 209. Pseudopygidial area. Fig. 210. Triepeolus subnitens, female, from California. Dorsal habitus.

164 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER FIGURE 211. Triepeolus subnitens, female, from California. Pseudopygidial area. Figs. 212 and 213. Triepeolus tanneri, female, from Kansas. 212. Dorsal habitus. 213. Pseudopygidial area. Figs. 214–216. Triepeolus tepanecus, females, from Jalisco (Figs. 214 and 216) and Colima (Fig. 215), México. 214. Dorsal habitus. 215. Pseudopygidial area. 216. Mesepisternum, lateral view. Figs. 217 and 218. Triepeolus texanus, females, from Colorado (Fig. 217) and Washington (Fig. 218). 217. Dorsal habitus. 218. Pseudopygidial area.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 165 FIGURES 219 and 220. Triepeolus totonacus, female, from Michoacán, México (Fig. 219) and Chiapas, México (Fig. 220). 219. Dorsal habitus. 220. Pseudopygidial area. Figs. 221 and 222. Triepeolus townsendi, females, from Arizona (Fig. 221) and New Mexico (Fig. 222). 221. Dorsal habitus. 222. Pseudopygidial area. Figs. 223 and 224. Triepeolus tristis, female, from Burgenland, Austria. 223. Dorsal habitus. 224. Pseudopygidial area. Figs. 225 and 226. Triepeolus ventralis, female, from Chūbu, Japan. 225. Dorsal habitus. 226. Pseudopygidial area.

166 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER FIGURES 227 and 228. Triepeolus vernus, female, from Texas. Fig. 227. Dorsal habitus. 228. Pseudopygidial area. Figs. 229–231. Triepeolus vicinus, females, from Pinar del Rio (Figs. 229 and 230) and Havana (Fig. 231), Cuba. 229. Mesepisternum, lateral view. 230. Dorsal habitus. 231. Pseudopygidial area. Figs. 232 and 233. Triepeolus victori, from El Seibo, Dominican Republic. 232. Dorsal habitus, male. 233. Pseudopygidial area. Fig. 234. Triepeolus warriti, female, from New Mexico. Dorsal habitus.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 167 FIGURES 235 and 236. Triepeolus warriti, female, from New Mexico. 235. Metasoma, dorsal view. 236. Pseudopy- gidial area. Figs. 237 and 238. Triepeolus wilsoni, female, from Pinar del Rio, Cuba. 237. Dorsal habitus. 238. Pseudopygidial area. Figs. 239 and 240. Triepeolus zacatecus, female, from Colima, México. 239. Dorsal habitus. 240. Pseudopygidial area. Figs. 241 and 242. Triepeolus grandis, male metasoma, showing apical fringes of setae on S3–S5. 241. Ventral view. 242. Lateral view.

168 · Zootaxa 1710 © 2008 Magnolia Press RIGHTMYER FIGURE 243. Triepeolus grandis, male, dorsal habitus. Figs. 244–247. Clypeus illustrations. 244. Triepeolus distinctus (from holotype of Triepeolus mesillae). 245. Triepeolus rufithorax. 246. Triepeolus subnitens. 247. Triepeolus martini.

REVIEW OF TRIEPEOLUS Zootaxa 1710 © 2008 Magnolia Press · 169 FIGURES 248–261. Female metasomata, dorsal views. 248. Triepeolus cressonii. 249. Triepeolus dacotensis. 250 and 251. Triepeolus distinctus. 252. Triepeolus donatus. 253. Triepeolus edwardi. 254. Triepeolus eldoradensis. 255. Trie- peolus grandis. 256. Triepeolus lunatus. 257. Triepeolus martini. 258. Triepeolus mojavensis. 259. Triepeolus occidentalis. 260. Triepeolus remigatus. 261. Triepeolus tristis. Figs. 262 and 263. Female sterna 5, ventral views (A), and terga 5, dorsal views (B). 262. Triepeolus penicilliferus. 263. Triepeolus concavus.