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Research Article
A revision of the cleptoparasitic bee genus Epeolus Latreille for Nearctic species, north of Mexico (Hymenoptera, Apidae)
expand article infoThomas M. Onuferko
‡ York University, Toronto, Canada
Open Access

Abstract

Herein, the cleptoparasitic (cuckoo) bee genus Epeolus (Hymenoptera: Apidae) is revised for species occurring in North America, north of Mexico, and an updated checklist of all species known to occur in Canada and the United States of America is provided with comprehensive descriptions, diagnoses, and a single dichotomous key (using the same couplets for both sexes) to aid in their identification. To increase their recognition among North American naturalists, English common names are also proposed for all North American Epeolus. A total of 43 species is confirmed as present in the region, 15 of which are newly recognized. The following new species are proposed based on unique morphological (and in most cases also molecular) attributes: E. andriyi sp. n., E. attenboroughi sp. n., E. axillaris sp. n., E. basili sp. n., E. brumleyi sp. n., E. chamaesarachae sp. n., E. deyrupi sp. n., E. diadematus sp. n., E. ferrarii sp. n., E. gibbsi sp. n., E. inornatus sp. n., E. nebulosus sp. n., E. packeri sp. n., E. splendidus sp. n., and E. tessieris sp. n. Of the 15, six (E. axillaris, E. brumleyi, E. chamaesarachae, E. diadematus, E. splendidus, and E. tessieris) were identified as new species under different names (nomina nuda) in an M.Sc. thesis by Richard L. Brumley in 1965, but until now they have not been formally described. Detailed morphological comparisons with some evidence from DNA barcoding support the following synonymies, one of which C was first proposed by Brumley (1965): a) E. melectimimus Cockerell and Sandhouse, syn. n., under E. asperatus Cockerell; b) E. crucis Cockerell, syn. n., under E. compactus Cresson; c) E. mesillae palmarum Linsley, syn. n., under E. mesillae (Cockerell); and d) E. weemsi Mitchell, syn. n., and e) E. vernalis Mitchell, syn. n., under E. ilicis Mitchell. Only one member of the almost entirely Neotropical “Trophocleptria group” (Epeolus bifasciatus Cresson) is confirmed as occurring north of Mexico, and is widespread East of the Rocky Mountains. Known floral associations are indicated for each species, as are suspected or known host species of Colletes Latreille. Evidence is presented that suggests further investigation into the possible synonymy of Colletes wickhami Timberlake under C. scopiventer Swenk is warranted.

Keywords

cleptoparasitic bee, DNA barcoding, Epeolus, morphology, taxonomic revision

Introduction

Epeolus Latreille (Hymenoptera: Apidae, subfamily Nomadinae) is one of the most widespread genera of cleptoparasitic bees (commonly referred to as cuckoo bees), occurring on all continents except Antarctica and Australia. The genus is also absent from Madagascar, Oceania, and parts of Southeast Asia, regions in which their host genus Colletes Latreille (Hymenoptera: Colletidae: Colletinae) is not present (Michener 2007). Other genera in the tribe Epeolini are largely restricted to the Americas, mostly to the Neotropical region. The similarly diverse bee genus Triepeolus Robertson has only two representatives in the Palearctic region, whereas Epeolus is represented across Africa, Asia, and Europe by about 48 species (Ascher and Pickering 2017). However, the genus is most diverse in North America, with 32 valid species confirmed as occurring north of Mexico before the date of this publication.

For North American species, the taxonomy of Epeolus has been in need of revision for some time. While Mitchell’s (1962) treatment of the Eastern United States fauna was fairly comprehensive, the Western species have been in much need of attention. In his M.Sc. thesis, Richard L. Brumley (1965) recognized several new species from the Western United States, but his names were never published and are therefore not considered valid. Recently, Onuferko (2017) identified 14 redundant names (most are of Western “species”), which were synonymized under the names of four valid species, but this treatment was limited to the Canadian fauna. The purpose of the present study is to resolve the taxonomy of Epeolus occurring in Canada and the USA by naming and describing new species and identifying which accepted names are valid and which are not, thereby standardizing name use, as well as to provide a user-friendly dichotomous identification key. To help amateur and professional entomologists become more familiar with these bees, English common names are proposed for all North American species of Epeolus. An additional objective is to present ecological information in terms of floral and Colletes hosts and phenology wherever possible, as well as comprehensive occurrence records to aid those interested in locating and identifying representatives of the species treated herein for further research.

Materials and methods

To revise Epeolus an integrative biosystematics approach was followed, using morphological and molecular evidence to distinguish intraspecific from interspecific variation (as in Gibbs 2009, 2010, 2011, Pauly et al. 2014, Rocha-Filho and Packer 2015, Ferrari 2017, Onuferko 2017). Morphological evidence was prioritized over molecular evidence when the two were not in agreement, as in Gibbs (2009). Sequence data from a 658 bp segment of the mitochondrial cytochrome c oxidase subunit I (COI) gene (DNA barcode, Hebert et al. 2003a, b) were obtained from specimens of nearly all (42 out of 43) species, and 37 have sequences that are barcode compliant (i.e., have met the criteria to be assigned automated barcode index numbers (BINs) given to unique barcode clusters, Ratnasingham and Hebert 2007, 2013). One or two legs were removed from each specimen to be “barcoded”, and sent to the Canadian Centre for DNA Barcoding in Guelph, Ontario, Canada for DNA extraction and gene amplification and sequencing. A neighbor-joining (NJ) tree, based on Kimura’s two-parameter distance model (Kimura 1980), was used to compare short, non-compliant and barcode-compliant sequences for the purpose of validating species designations of sequenced specimens and checking for contamination errors. Partial and BIN-compliant sequences are published in the “Epeolus of North America project” on the Barcode of Life Data Systems website (http://www.barcodinglife.org/) and have been deposited in the GenBank database (see Suppl. material 1 for accession numbers).

Terminology used herein is consistent with that used in the recent treatment of Canadian Epeolus (Onuferko 2017), which generally followed Michener (2007), except the terms frontal area and vertexal area are used instead of frons and vertex, respectively. Acronyms used herein (in bold) are as follows. Puncture density is described in terms of interspaces (i) relative to the diameters (d) of punctures. Median ocellar diameter (MOD) is a comparative unit of measurement for smaller structures. F followed by a number represents one of 10 (for female) or 11 (for male) flagellomeres of the antenna. T followed by a number represents one of six (for female) or seven (for male) exposed metasomal terga. S followed by a number represents one of six (for female) or eight (for male) metasomal sterna. Several terms used in Onuferko (2017), some of which were taken from Rightmyer (2008), are defined here again for clarity, and are indicated in bold. Length refers to measurements made along the longitudinal axis of the bee, except in reference to the longitudinal extent of the transverse metasomal fasciae, for which the term breadth is used, and width refers to measurements made along the lateral axis. The length and width of an anatomical feature refer to its longest and widest margins, respectively, and were recorded at the highest magnification that allowed measurement in ocular micrometer units. The scape was measured without the radicle. In Epeolus, the frontal line extends into the supraclypeal area as a pronounced carina on a convex surface, referred to herein as the frontal keel. Paramedian bands are the paired lines of off-white or yellow tomentum located anteriorly on the mesoscutum of most Epeolus species (Fig. 1). The term bigibbous is an adjective used in reference to the biconvexities present on the mesoscutellum of Epeolus species. The basal and apical fasciae of T1 are often connected by a longitudinal band of pale tomentum of varying width. Discal patch refers to the discal region of T1 that is typically covered in dark tomentum and is bordered by bands of pale tomentum. This area is not always clearly delineated because the surrounding bands of pale tomentum may be reduced or missing entirely.

Figure 1. 

Female E. chamaesarachae sp. n. illustrating mesosomal and metasomal bands of tomentum commonly present in North American Epeolini.

The species of Epeolus are, with the usual exceptions (differences in the number of antennal flagellomeres, number of exposed metasomal terga, length of the S4 and S5 subapical hairs [usually longer in males], and terminalia) and a few atypical ones, sexually monomorphic. For this reason, separate keys for females and males are not presented, and the few sex-specific features used to distinguish species are indicated as such in the couplets. The key to Nearctic Epeolus is heavily based on the structure of the axilla and the bands of pale tomentum forming the basal and apical fasciae on the metasomal terga. To limit the number of steps required to identify all species, efforts were made to make the key as close to fan shaped (evenly bifurcated) as possible, following the recommendations of Walter and Winterton (2007; see also Packer et al. 2016). When possible, couplets were based on more than a single feature (ideally one per tagma) should one be obscured or lost in the specimen being identified. However, avoiding monothetic couplets was not always possible. In such cases couplets were usually based on mesosomal features that should be visible even in damaged pinned specimens. In couplets that list multiple features, the most important (i.e., reliable) one for achieving a diagnosis is given first whereas features that do not always result in a positive identification (e.g., integument black vs integument black or ferruginous will resolve species with ferruginous but not black integument) are included but given at the end and always preceded by at least one feature that is fully contrasted between both halves of the couplet. The features referenced in the key were imaged. Quite often a single image or image plate was used to illustrate more than one feature, so a number of figures were cited two or more times within the key and elsewhere in the present monograph. As a result, it was not possible to put most illustrations near the couplets without duplicating them, and for practical reasons multiple versions of the same figures are not included herein. Many couplets rely on precise comparative measurements, and the key is meant to be used with the aid of an eyepiece graticule. None of the couplets require specimens to be dissected. Although the male S7, S8, and genital capsules of nearly all species were examined (except those represented by very few male specimens), the variation among them is minimal (illustrated in part in plates 2 and 3 in Onuferko 2017), and the terminalia have not proven useful in separating similar-looking species. Consequently, they have not been illustrated or imaged. The illustrations presented to aid in the identification of Epeolus species are my own. Images were taken with a digital camera (Canon EOS 40D SLR) using the Visionary Digital macro-imaging BK PLUS Lab System, focus stacked in Helicon Focus, and edited in Adobe Photoshop and PaintShop Pro.

Species descriptions follow the format of Onuferko (2017). A full description of the primary type specimen of each species is provided, except for the species occurring in Canada that were recently redescribed in Onuferko (2017). The physical name-bearing type specimens of all described North American Epeolus were seen and thoroughly examined, including those whose names are no longer considered valid, except in the case of E. mercatus Fabricius, for which the original type material cannot be traced and description is so insufficiently detailed that it is unclear if the species is an Epeolus or Triepeolus (Rightmyer 2008). Since most Epeolus species to date were described from female specimens, new species described herein are generally represented by a female holotype, male allotype, and paratypes. Given that Epeolus is a genus of largely sexually monomorphic species, descriptions of the sex opposite that of the name-bearing type list only key differences to avoid unnecessary duplication of text. In many, but not all, cases it is the female that is fully described. I have opted to propose new names for the species Brumley (1965) discovered rather than validate the ones he used. This will ensure that it is clear who made designations of type specimens (i.e., specimens used as types by Brumley (1965) and me have both our type labels, those unavailable to me but designated as types by Brumley (1965) have only his labels, and those seen exclusively by me and given type status have only my labels). This will also eliminate any possible confusion that could arise if Brumley’s (1965) names are published and registered in ZooBank long after their first appearance in his thesis.

The proposed common name for each species reflects its scientific name, which in most cases was easy to translate into English. Since there are many genera of cuckoo bees, epeolus is used herein as the common name for the genus instead of cuckoo bee or more specific but cumbersome names like Colletes cuckoo bee or polyester bee cuckoo bee.

Among the material examined were representatives of Epeolus from all Canadian provinces and territories except Newfoundland and Labrador and Nunavut, and all but six (Connecticut, Delaware, Kentucky, Rhode Island, Tennessee, and West Virginia) of the 49 states in the continental U.S. where the genus is expected to occur. Also examined were Epeolus records from 17 states in Mexico, and their data are included for species confirmed as occurring north of the Mexico–United States border. All examined records are presented in Suppl. material 1. Specimens were made available for study by curators and collections managers (in parentheses) from the following institutions:

ABS Archbold Biological Station, Venus, FL (M. Deyrup);

AMNH American Museum of Natural History, New York, NY (J.G. Rozen, Jr. and C. Smith);

ANSP Academy of Natural Sciences of Drexel University, Philadelphia, PA (J. Weintraub);

AUMNH Auburn University Museum of Natural History, Auburn, AL (C.H. Ray);

BBSL Utah State University USDA Bee Biology and Systematics Laboratory, Logan, UT (T.L. Griswold);

BIML Patuxent Wildlife Research Center USGS Native Bee Inventory and Monitoring Lab, Laurel, MD (S. Droege);

CAS California Academy of Sciences, San Francisco, CA (B. Fisher and R. Zuparko);

CNC Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, ON (S. Cardinal);

CTMI Central Texas Melittological Institute, Austin, TX (J.L. Neff);

CUIC Cornell University Insect Collection, Ithaca, NY (J. Dombroskie);

CUM University of Colorado Museum of Natural History, Boulder, CO (V. Scott);

DEBU University of Guelph Insect Collection, Guelph, ON (S.A. Marshall);

EMEC University of California Essig Museum of Entomology, Berkeley, CA (P. Oboyski);

FMNH Field Museum of Natural History, Chicago, IL (C. Maier);

FSCA Florida State Collection of Arthropods, Gainesville, FL (K. Schnepp and P.E. Skelley);

INHS Illinois Natural History Survey, Champaign, IL (C. Grinter);

JBWM University of Manitoba J.B. Wallis / R.E. Roughley Museum of Entomology, Winnipeg, MB (J. Gibbs);

KUNHM University of Kansas Biodiversity Institute and Natural History Museum, Lawrence, KS (M.S. Engel and J. Thomas);

LACM Natural History Museum of Los Angeles County, Los Angeles, CA (B.V. Brown and G.A. Kung);

MCZ Harvard University Museum of Comparative Zoology, Cambridge, MA (P.D. Perkins);

NCSU North Carolina State University Insect Museum, Raleigh, NC (R. Blinn);

NHMUK Natural History Museum, London, United Kingdom (D. Notton);

PCYU Packer Collection at York University, Toronto, ON (L. Packer);

ROM Royal Ontario Museum, Toronto, ON (A. Guidotti);

RSKM Royal Saskatchewan Museum, Regina, SK (C. Sheffield);

UCBME University of California Bohart Museum ​of Entomology, Davis, CA (S. Heydon and T.J. Zavortink);

UCR University of California Entomology Research Museum, Riverside, CA (D. Yanega); and

USNM U.S. National Entomological Collection, National Museum of Natural History, Washington, D.C. (S.G. Brady and B. Harris).

In lists of examined specimens, semi-colons separate records from different localities. Otherwise, commas are used between records from the same locality that are associated with a different collection date, collector(s), and/or repository. In such cases, the locality is not repeated and a comma appears after the specimen repository and before the collection date of the next record. If only the collection day and month were given, then “????” was used for the missing year. If the collection year was given to two digits but the century or millennium could not be inferred (e.g., from knowing who the collector was and the period in which he/she would have conducted field work), the two-digit year is still indicated but with “??” in front. All GPS coordinates indicated herein are taken directly from specimen labels. For approximate coordinates obtained post hoc for specimens with imprecise locality records used to construct range maps, see Suppl. material 1. For species reported from Canada, only total numbers of females and males from each province or state are shown for examined non-type specimens if the same records have already been published (Onuferko 2017).

Range maps were constructed as in Onuferko (2017) in RStudio (version 1.0.44) using the packages maptools (Bivand and Lewin-Koh 2014), raster (Hijmans 2014), rgdal (Bivand et al. 2014), and rgeos (Bivand and Rundel 2014) installed in R (version 3.3.2) (R Core Team 2016). The shapefiles used to plot projected maps of Canada, Mexico, and the USA were obtained from Statistics Canada (2015), DIVA-GIS (http://www.diva-gis.org/gdata), and the US Census Bureau (2015), respectively.

Floral associations are given for each species based on photo records, observations, and specimen labels. Records published in Onuferko (2017) are not repeated here, but they are included in Suppl. material 1. All floral records were checked against The Plant List (http://www.theplantlist.org/) to ensure that the scientific nomenclature is up to date.

Taxonomy

Epeolus Latreille, 1802

Epeolus Latreille, 1802: 427. Type species: Apis variegata Linnaeus, 1758, by monotypy.

Trophocleptria Holmberg, 1886: 233, 275. Type species: Trophocleptria variolosa Holmberg, 1886, by monotypy.

Epeolus (Diepeolus) Gribodo, 1894: 80. Type species: Epeolus giannellii Gribodo, 1894, by monotypy.

Epeolus (Monoepeolus) Gribodo, 1894: 80. Type species: Apis variegata Linnaeus, by monotypy.

Pyrrhomelecta Ashmead, 1899: 66. Type species: Epeolus glabratus Cresson, 1878, by original designation and monotypy.

Argyroselenis Robertson, 1903: 284. Type species: Triepeolus minimus Robertson, 1902, by original designation and monotypy.

Oxybiastes Mavromoustakis, 1954: 260. Type species: Oxybiastes bischoffi Mavromoustakis, 1954, by original designation and monotypy.

Remarks

In his original description, Latreille (1802) did not explain the etymology of Epeolus, but it seems likely that the name is a diminutive of Epeus/Epeius, the soldier in Greek mythology to whom building the Trojan Horse is attributed, and that it was inspired by the sinister nature of these cleptoparasitic bees. This was the first genus of Epeolini described, and ‘epeolus’ has since become the root in the names of many other nomadine and non-nomadine genera and tribes (e.g., Epeoloides Giraud (Osirini), Parepeolus Ducke (Osirini), Protepeolini, Pseudepeolus Holmberg (Epeolini), etc.).

Several species of Epeolus were previously described as belonging to different genera, in particular Triepeolus. On account of Rightmyer’s (2008) revision of Triepeolus, the generic placement of species that were once erroneously switched has been corrected. A few North American species were (initially or at some point in the past) described as belonging to genera that are no longer considered valid, including Argyroselenis Robertson, Phileremus (the name is a synonym of Ammobates Latreille subgenus Ammobates Latreille s. str. in Michener 2007), and Pyrrhomelecta Ashmead. These represented unnatural groupings of species by shared homoplasious morphological features: if the fore wing has two submarginal cells (Phileremus) instead of the usual three, if the maxillary palpus is three-segmented (Argyroselenis) rather than two-segmented (both states occur within Epeolus and Thalestriina, Rightmyer 2004), and if there is extensive red versus black integument coloration and reduced pubescence (Pyrrhomelecta).

Species of Epeolus are small to moderate-sized (body length 5.5–10.0 mm) relatively robust cleptoparasitic (epeoliform) bees. In North America, Epeolus may be confused with Triepeolus, which it resembles in general appearance, although Triepeolus may attain a much larger size (body length up to 18 mm in some species, Rightmyer 2008). The only other North American epeoline genus, Odyneropsis Schrottky, is rare (known only from the American Southwest) (Griswold and Parker 1999) and more likely to be confused with vespid wasps (hence the root ‘odynerus’) rather than Epeolus. Comprehensive overviews of the distinguishing features of Epeolus in reference to all other Epeolini are provided in Rightmyer (2004) and Michener (2007).

Diagnosis for Epeolus in North America

(Canada and the United States). Diagnostic for female Epeolus is a very distinct S6, which is usually retracted except sometimes for a pair of convergent spatulate lateral apical processes bearing setae modified into minute, pointed denticles (Onuferko 2017, Fig. 2A, B). Basally, the processes are separated by a large lobe-like disc, which in Triepeolus is reduced to a narrow transverse bar. In both Triepeolus (Onuferko 2017, Fig. 2C, D) and Odyneropsis, the lateral apical processes are subparallel and bear coarse, spine-like setae. Additionally, females may be separated on the basis of the pseudopygidial area (the apicomedial region of T5 that changes slope from the rest of the tergum), which in Epeolus is covered in a silvery band of short apically rounded setae. In Triepeolus, the pseudopygidial area is usually longer than in Epeolus and in most species the setae reflect a golden color. The T5 in female Odyneropsis is unique in that it is broadly notched posteriorly and has a distinct middorsal depressed area in the shape of a pointed oval outlined by ridges (Rightmyer 2004, fig. 180A).

Figure 2. 

Pygidial plate (in dorsal view) of male A E. australis (longer than wide and apically narrowed) B E. brumleyi paratype (nearly as long as wide and apically rounded) C E. flavofasciatus (longer than wide, with the lateral margins parallel) D E. asperatus (longer than wide and apically narrowed) E E. barberiellus (somewhat longer than wide and apically narrowed), and F T. concavus (longer than wide, with the lateral margins somewhat concave). Scale bars 1 mm.

Male Epeolus are more difficult to diagnose. As in females, the body lacks integumental white or yellow areas but the mesosoma and usually other tagmata have short appressed plumose white and/or yellow setae; the maxillary palpus is two or three segmented; the inner margins of the compound eyes are distinctly convergent below; the axilla is produced to a rounded lobe or angle or spine (i.e., not continuing the contour of the mesoscutellum); the distitarsi of all legs have arolia; the fore wing usually has three submarginal cells (if with two, then the second is at least nearly as long as the first), and the marginal cell is apically removed from the wing margin and much longer than the stigma; and a pygidial plate is present. In male Epeolus, the pygidial plate in most species is broadly rounded posteriorly (Fig. 2B); in Odyneropsis and Triepeolus it is usually more elongate and with a median constriction (Fig. 2F). It should be noted that males of some species of Epeolus in North America (notably E. australis Mitchell, E. flavofasciatus Smith, and some males in the “americanus group”) have a very narrow and distinctly Triepeolus-like pygidial plate (Fig. 2A, C, D), as opposed to the more broadly rounded/subtruncate pygidial plate typically associated with male Epeolus (Fig. 2B). The presence of a preapical tooth of the mandible (Fig. 3B, C, D, F) (often hidden from view because the mandibles are usually closed) confirms these and other species as Epeolus; all Triepeolus and only some Epeolus (in North America E. ainsliei, E. erigeronis, E. ilicis, E. inornatus, and E. zonatus) lack one (Fig. 3A, E) (Rightmyer 2004).

Figure 3. 

Mandible (in frontal view) of female A E. ainsliei without a preapical angulation or tooth B E. attenboroughi holotype with an inconspicuous, obtuse preapical tooth C E. carolinus with an inconspicuous, obtuse preapical tooth D E. gibbsi paratype with an obtuse angle appearing like a tooth E E. vernalis holotype (herein synonymized under E. ilicis) without a preapical angulation or tooth, and F E. compactus with a distinct preapical tooth with sides forming a right triangle. Scale bars 0.5 mm.

List of species with their proposed common names

Epeolus ainsliei Crawford, 1932 – Ainslie’s epeolus

Epeolus americanus (Cresson, 1878) – American epeolus

Epeolus andriyi Onuferko, sp. n. – Andrew’s epeolus

Epeolus asperatus Cockerell, 1909 – rough epeolus

Epeolus attenboroughi Onuferko, sp. n. – Attenborough’s epeolus

Epeolus australis Mitchell, 1962 – southern epeolus

Epeolus autumnalis Robertson, 1902 – fall epeolus

Epeolus axillaris Onuferko, sp. n. – spiny epeolus

Epeolus banksi (Cockerell, 1907) – Banks’ epeolus

Epeolus barberiellus Cockerell, 1907 – Barber’s epeolus

Epeolus basili Onuferko, sp. n. – Basil’s epeolus

Epeolus bifasciatus Cresson, 1864 – two-banded epeolus

Epeolus brumleyi Onuferko, sp. n. – Brumley’s epeolus

Epeolus canadensis Mitchell, 1962 – Canada epeolus

Epeolus carolinus Mitchell, 1962 – Carolina epeolus

Epeolus chamaesarachae Onuferko, sp. n. – five eyes crowned epeolus

Epeolus compactus Cresson, 1878 – compact epeolus

Epeolus deyrupi Onuferko, sp. n. – Deyrup’s epeolus

Epeolus diadematus Onuferko, sp. n. – Texas crowned epeolus

Epeolus erigeronis Mitchell, 1962 – fleabane epeolus

Epeolus ferrarii Onuferko, sp. n. – Ferrari’s epeolus

Epeolus flavofasciatus Smith, 1879 – yellow-banded epeolus

Epeolus floridensis Mitchell, 1962 – Florida epeolus

Epeolus gibbsi Onuferko, sp. n. – Gibbs’ epeolus

Epeolus glabratus Cresson, 1878 – smooth epeolus

Epeolus howardi Mitchell, 1962 – Howard’s epeolus

Epeolus ilicis Mitchell, 1962 – holly epeolus

Epeolus inornatus Onuferko, sp. n. – inornate epeolus

Epeolus interruptus Robertson, 1900 – interrupted epeolus

Epeolus lectoides Robertson, 1901 – Eastern prized epeolus

Epeolus lectus Cresson, 1878 – Great Plains prized epeolus

Epeolus mesillae (Cockerell, 1895) – Mesilla epeolus

Epeolus minimus (Robertson, 1902) – least epeolus

Epeolus nebulosus Onuferko, sp. n. – clouded epeolus

Epeolus novomexicanus Cockerell, 1912 – New Mexico epeolus

Epeolus olympiellus Cockerell, 1904 – Olympia epeolus

Epeolus packeri Onuferko, sp. n. – Packer’s epeolus

Epeolus pusillus Cresson, 1864 – dwarf epeolus

Epeolus rufulus Cockerell, 1941 – reddish epeolus

Epeolus scutellaris Say, 1824 – shield-backed epeolus

Epeolus splendidus Onuferko, sp. n. – polished epeolus

Epeolus tessieris Onuferko, sp. n. – Tessier’s epeolus

Epeolus zonatus Smith, 1854 – white-banded red epeolus

Epeolus ainsliei Crawford, 1932

Figs 3A, 4, 5, 95A

Epeolus ainsliei Crawford, 1932. Proc. Entomol. Soc. Wash. 34: 74 (♀).

Diagnosis

The following morphological features in combination can be used to tell E. ainsliei apart from all other North American Epeolus: the mandible lacks a preapical angle or tooth and the preoccipital ridge joins the hypostomal carina. In some specimens of E. scutellaris, the preoccipital ridge joins or nearly joins the hypostomal carina, in which case it is separated from the hypostomal carina by less than 1 MOD at its terminal, but the species has a blunt, obtuse preapical tooth on the mandible and the axillae are relatively straight along the medial margin whereas in E. ainsliei the free portion is distinctly hooked. Epeolus ainsliei is also very similar to E. attenboroughi and E. rufulus, which it resembles in that in all three species the axilla is dilated laterally and the free portion is distinctly hooked, and the T1–T4 apical fasciae are complete; however, in both E. attenboroughi and E. rufulus the mandible has a blunt, obtuse preapical tooth, the mesoscutum lacks the distinct paramedian bands present in E. ainsliei and is instead largely obscured by pale tomentum, and the preoccipital ridge does not join the hypostomal carina.

Figure 4. 

Epeolus ainsliei A female, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Redescription

This species was recently redescribed (Onuferko 2017).

Distribution

Great Plains to southwestern Ontario (Fig. 5).

Figure 5. 

Approximate geographic range of E. ainsliei (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: Epeolus ainsliei has been collected with possible host species Colletes susannae Swenk in Birds Hill Provincial Park (Gibbs et al. 2017) and Spruce Woods Provincial Park (J. Gibbs, personal communication, 2017), Manitoba, Canada and Spring Green Preserve in Sauk County, Wisconsin, USA (Wolf and Ascher 2009). In all cases at least one other species of Colletes was observed at the same locality and time as C. susannae and E. ainsliei, but observations of other Colletes were limited to one or two localities.

FLORAL RECORDS: Labels of examined voucher specimens indicate floral associations with Dalea purpurea Vent. (Leguminosae) and D. villosa (Nutt.) Spreng.

Discussion

Detailed morphological and taxonomic remarks about this species are given in Onuferko (2017).

Material studied

Type material. Primary: USA: Iowa: Sioux City, 15.vii.1922, C.N. Ainslie (holotype ♀ [USNM, catalog number: 534035]).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ACZ1957. Specimens examined and sequenced. Canada: Manitoba: 1♀ (PCYU); Birds Hill Provincial Park (50.0190°N; 96.8820°W) (Division 12), 05.viii.2017, J. Gibbs and Nozoe (1♀, JBWM); Ontario: Rondeau Provincial Park (42.2814°N; 81.8427°W) (Beach Access #10, near Visitor Centre), 08.viii.2017, R. Ferrari (1♂, PCYU).

Non-barcoded material examined

Canada: Alberta: 10♀, 1♂ (BBSL, CNC); Manitoba: Yellow Quill Mixed Grass Prairie Preserve (49.6911°N; 99.5747°W) (near Treesbank), 17.vii.2006, A.M. Patenaude (1♀, JBWM); Bald Head Hills (Spruce Woods Provincial Park), 01.viii.1983, W.E. Ralley (1♀, JBWM); Birds Hill Provincial Park (50.0100°N; 96.9100°W) (Division 12), 15.vii.2017, J. Gibbs and Nozoe (1♂, JBWM); Birds Hill Provincial Park (50.0115°N; 96.9065°W) (Division 12), 05.viii.2017, J. Gibbs and Nozoe (2♀, JBWM).

USA: Colorado: Longmont (Boulder County), 21.vii.1936, R. Bauer (1♂, CUM); Roggen, 08.vii.1933, M. and H. James and L. Ireland (1♂, CUM); Iowa: 1♀ (AMNH); Michigan: Edwin S. George Reserve (Livingston County), 12.viii.1960, U.N. Lanham (1♀, CUM); Minnesota: 1♀ (EMEC); Nebraska: 1♀ (AMNH); North Dakota: 7♀, 3♂ (AMNH, EMEC); Texas: 3♀, 2♂ (AMNH, CAS, CTMI); Wyoming: 1♀ (USNM).

Epeolus americanus (Cresson, 1878)

Figs 6, 7, 92K

Phileremus americanus Cresson, 1878. Trans. Am. Entomol. Soc. 7: 83 (♀, ♂); Cresson, 1916. Mem. Am. Entomol. Soc. 1: 111 (♀) [lectotype designation].

Phileremus montanus Cresson, 1878. Trans. Am. Entomol. Soc. 7: 83 (♂).

Epeolus lanhami Mitchell, 1962. N. C. Agric. Exp. Stn. Tech. Bull. 152: 450 (♀).

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. americanus apart from all other North American Epeolus except E. asperatus and E. barberiellus: in females, F2 is not more than 1.1 × as long as wide; the mesoscutum has distinct paramedian bands; the axilla is small to intermediate in size, not extending beyond the midlength of the mesoscutellum and the free portion is less than 1/4 as long as the entire medial length of the axilla, and like the mesoscutellum black; the mesopleuron is closely (i≤1d) and evenly punctate; T1 has a quadrangular discal patch, in dorsal view the longitudinal band is at least as wide as the breadth of the apical fascia; and the T1 and T2 apical fasciae are interrupted or at least greatly narrowed medially. Whereas in E. barberiellus the pronotal lobe and legs, at least from the tibiae to tarsi (sometimes the trochanters and femora as well), are reddish orange, in E. americanus the pronotal lobe and legs are brown or black. Epeolus americanus is also very similar to E. asperatus, but in E. asperatus the mesopleuron has much denser punctures ventrolaterally (most i<1d) than that of E. americanus and the T3 and T4 fasciae are never complete but broken or at least greatly narrowed laterally, as well as medially into separated or narrowly connected oval patches.

Figure 6. 

Epeolus americanus A female, lateral habitus (scale bar 3 mm) B female, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Redescription

This species was recently redescribed (Onuferko 2017).

Distribution

Widely distributed across Canada and the United States, including Alaska; not known to occur in parts of northeastern North America, the southeastern United States, or the high arctic (Fig. 7).

Figure 7. 

Approximate geographic range of E. americanus (orange) based on occurrence records known to the author (yellow circles).

Ecology

See Onuferko (2017) for host and floral records. Floral associations are also indicated in Suppl. material 1, which includes newly discovered associations with Leucanthemum vulgare (Vaill.) Lam. (Compositae), Plagiobothrys Fisch. & C.A. Mey. (Boraginaceae), Salix exigua Nutt. (Salicaceae), and S. interior Rowlee based on labels of examined voucher specimens.

Discussion

Detailed morphological and taxonomic remarks about this species are given in Onuferko (2017).

Material studied

Type material. Primary: USA: Colorado: H.K. Morrison (P. americanus lectotype ♀ [ANSP, catalog number: 2235]); Michigan: Near Saline, 26.vi.1954, U.N. Lanham (E. lanhami holotype ♀ [CUM, catalog number: 0000041]); Nevada: H. Edwards (P. montanus holotype ♂ [ANSP, catalog number: 2231]).

Secondary: USA: Michigan: Near Saline, 26.vi.1954, U.N. Lanham (E. lanhami allotype ♂ [CUM, catalog number: 0000042]).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:AAB9110. Specimens examined and sequenced. Canada: Quebec: 1♂ (RSKM); Yukon: 12♀, 2♂ (PCYU).

USA: Colorado: 2♀ (PCYU); Utah: 1♀ (BBSL).

Non-barcoded material examined

Canada: Alberta: 1♂ (CNC); British Columbia: 1♀, 2♂ (CNC); Manitoba: 1♀ (CNC); Adam Lake (Turtle Mountain Provincial Park), 27.vi.1987, T.D. Galloway (1♀, JBWM); Beaver Creek (Lake Winnipeg), 21.vi.1962, J.A. Garland (1♀, JBWM); Ontario: 6♀, 2♂ (CAS, CNC); Quebec: 1♀ (USNM); Saskatchewan: 2♀ (CNC); Yukon: 5♀, 1♂ (PCYU, RSKM).

USA: Alaska: 2♀, 3♂ (CNC); California: 1♂ (PCYU); 2 mi S Hilmar (Merced County), 14.iv.1961, R.R. Snelling (1♂, LACM); 3 mi SW Ash Creek (Siskiyou County), 16.vi.1974, D. Green (1♀, EMEC); Ash Creek Ranger Station (9 mi E McCloud, Siskiyou County), 07–09.vi.1974, J. Powell (1♂, EMEC), 10–12.vi.1974, R. Coville (4♀, 1♂, EMEC); Hayfork Ranger Station (Trinity County), 19.v.1973, J. Doyen (1♂, EMEC), 23.v.1973, J. Powell (1♀, EMEC); Independence Lake (Sierra County), 24.iv.1974, R.M. Bohart (1♂, UCBME); Lone Pine (Inyo County), 13.v.1969, J.A. Chemsak (1♀, EMEC); Sagehen Creek (Nevada County), 04.vii.??62, R.L. Westcott (1♀, LACM), 01.vii.??70, M.G. Axtman (1♂, LACM), 22.vi.1972, R.M. Bohart (1♀, 1♂, UCBME), 19.vi.1974, R.M. Bohart (4♀, UCBME), 23.vi.1976, N.J. Smith (1♀, UCBME), 23.vi.1976, R.M. Bohart (3♀, 2♂, UCBME), 23.vi.1976, R.M. Giblin (3♀, 1♂, UCBME), 23.vi.1976, R.E. Otondo (1♂, UCBME), 23.vi.1976, G.M. Streett (2♂, UCBME), 23.vi.1976, C.M. Bortfeid (1♂, UCBME), 30.vi.1976, N.J. Smith (1♀, UCBME), 14.vii.1976, R.M. Bohart (1♀, UCBME), 28.vi.1978, D.R. Smart (1♂, UCBME), 28.vi.1978, L.S. Kimsey (2♀, UCBME), 16.vii.1980, R.M. Bohart (1♀, UCBME); Colorado: 4♀ (PCYU); vi.1917 (1♀, AMNH); Cirque Meadows (Larimer County), 01.vii.1978, S. Hart (1♂, EMEC); Davenport Camp, 02.vii.1967, F., P., and M. Rindge (1♀, AMNH); Electra Lake, 28.vi.–01.vii.1919 (1♀, AMNH); Longmont (40.1507°N; 105.0385°W) (Weld County), 23.v.2012, V. Scott (1♂, CUM); Near Wolf Creek (37.4999°N; 106.7692°W) (Mineral County), 28.vii.2007, J. Gibbs and C. Sheffield (2♀, PCYU); Ouray (Summit road), 13.vii.1919 (1♂, AMNH); Idaho: 1♂ (USNM); Nevada: Reno, v.1940, U.N. Lar (1♀, CUM); Utah: 2♀ (PCYU); Virginia: 1♀ (USNM); Wyoming: 13 mi SE Cooke City, 27.vii.1962, F., P., and M. Rindge (1♀, AMNH); Yellowstone River (between Knowles Falls and Gardiner, Yellowstone National Park), 24.vi.1979, R.E. Dietz (1♂, EMEC).

Epeolus andriyi sp. n.

Figs 8, 9

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. andriyi apart from all other North American Epeolus: the axilla is large, with the tip extending well beyond the midlength of the mesoscutellum but not as far back as its posterior margin, dilated laterally but relatively straight along the medial margin, and like the mesoscutellum ferruginous; the axilla’s free portion is clearly less than 2/5 as long as its entire medial length; the mesopleuron is closely (i≤1d) and evenly punctate; the metasomal terga are black; T1 has a distinct basal fascia, which may be narrowly interrupted medially; the mesoscutum and metasomal terga have bands of bright or pale yellow short appressed setae; at least the T1–T3 apical fasciae are distinctly interrupted medially; and the pseudopygidial area of the female is lunate with the apex <2 × the medial length. Epeolus andriyi is most similar to E. howardi, but in E. howardi the axillae extend further posteriorly, as far back as or beyond the posterior margin of the mesoscutellum, and both the axillae and mesoscutellum are entirely red whereas in E. andriyi the mesoscutellum is dark brown or black along the anterior margin. Epeolus andriyi is also similar to E. scutellaris, but in E. scutellaris the T1–T3 apical fasciae are complete or only very narrowly interrupted medially, and the pseudopygidial area of the female is lunate with the apex >2 × the medial length.

Description

FEMALE: Length 8.2 mm; head length 1.9 mm; head width 2.6 mm; fore wing length 5.5 mm (margins of both worn in holotype).

Integument coloration. Mostly black; notable exceptions as follows: partially to entirely ferruginous on mandible, antenna, pronotal lobe, tegula, axilla, mesoscutum, mesoscutellum, mesopleuron, and legs. Mandible with apex darker than all but extreme base; preapical tooth lighter than mandibular apex. Antenna brown except scape, pedicel, and F1 extensively orange. F2 with orange spot basally. Pronotal lobe and tegula pale ferruginous to amber. Mesoscutum with reddish-brown spot anterolaterally between pronotal lobe and tegula. Wing membrane dusky subhyaline, slightly darker at apex. Legs more extensively reddish orange than brown or black.

Pubescence. Face with tomentum densest around antennal socket. Clypeus, upper paraocular and frontal areas, and vertexal area mostly exposed. Dorsum of mesosoma and metasoma with bands of off-white to pale yellow short appressed setae. Mesoscutum with paramedian band. Mesopleuron with upper half hairy, except beneath base of fore wing (hypoepimeral area); ventrolateral half nearly bare. Metanotum with tomentum sparser medially, uniformly off white. T1 with discal patch quadrangular and very wide, the basal and apical fasciae only narrowly joined laterally by few sparsely scattered pale hairs. T1–T3 with apical fasciae interrupted medially and narrowed before becoming somewhat broader laterally; T2 with fascia without anterolateral extensions of tomentum, although few sparsely scattered pale hairs present. T4 with fascia narrowed medially. T5 with two patches of pale tomentum (both quite faint in holotype because much of pubescence discolored or rubbed off) lateral to and contacting pseudopygidial area. T5 with pseudopygidial area lunate, its apex less than twice as wide as medial length, indicated by silvery setae on impressed disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs not extending beyond apex of sternum by more than 1/4 MOD.

Surface sculpture. Punctures dense. Labrum with larger and sparser punctures (i=1–2d) than clypeus (i<1d). Small impunctate matte spot lateral to lateral ocellus. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula densely punctate mesally (i≤1d), less so laterally (i=1–2d). Mesopleuron with ventrolateral half densely punctate (i≤1d), the interspaces shining; mesopleuron with punctures more or less equally dense throughout. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc; the interspaces shining somewhat.

Structure. Preapical tooth inconspicuous, blunt and obtuse. Labrum with pair of small subapical denticles, each preceded by small discrete longitudinal ridge. Frontal keel not strongly raised. Scape with greatest length 1.8 × greatest width. F2 noticeably longer than wide (L/W ratio = 1.5). Preoccipital ridge not joining hypostomal carina, from which it is separated by no less than 1 MOD at its terminal. Mesoscutellum weakly bigibbous. Axilla large, its lateral margin (L) half as long as mesoscutellar width (W) (L/W ratio = 0.5) and tip extending well beyond midlength of mesoscutellum but not as far back as its posterior margin; axilla with tip clearly visible, but unattached to mesoscutellum for less than 2/5 the medial length of axilla; axilla with lateral margin arcuate. Fore wing with three submarginal cells. Pygidial plate apically truncate.

MALE: Description as for female except for usual secondary sexual characters and as follows: F2 shorter, not noticeably longer than wide (L/W ratio = 1.1); S4 and S5 with much longer coppery to silvery subapical hairs; pygidial plate apically rounded, with large deep punctures more or less evenly spaced throughout, with the interspaces shining.

Figure 8. 

Epeolus andriyi A female holotype, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male allotype, lateral habitus (scale bar 3 mm), and D female holotype axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Etymology

This species is named in honor of my father, Rev. Andriy Onuferko, in gratitude for encouraging my interests in the natural world and for his assistance in collecting Epeolus in the field.

Distribution

Presently known from a single location along the Patuxent River in Maryland, USA (Fig. 9).

Figure 9. 

Occurrence record of E. andriyi known to the author (yellow circle).

Ecology

HOST RECORDS: The host species of E. andriyi is/are presently unknown.

FLORAL RECORDS: Unknown.

Discussion

Epeolus andriyi and E. howardi are very similar to one another, and both species have been collected in Maryland, USA in late August. Although E. andriyi is known from only two specimens, in both the axillae are shorter than in any examined specimen of E. howardi. The status of E. andriyi as a separate species is further supported by a separate BIN, but unusually its nearest neighbor is E. lectoides, from which E. andriyi exhibits a large barcode sequence divergence (7.1%).

Material studied

Type material. Primary: USA: Maryland: Jug Bay Wetlands Sanctuary (38.7839°N; 76.7014°W) (Anne Arundel County), 31.viii.2004, B. Hollister (♀ holotype [04-MD-1692], RSKM).

Secondary: USA: Maryland: Jug Bay Wetlands Sanctuary (38.7839°N; 76.7014°W) (Anne Arundel County), 31.viii.2004, B. Hollister (♂ allotype [04-MD-1691], RSKM).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:AAX7179. See Type material for specimens examined and sequenced (indicated by unique identifier number in square brackets).

Epeolus asperatus Cockerell, 1909

Figs 2D, 10, 11, 92L

Epeolus asperatus Cockerell, 1909. Ann. Mag. Nat. Hist. 5: 25 (♀).

Epeolus melectimimus Cockerell & Sandhouse, 1924. Proc. Calif. Acad. Sci. (4) 13: 317 (♂), syn. n.

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. asperatus apart from all other North American Epeolus except E. americanus and E. barberiellus: in females, F2 is not more than 1.1 × as long as wide; the mesoscutum has distinct paramedian bands; the axilla is small to intermediate in size, not extending beyond the midlength of the mesoscutellum and the free portion is less than 1/4 as long as the entire medial length of the axilla, and like the mesoscutellum black; the mesopleuron is closely (most i<1d) and evenly punctate; T1 has a quadrangular discal patch, in dorsal view the longitudinal band is at least as wide as the breadth of the apical fascia; and the T1 and T2 apical fasciae are interrupted or at least greatly narrowed medially. Whereas in E. barberiellus the legs, at least from the tibiae to tarsi (sometimes the trochanters and femora as well), are reddish orange and the metasomal terga are fasciate, in E. asperatus the legs are brown or black and the T3 and T4 fasciae are broken or at least greatly narrowed laterally, as well as medially into separated or narrowly connected oval patches. Epeolus asperatus is most similar to E. americanus, but in E. americanus the mesopleuron has sparser punctures ventrolaterally (i≤1d) than that of E. asperatus, with the interspaces shining, and the T3 and T4 fasciae are complete or broken medially and/or laterally, but rarely into separated oval patches.

Redescription

FEMALE: Length 7.8 mm; head length 2.0 mm; head width 2.8 mm; fore wing length 5.4 mm.

Integument coloration. Mostly black; notable exceptions as follows: at least partially ferruginous on mandible, labrum, antenna, pronotal lobe, tegula, and legs. Mandible with apex darker than rest of mandible; preapical tooth lighter than mandibular apex (difficult to see in the E. asperatus holotype; described from non-type specimens). Antenna brown except F1 and F2 orange in part. Flagellum slightly lighter than conspicuously dark brown scape and pedicel, primarily due to extensive pilosity on flagellum. Pronotal lobe and tegula pale ferruginous to amber. Wing membrane subhyaline, apically dusky. Legs with brown or black more extensive than reddish orange.

Pubescence. Face with tomentum densest around antennal socket. Dorsum of mesosoma and metasoma with bands of off-white to pale yellow short appressed setae. Mesoscutum with paramedian band. Mesopleuron with upper half hairy, ventrolateral half nearly bare. Metanotum with tomentum rubbed off medially in the E. asperatus holotype, but somewhat sparser medially and uniformly off white in non-type specimens. T1 with median quadrangular black discal patch enclosed by pale tomentum, except for medial separation at apex, and narrow, such that longitudinal band nearly half as wide as width of discal patch in dorsal view. T2–T4 with fasciae interrupted medially and with anterolateral extensions of sparser tomentum. T3 and T4 with fasciae also interrupted laterally, appearing as pair of oval patches between medial and lateral interruptions. T5 with two patches of pale tomentum lateral to and separate from pseudopygidial area (difficult to see in the E. asperatus holotype because T5 mostly retracted; described from non-type specimens). T5 with pseudopygidial area lunate, its apex more than twice as wide as medial length, indicated by silvery setae on impressed disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs not extending beyond apex of sternum by more than 1/4 MOD.

Surface sculpture. Punctures dense. Labrum with larger and sparser punctures (i=1–2d) than clypeus (i<1d). Small impunctate shiny spot lateral to lateral ocellus. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula very densely punctate (i<1d). Mesopleuron with ventrolateral half densely punctate (i<1d); mesopleuron with punctures more or less equally dense throughout. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc.

Structure. Preapical tooth with blunt tip. Labrum with pair of small subapical denticles, each preceded by small discrete longitudinal ridge. Frontal keel not strongly raised. Scape with greatest length 1.9 × greatest width. F2 as long as wide (L/W ratio = 1.0). Preoccipital ridge not joining hypostomal carina, from which it is separated by about 1.5–2 MOD at its terminal (difficult to see in the E. asperatus holotype; described from non-type specimens). Mesoscutellum moderately bigibbous. Axilla small to intermediate in size, its lateral margin (L) less than half as long as mesoscutellar width (W) (L/W ratio = 0.4) and tip not extending beyond midlength of mesoscutellum; axilla with tip visible, but unattached to mesoscutellum for less than 1/3 the medial length of axilla; axilla with lateral margin relatively straight and without carina. Fore wing with second submarginal crossvein incomplete in the E. asperatus holotype; with submarginal cells two or three and closed or second submarginal crossvein incomplete in non-type specimens. Pygidial plate apically truncate.

MALE: Description as for female except for usual secondary sexual characters and as follows: F2 shorter, nearly as long as wide (L/W ratio = 0.8); S4 and S5 with much longer coppery to silvery subapical hairs; pygidial plate V-shaped but apically rounded, with large deep, well-separated punctures, with the interspaces shining.

Figure 10. 

Epeolus asperatus A female, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Distribution

Central and southern California (Fig. 11).

Figure 11. 

Approximate geographic range of E. asperatus (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: Nine representatives of this species were collected at the Robert J. Bernard Biological Field Station in Claremont, California, USA in the spring of 2016 (see Material studied), and the only Colletes collected or observed was a single female of a predominantly black species with pale pubescence limited to the mesosoma. The collected female of the possible host species was barcoded, and using Stephen’s (1954) key identified as C. californicus Provancher. However, its sequence clusters with sequences of specimens collected in New Mexico (also in the spring of 2016) and identified as C. sphaeralceae Timberlake (with entirely/predominantly pale pubescence) through the use of Stephen’s (1954) key, dissection of the male terminalia, and collection from red Sphaeralcea A. St.-Hil. (Malvaceae) flowers, and all were assigned the same BIN (BOLD:ABZ4529). Another predominantly black female specimen from the San Diego National Wildlife Refuge Otay-Sweetwater Unit in California was barcoded (its image and 601 bp sequence are available on the Barcode of Life Data Systems website [http://www.barcodinglife.org/]), and was assigned the same BIN as the female from Claremont and specimens from New Mexico.

FLORAL RECORDS: Labels of examined voucher specimens indicate floral associations with Lasthenia Cass. (Compositae) and Plagiobothrys.

Discussion

Brumley (1965) synonymized E. asperatus and E. melectimimus under E. americanus, but current evidence suggests that the holotypes of E. asperatus and E. melectimimus belong to a cryptic species within the “americanus group”, distinct from E. americanus and E. barberiellus. In addition to the subtle diagnostic morphological features that separate E. asperatus from E. americanus and E. barberiellus, the status of E. asperatus as a separate species is supported by a separate BIN and large barcode sequence divergence (4.4%) from its nearest neighbor, E. barberiellus.

Epeolus melectimimus, with three submarginal cells, was described by Cockerell and Sandhouse (1924), who claimed that the species resembles a small Pseudomelecta Radoszkowski (a subgenus of Melecta Latreille in Michener 2007), from which it can be readily distinguished based on differences in the marginal cell. In the E. asperatus holotype, the second submarginal crossvein on each side is incomplete and inconspicuous. A series of E. asperatus was collected from the Robert J. Bernard Biological Field Station in Claremont, California, USA, which is in the same county as the type locality (Los Angeles). In some specimens, the fore wing has three submarginal cells whereas in others, the second submarginal crossvein is incomplete or lacking entirely. In some specimens, one fore wing has three submarginal cells and the other has an incomplete second submarginal crossvein. The male holotype of E. melectimimus was examined, and excluding sex-specific features the specimen with few exceptions agrees with the present redescription based on the female holotype of E. asperatus. Along with the abovementioned differences in wing venation, the pronotal lobe and tegula are darker in the holotype of E. melectimimus than in that of E. asperatus, but these differences fall within the range of observed intraspecific morphological variation among sequenced specimens. Although both E. americanus and E. asperatus are present in California, E. americanus appears to be absent from the southern part of the state.

Material studied

Type material. Primary: USA: California: Huntington Lake (Fresno County), 07.vii.1919, E.P. Van Duzee (E. melectimimus holotype ♂ [CAS, catalog number: 01612]); Los Angeles (Los Angeles County), 24.iv.1909, F. Grinnell, Jr. (E. asperatus holotype ♀ [USNM, catalog number: 534036]).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ACZ2142. Specimens examined and sequenced. USA: California: Robert J. Bernard Biological Field Station (Claremont, Los Angeles County), 18.iv.2002, M.G. Rightmyer (1♀, KUNHM); Robert J. Bernard Biological Field Station (34.1083°N; 117.7100°W) (Claremont, Los Angeles County), 13.iv.2016, T.M. Onuferko (2♂, PCYU).

Non-barcoded material examined

USA: California: 2 mi S Hilmar (Merced County), 19.iv.1960, R.R. Snelling (1♀, AMNH); 2 mi S Pearblossom (Los Angeles County), 01–02.v.1977, R.R. Snelling (1♂, LACM); Arroyo Seco Campground (Monterey County), 01.v.1960, F.D. Parker (1♂, UCBME), 19.v.1964, R.M. Bohart (1♂, UCBME), 11.v.1971, R.M. Bohart (3♀, 2♂, UCBME); Claremont (Los Angeles County), Baker (1♂, USNM), Metz (1♀, AMNH); Devore (San Bernardino County), 21.vi.1974, J.C. and E.M. Hall (1♂, UCR); East Fork Kaweah River (Tulare County), 02.vii.1976, T.L. Griswold (1♀, BBSL); Millard Canyon (Riverside County), 07.iv.1974, J.C. and E.M. Hall (1♀, UCR); Moreno Valley (base of Box Springs Mountains, Riverside County), 26.iv.1992, R.K. Velten (1♀, UCR); Robert J. Bernard Biological Field Station (34.1083°N; 117.7100°W) (Claremont, Los Angeles County), 13.iv.2016, T.M. Onuferko (2♀, 1♂, PCYU), 14.iv.2016, T.M. Onuferko (1♀, PCYU), 26.iv.2016, T.M. Onuferko (3♂, PCYU); W L Jepson Prairie Preserve (TNC) (13 mi S Dixon, Solano County), 20.v.1983, J.D. Barbour (1♂, UCBME).

Epeolus attenboroughi sp. n.

Figs 3B, 12, 13, 94B, 95B, 96A

Diagnosis

The following morphological features in combination can be used to tell E. attenboroughi apart from all other North American Epeolus except E. rufulus: the mandible has a blunt, obtuse preapical tooth; the preoccipital ridge does not join the hypostomal carina; the mesoscutum is largely obscured by pale tomentum; the axilla is elongate, extending well beyond the midlength of the mesoscutellum but not as far back as its posterior margin, and the free portion is distinctly hooked; the mesopleuron is closely (most i<1d) and evenly punctate; and T1–T4 have complete apical fasciae. Whereas in E. rufulus the discal patch is so wide that the longitudinal band is barely visible in dorsal view and in females F2 is noticeably longer than wide, in E. attenboroughi T1 has a comparatively narrow discal patch (the longitudinal band is more than half as wide as the breadth of the apical fascia in dorsal view) and in females F2 is less than 1.2 × as long as wide. Epeolus attenboroughi is also similar to E. ainsliei in that in both species the axilla is dilated laterally and the free portion is distinctly hooked, and the T1–T4 apical fasciae are complete; however, in E. ainsliei the mandible is simple, the preoccipital ridge joins the hypostomal carina, and the mesoscutum has distinct paramedian bands.

Description

FEMALE: Length 6.8 mm; head length 1.7 mm; head width 2.2 mm; fore wing length 4.5 mm.

Integument coloration. Black in part, at least partially ferruginous on mandible, labrum, clypeus, antenna, pronotal lobe, tegula, axilla, mesopleuron, legs, metasomal terga (including pygidial plate), and metasomal sterna. Mandible with apex darker than rest of mandible; preapical tooth slightly lighter than mandibular apex. Antenna brown and orange in part. Pronotal lobe and tegula pale ferruginous to amber. Wing membrane subhyaline, apically dusky. Legs entirely reddish orange.

Pubescence. Face with tomentum densest around antennal socket, slightly sparser on clypeus, upper paraocular and frontal areas, and vertexal area. Dorsum of mesosoma and metasoma with bands of off-white to pale yellow short appressed setae. Mesoscutum, mesoscutellum, and axilla largely obscured by pale tomentum. Mesopleuron densely hairy, except for sparsely hairy circular patch occupying much of ventrolateral half of mesopleuron. Metanotum with tomentum uninterrupted, uniformly off white. T1 with median quadrangular reddish-brown discal patch entirely enclosed by pale tomentum and narrow, such that longitudinal band more than half as wide as breadth of apical fascia in dorsal view. T2–T4 with fasciae complete, T2 with fascia with anterolateral extensions of sparser tomentum. T5 with two patches of pale tomentum lateral to and separate from pseudopygidial area. T5 with pseudopygidial area lunate, its apex more than twice as wide as medial length, indicated by silvery setae on impressed disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs extending beyond apex of sternum by ~1/3 MOD.

Surface sculpture. Punctures dense. Labrum and clypeus with punctures equally dense (i<1d). Impunctate spot lateral to lateral ocellus absent. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula very densely punctate (i<1d). Mesopleuron with ventrolateral half densely punctate (i<1d) to rugose; mesopleuron with punctures more or less equally dense throughout. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc.

Structure. Preapical tooth blunt and obtuse. Labrum with pair of small subapical denticles not preceded by carinae. Frontal keel not strongly raised. Scape with greatest length 1.7 × greatest width. F2 not noticeably longer than wide (L/W ratio = 1.1). Preoccipital ridge not joining hypostomal carina, from which it is separated by no less than 1 MOD at its terminal. Mesoscutellum weakly bigibbous. Axilla large, its lateral margin (L) more than half as long as mesoscutellar width (W) (L/W ratio = 0.6) and tip extending well beyond midlength of mesoscutellum but not as far back as its posterior margin; axilla with tip conspicuously diverging from side of mesoscutellum, distinctly hooked, and axilla with free portion approximately half its medial length; axilla with lateral margin arcuate and carinate. Fore wing with three submarginal cells. Pygidial plate apically truncate.

MALE: Description as for female except for usual secondary sexual characters and as follows: F2 shorter, as long as wide (L/W ratio = 1.0); mesopleuron almost entirely obscured by white tomentum; S4 and S5 with much longer coppery to silvery subapical hairs, which individually are often darker apically; pygidial plate apically rounded, with large deep, well-separated punctures, with the interspaces shining.

Etymology

This species is named in honor of English broadcaster and naturalist Sir David Attenborough in recognition of his inspiring books and television programs on natural history.

Figure 12. 

Epeolus attenboroughi A female holotype, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male allotype, lateral habitus (scale bar 3 mm), and D female holotype axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Distribution

New Mexico and southern Colorado (Fig. 13).

Figure 13. 

Occurrence records of E. attenboroughi known to the author (yellow circles).

Ecology

HOST RECORDS: The host species of E. attenboroughi is/are presently unknown.

FLORAL RECORDS: Unknown.

Discussion

Epeolus attenboroughi is similar in overall appearance to E. ainsliei and E. rufulus, and the ranges of the three species overlap to some extent. Although BIN-compliant sequences are presently not available for E. attenboroughi, partial sequences 421 bp and 289 bp in length are available for two specimens (male and female respectively) collected at the same locality and within one day of each other, and there is virtually no divergence (<1%) between the two. Moreover, the 421 bp sequence does not cluster closely with any sequences from other Epeolus species in a NJ tree of sequences >300 bp in length (Suppl. material 2). The longer of the two partial sequences is most similar (95.2%) to sequences from E. glabratus and E. lectoides (very different species).

In general, there is little morphological variation among examined specimens except in integument coloration; the axillae and mesoscutellum range from entirely black to partially ferruginous. Based on known records, adults of E. attenboroughi are active in summer.

Material studied

Type material. Primary: USA: Colorado: Great Sand Dunes National Monument (Alamosa County), 03–13.vii.1989, W.J. Bell (holotype ♀, KUNHM).

Secondary: USA: Colorado: Great Sand Dunes National Monument (Alamosa County), 10.vii.1991, B. Cutler (paratype ♀, KUNHM), 03–13.vii.1989, W.J. Bell (paratypes 1♀, 1♂, KUNHM), 11.vii.1991, B. Alexander and B. Cutler (allotype ♂, KUNHM), 11.vii.1991, B. Alexander and B. Cutler (paratypes 3♂, KUNHM); New Mexico: 24 km W Quemado (Catron County), 02.ix.1990, T.L. Griswold (paratype ♀, BBSL).

DNA barcoded material with BIN-compliant sequences

Unavailable.

Epeolus australis Mitchell, 1962

Figs 2A, 14, 15, 97I, 103A

Epeolus australis Mitchell, 1962. N. C. Agric. Exp. Stn. Tech. Bull. 152: 441 (♀).

Diagnosis

The following morphological features in combination can be used to tell E. australis apart from all other North American Epeolus: the frontal carina is strongly convex, such that the supraclypeal area is distinctly protuberant in lateral view; T1–T4 have complete fasciae; and the T2 fascia has a pair of anterolateral extensions of tomentum that are strongly convergent basally. In E. chamaesarachae and E. diadematus and commonly in E. bifasciatus the frontal carina is also strongly convex, but in the first two species the vertexal area has two pairs of shiny (usually impunctate) protrusions and in E. bifasciatus the frontal area bears a pair of granulose protrusions whereas in E. australis the frontal and vertexal areas lack protrusions. Epeolus australis most closely resembles E. brumleyi, but in E. brumleyi the frontal carina is only weakly convex and the pygidial plate of the male is wider (the medial length ≈ the basal width) than in E. australis (the medial length is ~1.5 × the basal width).

Redescription

FEMALE: Length 7.5 mm; head length 2.0 mm; head width 2.8 mm; fore wing length 5.7 mm.

Integument coloration. Mostly black; notable exceptions as follows: partially to entirely ferruginous on mandible, antenna, pronotal lobe, tegula, axilla, mesoscutellum, legs, pygidial plate, and metasomal sterna. Mandible with apex darker than rest of mandible; preapical tooth slightly lighter than mandibular apex. Both antennae missing in holotype, but brown and orange in part in paratype. Pronotal lobe and tegula pale ferruginous to amber. Wing membrane subhyaline, apically dusky. Legs more extensively reddish orange than brown or black.

Pubescence. Face with tomentum densest around antennal socket, slightly sparser on clypeus, upper paraocular and frontal areas, and vertexal area. Dorsum of mesosoma and metasoma with bands of off-white to pale yellow short appressed setae. Mesoscutum with paramedian band. Mesopleuron with upper half densely hairy, except beneath base of fore wing (hypoepimeral area); ventrolateral half sparsely hairy. Metanotum with tomentum uninterrupted, uniformly off white. T1 with discal patch elliptical and very wide, the basal and apical fasciae only narrowly joined laterally. T1 with basal and apical fasciae and T2–T4 with apical fasciae complete, T2 with fascia with basomedially convergent anterolateral extensions of tomentum. T5 with two large patches of pale tomentum lateral to and separate from pseudopygidial area, enclosing pseudopygidial area in triangle, except for medial separation at base. T5 with pseudopygidial area lunate, its apex more than twice as wide as medial length, indicated by silvery setae on disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs extending beyond apex of sternum by ~1/3 MOD.

Surface sculpture. Punctures dense. Labrum with larger punctures than clypeus, but punctures of both equally dense (i≤1d). Impunctate spot lateral to lateral ocellus absent in holotype, but shiny spot present in some non-type specimens. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula densely punctate mesally (i≤1d), less so laterally (i=1–2d). Mesopleuron with ventrolateral half densely punctate (i<1d); mesopleuron with punctures more or less equally dense throughout. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc.

Structure. Preapical tooth inconspicuous, blunt and obtuse. Labrum with pair of small subapical denticles (approximately at 1/4 length of labrum from apical margin) not preceded by carinae. Frontal keel strongly raised. Scape (missing in holotype) with greatest length 1.6 × greatest width in paratype. F2 (missing in holotype) not noticeably longer than wide (L/W ratio = 1.1) in paratype. Preoccipital ridge not joining hypostomal carina, from which it is separated by no less than 1 MOD at its terminal. Mesoscutellum moderately bigibbous. Axilla intermediate in size, its lateral margin (L) nearly half as long as mesoscutellar width (W) (L/W ratio = 0.4–0.5) and tip not extending beyond midlength of mesoscutellum; axilla with tip visible, but unattached to mesoscutellum for less than 2/5 the medial length of axilla; axilla with lateral margin relatively straight and without carina. Fore wing with three submarginal cells. Pygidial plate apically truncate.

MALE: Description as for female except for usual secondary sexual characters and as follows: F2 shorter, as long as wide (L/W ratio = 1.0); S4 and S5 with much longer coppery to silvery subapical hairs, which individually are often darker apically; pygidial plate unusually narrow (Triepeolus-like) and apically rounded, with large deep punctures closely clustered.

Figure 14. 

Epeolus australis A female, lateral habitus (scale bar 3 mm) B female, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Distribution

Mid-Atlantic states to Texas and presumably Mexico, given the close proximity of some collection localities (e.g., Eagle Pass, Texas) to the Mexico–United States border (Fig. 15).

Figure 15. 

Approximate geographic range of E. australis (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: The host species of E. australis is/are presently unknown.

FLORAL RECORDS: Mitchell (1962) indicated floral associations with Ceanothus L. (Rhamnaceae), Rubus L. (Rosaceae), Senecio L. (Compositae), and Specularia (now Triodanis? Raf. ex Greene) (Campanulaceae). Labels of examined voucher specimens further indicate associations with Chaetopappa asteroides (Nutt.) Nutt. ex DC. (Compositae), Hymenopappus artemisiifolius DC. (Compositae), and Sphaeralcea.

Discussion

This southeastern species displays minor sexual dimorphism in the coloration of the mesoscutellum, which is bright ferruginous in females and dark ferruginous to black in males. Otherwise, there is very little morphological variation among examined specimens. Although BIN-compliant sequences are presently not available for E. australis, 422 bp sequences were obtained from two male specimens (one from New Jersey, USA and one from South Carolina, USA), and there is virtually no divergence (<1%) between the two. Moreover, these sequences do not cluster with any sequences from other Epeolus species in a NJ tree (Suppl. material 2). Based on known records, adults of E. australis are active in spring.

Material studied

Type material. Primary: USA: North Carolina: Raleigh, 19.v.1950, T.B. Mitchell (holotype ♀, NCSU).

Secondary: USA: North Carolina: Raleigh, 09.v.1948, T.B. Mitchell (paratype ♀, NHMUK), 19.v.1950, T.B. Mitchell (paratype ♀, USNM).

DNA barcoded material with BIN-compliant sequences

Unavailable.

Non-barcoded material examined

USA: Florida: Alachua (Alachua County), 29.iv.1974, E.E. Grissell (2♀, UCBME); Georgia: Augusta (Richmond County), 18.v.1959, R.R. Snelling (1♀, LACM), 17.v.1959, R.R. Snelling (1♀, LACM), 03.v.1959, R.R. Snelling (1♂, LACM), 26.iv.1959, R.R. Snelling (1♂, LACM); Fort Gordon (Richmond County), 08.v.1958, R.R. Snelling (1♀, LACM); Maryland: Bowie (Prince George’s County), 08.vi.1968, R.R. Snelling (1♂, LACM); New Jersey: Forsythe (39.5296°N; 74.3421°W) (Atlantic and Ocean counties), 01–30.vi.2008, M. Springer (1♀, BIML); South Carolina: Carolina Sandhills National Wildlife Refuge (34.6043°N; 80.2469°W) (Chesterfield County), 18–19.v.2006, S.W. Droege (1♂, BIML); Texas: 10.7 mi S Dryden (Terrell County), 21.iv.1973, R.R. Snelling (1♂, LACM); 12 mi S Seguin (29.4060°N; 97.8550°W) (TX-123, Guadalupe County), 03.v.2014, J.L. Neff (1♀, CTMI); 8–25 km N Castroville (Medina County), 12.v.1988, B.N. Danforth (1♀, KUNHM); Camp Swift (30.2910°N; 97.3060°W) (Bastrop County), 24.iv.2003, J.L. Neff (1♀, CTMI); Eagle Pass (Maverick County), 28.iii.1946, C.D. Michener (2♂, AMNH); Hwy 83 (14 mi S Jct. Texas State Hwy 44, Webb County), 21.iv.1973, R.R. Snelling (1♀, LACM); Nacogdoches (Nacogdoches County), 14.iv.1960 (1♀, KUNHM); Stengl Lost Pines Research Station (30.0800°N; 97.1830°W) (Bastrop County), 02.iv.2006, J.L. Neff (1♀, CTMI).

Epeolus autumnalis Robertson, 1902

Figs 16, 17

Epeolus autumnalis Robertson, 1902. Entomol. News 13: 81 (♀, ♂). Webb, 1980. Ill. Nat. Hist. Surv. Bull. 32: 108 (♀) [lectotype designation (by W.E. LaBerge)].

Diagnosis

The following morphological features in combination can be used to tell E. autumnalis apart from all other North American Epeolus: the axilla is large, with the tip extending well beyond the midlength of the mesoscutellum but not as far back as its posterior margin, dilated laterally, and like the mesoscutellum black; the mesopleuron is closely (i≤1d) and evenly punctate; the T1 discal patch is so wide that the longitudinal band is barely visible in dorsal view; and the T2 fascia lacks lobe-like anterolateral extensions of tomentum, although a few sparsely scattered pale hairs are sometimes present. Epeolus autumnalis is similar to E. scutellaris in terms of surface sculpture and the patterns of pubescence on the mesosoma and metasoma, but in E. scutellaris at least the axilla is partially to entirely ferruginous (as is often the mesoscutellum), and the axilla is more elongate, extending to or beyond the band of pale tomentum along the posterior margin of the mesoscutellum.

Figure 16. 

Epeolus autumnalis A female, lateral habitus (scale bar 3 mm) B female, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Redescription

This species was recently redescribed (Onuferko 2017).

Distribution

Eastern North America (Fig. 17).

Figure 17. 

Approximate geographic range of E. autumnalis (orange) based on occurrence records known to the author (yellow circles).

Ecology

See Onuferko (2017) for host and floral records. Floral associations are also indicated in Suppl. material 1.

Discussion

Detailed morphological and taxonomic remarks about this species are given in Onuferko (2017).

Material studied

Type material. Primary: USA: Illinois: Carlinville (Macoupin County), C.A. Robertson (lectotype ♀ [INHS, catalog number: 44381]).

Secondary: USA: Illinois: Carlinville (Macoupin County), C.A. Robertson (lectoallotype ♂ [INHS, catalog number: 44382]).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:AAF2361. Specimens examined and sequenced. Canada: Nova Scotia: 2♀, 1♂ (PCYU, RSKM); Ontario: 1♀ (PCYU).

USA: New York: 1♀ (AMNH).

Non-barcoded material examined

Canada: Nova Scotia: 2♀ (PCYU, RSKM); Avonport (45.1189°N; 64.2634°W) (Kings County), 27.viii.2000, C. Sheffield (1♂, PCYU); Ontario: 14♀, 24♂ (DEBU, PCYU, ROM); King (44.0410°N; 79.5060°W), 23.viii.2000, V. Kushnir (1♂, PCYU); King (44.0430°N; 79.3100°W), 28.viii.2002, V. Kushnir (1♂, PCYU); King (44.0430°N; 79.5410°W), 06.ix.2003, J. Grixti (1♂, PCYU).

USA: Maryland: 2♂ (BIML); Massachusetts: 1♀, 2♂ (AMNH, BIML); New York: 1♀, 1♂ (AMNH, CAS); Lime Hollow (42.5650°N; 76.2550°W) (Cortland County), 03.ix.2011, J. Gibbs (1♂, JBWM); Virginia: Glencarlyn, 06.ix.???? (1♂, CUM).

Epeolus axillaris sp. n.

Figs 18, 19, 94A

Epeolus scopulus Brumley, 1965. M.S. thesis, Utah State University, Logan 66 (♀) [nomen nudum].

Diagnosis

Epeolus axillaris can be differentiated from all other Epeolus species in North America by the distinct posteromedial depression of the metanotum; in all other species the metanotum is flat, strongly convex, or weakly convex. Epeolus axillaris closely resembles E. banksi, E. minimus, and E. olympiellus in that the axilla (except sometimes the tip) and mesoscutellum are black; T1 has a quadrangular discal patch, in dorsal view the longitudinal band is at least half as wide as the breadth of the apical fascia; and the T2 fascia has lobe-like anterolateral extensions of tomentum. However, in all three species the metanotum is flat and the axilla does not extent much beyond the midlength of the mesoscutellum, whereas in E. axillaris the axilla is more elongate, extending well beyond the midlength of the mesoscutellum but not as far back as its posterior margin.

Description

FEMALE: Length 10.0 mm; head length 2.1 mm; head width 2.9 mm; fore wing length 6.9 mm.

Integument coloration. Mostly black; notable exceptions as follows: partially to entirely ferruginous on mandible, antenna, pronotal lobe, tegula, axilla, legs, T5, and pygidial plate. Mandible with apex darker than all but extreme base; preapical tooth slightly lighter than mandibular apex (difficult to see in holotype because mandible closed; described from paratypes). Flagellum brown and (except F1) slightly lighter than partially dark brown (otherwise orange) scape, pedicel, and F1, primarily due to extensive pilosity on flagellum. Axilla only with tip orange. Pronotal lobe and tegula pale ferruginous to amber. Wing membrane subhyaline, apically dusky. Legs, except reddish-orange mesotibia, metatibia, and tarsi, with brown or black more extensive than reddish orange.

Pubescence. Face with tomentum densest around antennal socket. Dorsum of mesosoma and metasoma with bands of off-white to pale yellow short appressed setae. Mesoscutum with paramedian band wider and joined posteriorly. Mesopleuron densely hairy, except for two sparsely hairy circular patches (one behind pronotal lobe, a larger one occupying much of ventrolateral half of mesopleuron). Metanotum with tomentum uninterrupted except for median bare patch in posterior half, uniformly off white. T1 with median quadrangular black discal patch enclosed by pale tomentum, except for medial separation at apex. T2–T4 with fasciae interrupted medially and narrowed before becoming somewhat broader laterally, T2 with fascia with anterolateral extensions of equally dense tomentum. T5 with two patches of pale tomentum bordering and separate from pseudopygidial area. T5 with pseudopygidial area lunate, its apex more than twice as wide as medial length, indicated by silvery setae on impressed disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs extending beyond apex of sternum by ~2/5 MOD.

Surface sculpture. Punctures dense. Labrum with larger and sparser punctures (i=1–2d) than clypeus (i<1d). Small impunctate shiny spot lateral to lateral ocellus. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula very densely punctate mesally (i<1d), less so laterally (i=1–2d). Mesopleuron largely obscured by tomentum, but ventrolateral half densely punctate (i<1d) to rugose where exposed; mesopleuron with punctures more or less equally dense throughout where exposed. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc.

Structure. Labrum with pair of small subapical denticles, each preceded by small discrete longitudinal ridge. Frontal keel not strongly raised. Scape with greatest length 1.7 × greatest width. F2 noticeably longer than wide (L/W ratio = 1.4). Preoccipital ridge not joining hypostomal carina, from which it is separated by about 1.5–2 MOD at its terminal. Mesoscutellum moderately bigibbous. Axilla large, its lateral margin (L) half as long as mesoscutellar width (W) (L/W ratio = 0.5) and tip extending well beyond midlength of mesoscutellum but not as far back as its posterior margin; axilla with tip conspicuously diverging from side of mesoscutellum, distinctly hooked, and axilla with free portion 2/5 its medial length; axilla with lateral margin relatively straight and without carina. Metanotum with posteromedial depression beneath overhanging anterior portion. Fore wing with three submarginal cells. Pygidial plate apically truncate.

MALE: Description as for female except for usual secondary sexual characters and as follows: F2 shorter, not noticeably longer than wide (L/W ratio = 1.1); S4 and S5 with much longer coppery to silvery subapical hairs; pygidial plate apically rounded, with large deep punctures more or less evenly spaced throughout, with the interspaces shining.

Figure 18. 

Epeolus axillaris A female paratype, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male allotype, lateral habitus (scale bar 3 mm), and D female paratype axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Etymology

The name is in reference to the axillae of this species, which are distinctly longer than those of the similar E. minimus and E. olympiellus.

Distribution

California and western Nevada. According to Brumley (1965), this species also ranges into Oregon, but its presence in that state could not be verified in the present study (Fig. 19).

Figure 19. 

Approximate geographic range of E. axillaris (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: The host species of E. axillaris is/are presently unknown.

FLORAL RECORDS: Labels of examined voucher specimens indicate floral associations with Chrysothamnus Nutt. (Compositae) (possibly in reference to plants that now are in the genus Ericameria Nutt. (Compositae)), Ericameria nauseosa var. nauseosa (Pall. ex Pursh) G.L. Nesom & Baird, E. nauseosa var. oreophila (A. Nelson) G.L. Nesom & Baird, and E. parryi (A. Gray) G.L. Nesom & Baird.

Discussion

This species is most similar to E. minimus and E. olympiellus, and there is overlap in the ranges of all three species. Brumley (1965) recognized E. axillaris as a separate species in which the axilla is more elongate and the metanotum is uniquely depressed posteromedially. The morphological distinction is supported by molecular data, as sequenced specimens exhibiting these attributes were assigned a separate BIN from either of the other two species.

Material studied

Type material. Primary: USA: Nevada: Cottonwood Creek (38.6013°N; 118.8280°W) (Mineral County), 14.viii.1998, F.D. Parker (holotype ♀ [CCDB-28237 D01], BBSL).

Secondary: USA: California: Antioch (Contra Costa County), x.1938, J.A. Downes (paratype ♂, CNC), 10.ix.1947, P.D. Hurd (paratype ♂, BBSL), 10.ix.1947, U.N. Lanham (paratype ♀, CUM); Bodie (Mono County), 21.ix.1958, A.S. Menke and L.A. Stange (paratype ♀, LACM); Hot Creek (Mono County), 29.viii.1969, E.E. Grissell (paratypes 3♀, UCBME), 29.viii.1969, R.M. Bohart (paratype ♂, UCBME); Parker Creek at Walker Lake Road (37.8768°N; 119.1203°W) (Mono County), 02.ix.2009, G.R. Ballmer (allotype ♂ [CCDB-28313 H10], UCR), 02.ix.2009, G.R. Ballmer (paratypes 2♂ (1 barcoded [CCDB-28313 H08]), UCR); Upper Santa Ana River (San Bernardino County), 22.ix.1946, G.H. and J.L. Sperry (paratype ♂, KUNHM); Nevada: 17 mi N Sparks (Washoe County), 02.ix.1957, E.G. Linsley (paratype ♀, BBSL), 02.ix.1957, E.G. Linsley (paratype ♀, USNM); 3 mi N Minden (Douglas County), 10.ix.1957, R.C. Bechtel (paratype ♀, AMNH); Reno, 09.ix.1961, F.D. Parker (paratype ♂, UCBME).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ACZ2412. See Type material for specimens examined and sequenced (indicated by unique CCDB-plate and well number).

Epeolus banksi (Cockerell, 1907)

Figs 20, 21, 96F

Triepeolus banksi Cockerell, 1907a. Entomologist 40: 135 (♂).

Epeolus banksi Mitchell, 1962. N. C. Agric. Exp. Stn. Tech. Bull. 152: 442.

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. banksi apart from all other North American Epeolus except E. minimus and E. olympiellus: in females, F2 is at least 1.2 × as long as wide; the mesoscutum has distinct paramedian bands; the axilla is small to intermediate in size, not extending much beyond the midlength of the mesoscutellum (extending to <2/3 its length) but the free portion is more than 1/4 as long as the entire medial length of the axilla, and the axilla and mesoscutellum are black; the mesopleuron is closely (most i<1d) and evenly punctate; T1 has a quadrangular discal patch, in dorsal view the longitudinal band is at least half as wide as the breadth of the apical fascia; and the T2 fascia has anterolateral extensions of tomentum. Whereas in E. minimus and E. olympiellus the mesoscutum and metasomal terga have bands of off-white to pale yellow short appressed setae, in E. banksi the mesoscutum and metasomal terga have bands of gray short appressed setae. In E. banksi, the integument is entirely dark brown or black. In E. olympiellus, at least the pronotal lobe is ferruginous. In E. minimus from California, the integument is often entirely dark brown or black, but throughout most of its range E. minimus exhibits reddish-orange coloration on the labrum, antenna, pronotal lobe, and/or legs, except foreleg, from trochanters to tarsi. Both sexes of E. banksi are larger (~10 mm in length) on average than E. minimus or E. olympiellus (7–8 mm in length).

Redescription

MALE: Length 9.4 mm; head length 2.3 mm; head width 3.3 mm; fore wing length 7.5 mm.

Integument coloration. Mostly black; notable exceptions as follows: at least partially ferruginous on mandible, antenna, tegula, and legs. Mandible black except apex reddish brown; preapical tooth same color as mandibular apex (difficult to see in holotype; described from non-type specimens). Flagellum, except right F1 and F2, missing in holotype, but brown and (except F1) slightly lighter than conspicuously dark brown scape and pedicel, primarily due to extensive pilosity on flagellum, in non-type specimens. Wing membrane subhyaline, apically dusky. Legs, except reddish-orange tarsi, with brown or black more extensive than reddish orange.

Pubescence. Face with tomentum densest on clypeus and around antennal socket, sparser on upper paraocular area and vertexal area. Dorsum of mesosoma and metasoma with bands of off-white to pale gray short appressed setae. Mesoscutum with paramedian band. Mesopleuron densely hairy, except for two sparsely hairy circular patches (one behind pronotal lobe, a larger one occupying much of ventrolateral half of mesopleuron). Metanotum with tomentum uninterrupted, uniformly off white. T1 with median quadrangular black discal patch enclosed by pale tomentum, except for medial separation at apex. T2–T6 with fasciae interrupted medially, those of T2–T4 narrowed before becoming somewhat broader laterally, T2 with fascia with anterolateral extensions of sparser tomentum. S4 and S5 with long coppery to silvery subapical hairs, which individually are often darker apically.

Surface sculpture. Punctures dense. Labrum with larger and sparser punctures (i=1–2d) than clypeus (i<1d). Small impunctate matte spot lateral to lateral ocellus. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula very densely punctate mesally (i<1d), less so laterally (i=1–2d). Mesopleuron with ventrolateral half densely punctate (i<1d); mesopleuron with punctures more or less equally dense throughout. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc.

Structure. Labral apex with pair of small denticles, each preceded by longitudinal carina. Frontal keel not strongly raised. Scape with greatest length 1.6 × greatest width. F2 noticeably longer than wide (L/W ratio = 1.2). Preoccipital ridge not joining hypostomal carina, from which it is separated by about 1.5–2 MOD at its terminal. Mesoscutellum moderately bigibbous. Axilla intermediate in size, its lateral margin (L) nearly half as long as mesoscutellar width (W) (L/W ratio = 0.4–0.5) and tip not extending much beyond midlength of mesoscutellum (extending to <2/3 its length); axilla with tip clearly visible, but unattached to mesoscutellum for less than 2/5 the medial length of axilla; axilla with lateral margin relatively straight and without carina. Fore wing with three submarginal cells. Pygidial plate apically rounded, with large deep punctures closely clustered.

FEMALE: Description as for male except for usual secondary sexual characters and as follows: F2 even longer than wide (L/W ratio = 1.4); T5 with two patches of pale tomentum bordering and separate from pseudopygidial area present only in female; T5 with pseudopygidial area lunate, its apex more than twice as wide as medial length, indicated by silvery setae on flat disc of apicomedial region elevated from rest of tergum; S4 and S5 with much shorter hairs (S5 with apical fimbria of coppery to silvery hairs extending beyond apex of sternum by ~2/5 MOD); pygidial plate apically truncate, with small, denser punctures.

Figure 20. 

Epeolus banksi A female, lateral habitus (scale bar 3 mm) B female, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Distribution

Maryland to North Carolina (Fig. 21).

Figure 21. 

Occurrence records of E. banksi known to the author (yellow circles).

Ecology

HOST RECORDS: The host species of E. banksi is/are presently unknown.

FLORAL RECORDS: Mitchell (1962) indicated a floral association with Fragaria L. (Rosaceae). Labels of examined voucher specimens further indicate associations with Solidago L. (Compositae) and Symphyotrichum ericoides (L.) G.L. Nesom (Compositae).

Discussion

Most of the specimens of this species that were examined were collected in the Washington metropolitan area. While Mitchell (1962) indicated Epeolus banksi as being quite prevalent across the Eastern United States, reportedly ranging from Minnesota to New Jersey and North Carolina, it seems that the name has been commonly misapplied to specimens of E. minimus (as in MacKay and Knerer (1979) for example, and probably by Mitchell (1962) as well). Epeolus banksi is much larger than E. minimus, and has completely black integument, but unlike similarly dark specimens of E. minimus from California, E. banksi has gray as opposed to pale yellow bands of tomentum on the mesosoma and metasoma. Unfortunately, no recently collected material was available for barcode sequencing, and the specimens seen are all from the early 1900s. The absence of this species from recent collections has not gone unnoticed (e.g in Colla et al. 2012 it is listed among the bee species not collected since 1990). Increased urbanization in and around Washington D.C. may have resulted in the extirpation of this species there, and perhaps it has even disappeared entirely throughout its earlier range. Hence, extensive efforts should be made to rediscover this species, by sampling its apparent historical range between North Carolina and Maryland, to assess its conservation status. The flight season of E. banksi appears to be late summer/early autumn.

Material studied

Type material. Primary: USA: Virginia: Falls Church, 26.viii.????, N. Banks (holotype ♂ [USNM, catalog number: 534038]).

Secondary: USA: Virginia: Falls Church, 07.ix.????, N. Banks (paratype ♂, CAS).

DNA barcoded material with BIN-compliant sequences

Unavailable.

Non-barcoded material examined

USA: Maryland: Glen Echo (Montgomery County), 30.viii.1923, J.R. Malloch (1♂, USNM); North Carolina: Valley of Black Mountains, 30.ix.1906, W. Beutenmuller (1♂, AMNH); Virginia: Chain Bridge, 10.ix.1922, J.R. Malloch (1♂, USNM); Falls Church, G.G. Rohwer (1♂, USNM); Glencarlyn?, 20.ix.??30 (1♂, USNM); Washington, D.C. (2♀, BBSL); Rock Creek Park, 28.viii.1919, J.C. Crawford (1♂, AMNH).

Epeolus barberiellus Cockerell, 1907

Figs 2E, 22, 23, 96E

Epeolus barberiellus Cockerell, 1907b. Entomologist 40: 266 (♀).

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. barberiellus apart from all other North American Epeolus except E. americanus and E. asperatus: in females, F2 is not more than 1.1 × as long as wide; the mesoscutum has distinct paramedian bands; the axilla is small to intermediate in size, not extending beyond the midlength of the mesoscutellum and the free portion is less than 1/4 as long as the entire medial length of the axilla, and like the mesoscutellum black; the mesopleuron is closely (i≤1d) and evenly punctate; T1 has a quadrangular discal patch, in dorsal view the longitudinal band is at least as wide as the breadth of the apical fascia; and the T1 and T2 apical fasciae are interrupted or at least greatly narrowed medially. In E. asperatus the mesopleuron has much denser punctures ventrolaterally (most i<1d) than that of E. barberiellus and the T3 and T4 fasciae are never complete but broken or at least greatly narrowed laterally, as well as medially into separated or narrowly connected oval patches. Epeolus barberiellus is most similar to E. americanus, but in E. americanus the pronotal lobe and legs are brown or black, not reddish orange.

Redescription

FEMALE: Length 5.7 mm; head length 1.8 mm; head width 2.3 mm; fore wing length 5.0 mm.

Integument coloration. Mostly black; notable exceptions as follows: at least partially ferruginous on mandible, labrum, antenna, pronotal lobe, tegula, mesopleuron, metapleuron, propodeum, legs, metasomal terga (including pygidial plate), and metasomal sterna. Mandible with apex darker than rest of mandible; preapical tooth as dark as mandibular apex (difficult to see in holotype because mandible closed; described from non-type specimens). Pedicel and flagellum brown and orange in part, slightly lighter than dark brown scape. Pronotal lobe reddish brown. Tegula pale ferruginous to amber. Wing membrane subhyaline, apically dusky. Legs more extensively reddish orange than brown or black. T5 and pygidial plate reddish orange.

Pubescence. Face with tomentum densest around antennal socket. Dorsum of mesosoma and metasoma with bands of off-white to pale yellow short appressed setae. Mesoscutum with paramedian band and moderately dense pale tomentum along margins. Mesopleuron densely hairy, except for almost entirely bare circular patch occupying much of ventrolateral half of mesopleuron. Metanotum with tomentum uninterrupted, uniformly off white. T1 with median quadrangular reddish-brown discal patch enclosed by pale tomentum, except for medial separation at apex, and narrow, such that longitudinal band more than half as wide as width of discal patch in dorsal view. T2 with fascia interrupted medially and without anterolateral extensions of tomentum, although fascia broader laterally with hairs sparser basally. T3 and T4 with fasciae complete and narrowed laterally. T5 with two patches of pale tomentum lateral to and separate from pseudopygidial area. T5 with pseudopygidial area lunate, its apex more than twice as wide as medial length, indicated by silvery setae on impressed disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs not extending beyond apex of sternum by more than 1/4 MOD.

Surface sculpture. Punctures dense. Labrum with larger and sparser punctures (i=1–2d) than clypeus (i<1d). Impunctate spot lateral to lateral ocellus absent in holotype, but shiny spot present in non-type specimens. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula densely punctate mesally (i≤1d), less so laterally (i=1–2d). Mesopleuron with ventrolateral half densely punctate (i≤1d), the interspaces shining; mesopleuron with punctures more or less equally dense throughout. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc.

Structure. Labrum with pair of small subapical denticles not preceded by carinae. Frontal keel not strongly raised. Scape with greatest length 1.9 × greatest width. F2 as long as wide (L/W ratio = 1.0). Preoccipital ridge not joining hypostomal carina, from which it is separated by about 1.5–2 MOD at its terminal (difficult to see in holotype; described from non-type specimens). Mesoscutellum moderately bigibbous. Axilla small to intermediate in size, its lateral margin (L) less than half as long as mesoscutellar width (W) (L/W ratio = 0.3) and tip not extending beyond midlength of mesoscutellum; axilla with tip visible, but unattached to mesoscutellum for less than 1/4 the medial length of axilla; axilla with lateral margin relatively straight and without carina. Fore wing with three submarginal cells. Pygidial plate apically truncate.

MALE: Description as for female except for usual secondary sexual characters and as follows: F2 shorter, nearly as long as wide (L/W ratio = 0.8); S4 and S5 with much longer coppery to silvery subapical hairs, which individually are often darker apically; pygidial plate orange and V-shaped but apically rounded, with large deep punctures closely clustered.

Figure 22. 

Epeolus barberiellus A female, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female holotype axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Distribution

Arizona to west Texas (Fig. 23).

Figure 23. 

Approximate geographic range of E. barberiellus (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: The host species of E. barberiellus is/are presently unknown.

FLORAL RECORDS: Labels of examined voucher specimens indicate floral associations with Aster (possibly in reference to a plant that is in a different genus now) (Compositae) and Sphaeralcea.

Discussion

Epeolus barberiellus is most similar to E. americanus, from which it differs consistently only in integument coloration. Although sequenced representatives of both forms share the same BIN, specimens identified as E. barberiellus cluster separately from those identified as E. americanus (Suppl. material 2). Whereas E. americanus is widely distributed across North America, E. barberiellus appears to be restricted to the Southwestern United States (and possibly adjacent Mexico), where it replaces the much darker form that characterizes E. americanus. Taken together, these differences are indicative of divergence, and therefore the two forms are herein considered to be heterospecific. Brumley (1965) also considered E. americanus and E. barberiellus as separate species, but synonymized E. asperatus and E. melectimimus under E. americanus. In the present study, three valid species in the “americanus group” (E. americanus, E. asperatus, and E. barberiellus) are recognized, of which only E. asperatus has been assigned a separate BIN, suggesting that E. americanus and E. barberiellus are sister species.

The male of E. barberiellus is described here for the first time. Of the Epeolus in the “americanus group”, this appears to be the least commonly collected species.

Material studied

Type material. Primary: USA: New Mexico: Mesilla Park, 22.iv.????, C.M. Barber (holotype ♀ [USNM, catalog number: 534039]).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:AAB9110. Specimens examined and sequenced.—USA: New Mexico: Sagebrush Valley Rd (32.9500°N; 104.8333°W) (Artesia), 01–10.v.2004, M.E. Irwin (1♂, BBSL).

Non-barcoded material examined

USA: Arizona: 2 mi SW Apache (Cochise County), 19.iv.1961, Gertsch, Rozen, and Schrammel (1♀, AMNH); 31 mi N Wickenburg, 21.iv.1967, P. Torchio and N. Youssef (1♂, LACM); 40 mi S Kingman (Mohave County), 21.iv.1967, P. Torchio and N. Youssef (1♀, BBSL); New Mexico: 12 mi N Las Cruces (Doña Ana County), 11.iv.1965, F.D. Parker (1♂, BBSL); Texas: 9.4 mi E Cornudas (Hudspeth County), 27.iv.1998, T., S., and L. Griswold (1♀, BBSL).

Epeolus basili sp. n.

Figs 24, 25, 97D, 98B

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. basili apart from all other North American Epeolus except E. nebulosus, E. novomexicanus, and E. pusillus: the axilla is large, with the tip extending well beyond the midlength of the mesoscutellum but at most to the band of pale tomentum along its posterior margin, dilated laterally, and usually ferruginous to some degree (rarely all black) whereas the mesoscutellum ranges from entirely black to partially ferruginous; the axilla’s free portion is clearly less than 2/5 as long as its entire medial length; the mesopleuron is closely (most i<1d) and evenly punctate, that of the female is obscured by white tomentum only in the upper half (with a large, sparsely hairy circle occupying much of the ventrolateral half) whereas that of the male (excluding the hypoepimeral area) is entirely obscured by white tomentum; the T1–T3 apical fasciae are complete or only very narrowly interrupted medially; the T2 fascia has lobe-like anterolateral extensions of tomentum; and the pseudopygidial area of the female is lunate with the apex at least 2 × and clearly <2.5 × the medial length. Epeolus basili, E. nebulosus, E. novomexicanus, and E. pusillus are all extremely similar to one another. Whereas in E. pusillus the flagellum, except sometimes F1, and metasomal sterna are consistently brown or black and clearly not the same reddish-orange color as the legs (tibiae to tarsi), in E. basili the flagellum, at least ventrally, is the same reddish-orange color as the legs (tibiae to tarsi) as are usually the metasomal sterna. In E. nebulosus and E. novomexicanus the T2–T4 fasciae are on or very little removed from the apical margin, and in both species as well as in E. pusillus the pseudopygidial area of the female is commonly less and no more than 2 × the medial length. By contrast, in E. basili the T2 and T3 (for female) or T2–T4 (for male) fasciae are narrowed medially and removed from the apical margin, and the pseudopygidial area of the female is ≥2 × the medial length. Epeolus basili is also similar to E. scutellaris in that the axilla is large, with the lateral margin arcuate, and that the apical fasciae are complete or only very narrowly interrupted medially. However, in E. scutellaris the pseudopygidial area of the female is even wider (the apex ~2.5–3 × the medial length) than in E. basili, and the mesopleuron of both the female and male is obscured by white tomentum only in the upper half (with a large, sparsely hairy circle occupying much of the ventrolateral half).

Description

FEMALE: Length 7.0 mm; head length 1.8 mm; head width 2.5 mm; fore wing length 4.8 mm.

Integument coloration. Mostly black; notable exceptions as follows: at least partially ferruginous on mandible, labrum, antenna, pronotal lobe, tegula, axilla, legs, and metasomal sterna. Mandible with apex darker than rest of mandible; preapical tooth slightly lighter than mandibular apex (difficult to see in holotype; described from paratypes). Antenna brown and orange in part. Pronotal lobe and tegula pale ferruginous to amber. Wing membrane subhyaline, apically dusky. Legs more extensively reddish orange than brown or black. S1–S5 reddish orange.

Pubescence. Face with tomentum densest around antennal socket, slightly sparser on clypeus, upper paraocular and frontal areas, and vertexal area. Dorsum of mesosoma and metasoma with bands of off-white to pale yellow short appressed setae. Mesoscutum with paramedian band. Mesopleuron densely hairy, except for sparsely hairy circular patch occupying much of ventrolateral half of mesopleuron. Metanotum with tomentum uninterrupted, uniformly off white. T1 with discal patch quadrangular and very wide, the basal and apical fasciae only narrowly joined laterally. T1–T3 with apical fasciae complete (basal fascia of T1 also), narrowed medially, and removed from apical margin, most noticeably at midline; T2 with fascia with anterolateral extensions of tomentum. T4 with fascia complete. T5 with large, continuous patch of pale tomentum bordering and separate from pseudopygidial area. T5 with pseudopygidial area lunate, its apex twice as wide as medial length, indicated by silvery setae on flat disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs extending beyond apex of sternum by ~2/5 MOD.

Surface sculpture. Punctures dense. Labrum with larger and sparser punctures (i=1–2d) than clypeus (i<1d). Small impunctate shiny spot lateral to lateral ocellus. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula densely punctate mesally (i≤1d), less so laterally (i=1–2d). Mesopleuron with ventrolateral half densely punctate (i≤1d) to rugose; mesopleuron with punctures more or less equally dense throughout. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc.

Structure. Preapical tooth obtuse. Labrum with pair of small subapical denticles not preceded by carinae. Frontal keel not strongly raised. Scape with greatest length 1.9 × greatest width. F2 noticeably longer than wide (L/W ratio = 1.4). Preoccipital ridge not joining hypostomal carina, from which it is separated by no less than 1 MOD at its terminal. Mesoscutellum weakly bigibbous. Axilla large, its lateral margin (L) half as long as mesoscutellar width (W) (L/W ratio = 0.5) and tip extending well beyond midlength of mesoscutellum but not as far back as its posterior margin; axilla with tip clearly visible, but unattached to mesoscutellum for less than 2/5 the medial length of axilla; axilla with lateral margin arcuate. Fore wing with three submarginal cells. Pygidial plate apically truncate.

MALE: Description as for female except for usual secondary sexual characters and as follows: F2 shorter, but still longer than wide (L/W ratio = 1.2); mesopleuron (excluding hypoepimeral area) entirely obscured by white tomentum; S4 and S5 with much longer coppery to silvery subapical hairs; pygidial plate apically rounded, with large deep, well-separated punctures, with the interspaces shining.

Figure 24. 

Epeolus basili A female holotype, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male allotype, lateral habitus (scale bar 3 mm), and D female paratype axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Etymology

This species is named in honor of my brother, Basil V. Onuferko (1986–2013).

Distribution

Northwestern Mexico and southwestern United States (Fig. 25).

Figure 25. 

Approximate geographic range of E. basili (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: This species has been collected east of Willcox, Arizona, USA in the presence of large numbers of Colletes tectiventris Timberlake (E. Wyman, personal communication, 2014).

FLORAL RECORDS: Labels of examined voucher specimens indicate floral associations with Isocoma hartwegii (A. Gray) Greene (Compositae), I. tenuisecta Greene, Pectis papposa Harv. & A. Gray (Compositae), Psorothamnus scoparius (A. Gray) Rydb. (Leguminosae), and Wislizenia refracta Engelm. (Cleomaceae).

Discussion

Structurally, this species is indistinguishable from the other three members of the “pusillus group”, and although consistent, the features (differences in integument coloration and patterns of pubescence) that in combination may be used to distinguish E. basili from E. nebulosus, E. novomexicanus, and E. pusillus are subtle. Its status as a separate species is supported by a separate BIN and large barcode sequence divergence (>7.3%) from its nearest neighbor, E. pusillus. In the United States, Epeolus basili appears to be restricted to parts of the American Southwest, east of California.

Material studied

Type material. Primary: USA: Arizona: 4 mi E Willcox (Cochise County), 29.viii.2013, J.S. Ascher (holotype ♀ [CCDB-22791 A05], AMNH).

Secondary: Mexico: Chihuahua: 9 mi S Hidalgo del Parral, 31.vii.1967, R.C. Gardner, C.R. Kovacic, and K. Lorenzen (paratype ♂, UCBME); Durango: Nombre de Dios, 01.viii.1951, P.D. Hurd (paratypes 1♀, 1♂, EMEC); Otinapa, 11.viii.1947, D. Rockefeller Exp. Michener (paratype ♀, AMNH); Tepehuanes, 1933, Wickham (paratype ♀, USNM).

USA: Arizona: 11 mi S San Simon, 02.ix.2013, G. Rowe (paratype ♀, PCYU); 1–3 mi SE Willcox (Cochise County), 25.viii.1994, J.G. Rozen and J.S. Ascher (paratype ♂, AMNH); 2 mi SE Willcox (Cochise County), 05.ix.1986, J.G. and B.L. Rozen (paratype ♀, AMNH); 4 mi E Willcox (Cochise County), 02.ix.2013, C. Lin (paratype ♂, AMNH), 02.ix.2013, Z. Soh (paratypes 2♂, AMNH), 03.ix.2015, R. González Vaquero (paratype ♂, PCYU), 06.ix.2012, J.G Rozen (paratypes 2♀, AMNH), 09.ix.1991, J.G. and B.L. Rozen (paratype ♀, AMNH), 11.ix.1991, J.G. and B.L. Rozen (paratypes 1♀, 2♂, AMNH), 16.ix.2012, E.S. Wyman (paratypes 2♂, AMNH), 16.ix.2012, J.G. and M.A. Rozen (paratype ♀, AMNH), 26.viii.1994, J.G. Rozen and J.S. Ascher (paratypes 3♂, AMNH), 27.viii.2013, E.S. Wyman (allotype ♂ [CCDB-22791 A11], AMNH), 27.viii.2013, E.S. Wyman (paratypes 8♂, AMNH), 27.viii.2013, W.J. Cromartie (paratype ♂, AMNH), 27.viii.2013, G. Rowe (paratypes 7♂ (1 barcoded [CCDB-24580 G03]), PCYU), 28.viii.1985, J.G. and B.L. Rozen (paratypes 8♂, AMNH), 29.viii.2013, J.S. Ascher (paratypes 3♂, AMNH), 30.viii.1993, J.G. Rozen (paratypes 1♀, 9♂, AMNH); E Moore Ranch Rd (32.2391°N; 109.7722°W) (Willcox), 29.viii.2017, R. Oram (paratype ♀, RSKM); Phoenix (Maricopa County), 13.x.1997, K.C. Rozen (paratypes 3♂, AMNH); San Simon (Cochise County), 01.ix.1976, R.M. Bohart (paratype ♂, UCBME); SE Willcox (Cochise County), 30.ix.2016, L. Packer (paratype ♀, PCYU); Willcox (Cochise County), 02.ix.2003, J.G. Rozen, J.S. Ascher, R.L. Staff, and R.E. Edwards (paratypes 2♂, AMNH), 22.ix.1984, J.G. Rozen (paratype ♀, AMNH), 26.ix.1980, J.G. Rozen (paratypes 6♀, AMNH), 28.viii.1958, P.D. Hurd (paratype ♂, UCBME), 28–29.viii.1988, K.V. Krombein and B. Norden (paratype ♂, USNM); New Mexico: 5 mi E Laguna (Valencia County), 07.viii.1966, C.R. Kovacic (paratype ♀, UCBME); 20 mi N Animas (Hidalgo County), 05.ix.1981, R.M. Bohart (paratype ♀, UCBME); Mesilla Park, 17.ix.????, T.D. Cockerell (paratype ♀, USNM).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ACR5356. See Type material for specimens examined and sequenced (indicated by unique CCDB-plate and well number).

Epeolus bifasciatus Cresson, 1864

Figs 26, 27, 91A

Epeolus bifasciatus Cresson, 1864a. Proc. Entomol. Soc. Phil. 3: 38 (♂); Cresson, 1916. Mem. Am. Entomol. Soc. 1: 113 (♂) [lectotype designation].

Diagnosis

Unique to E. bifasciatus among North American species of Epeolus are each of the following morphological features: the frontal area bears a pair of granulose protrusions, each located near the upper mesal margin of the compound eye; the pronotal collar is elongate, dilated laterally to about 2 × the medial length in dorsal view; and the dorsum of the metasoma has at most two bright orange-yellow fasciae (usually a basal fascia on T1 and always an apical fascia on T2). Similar species occur in Mexico and Central America, but their occurrence in Canada and the United States has not been confirmed.

Figure 26. 

Epeolus bifasciatus A female, lateral habitus (scale bar 3 mm) B female, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Redescription

This species was recently redescribed (Onuferko 2017).

Distribution

United States, east of the Continental Divide, into central Canada (Fig. 27).

Figure 27. 

Approximate geographic range of E. bifasciatus (orange) based on occurrence records known to the author (yellow circles).

Ecology

See Onuferko (2017) for host and floral records. Floral associations are also indicated in Suppl. material 1, which includes newly discovered associations with Coreopsis tinctoria Nutt. (Compositae) and Verbena hastata L. (Verbenaceae) based on labels of examined voucher specimens.

Discussion

Epeolus bifasciatus is the only species within the “Trophocleptria group” verified as occurring north of Mexico. Originally a genus, Trophocleptria Holmberg was later considered a subgenus of Epeolus (Michener 2000). Although its constituent species seem to form a natural group, a phylogenetic study by Rightmyer (2004) found that maintaining the subgeneric designation rendered Epeolus (Epeolus) paraphyletic, so Michener (2007) treated Trophocleptria as a distinct species group within Epeolus.

Epeolus fumipennis Say has been listed as occurring in Kansas (Snow 1879, in which E.T. Cresson was acknowledged for aiding in identification), but was probably confused with E. bifasciatus, a species that is common in that state (Ascher and Pickering 2017). Brumley (1965) examined specimens at the ANSP and KUNHM from the Midwestern and Southeastern United States labelled as E. fumipennis that according to him were clearly E. bifasciatus. The primary type of E. fumipennis was probably destroyed along with much of Thomas Say’s insect collection (LeConte 1859:v–vi, xix [footnote]), but the medially-narrowed ferruginous pronotal collar and yellow fasciae on T1 and T2 (contrasting with the whitish fasciae on the remaining terga), as well as its occurrence in Mexico, strongly suggest that this species is in the “Trophocleptria group”. However, in E. fumipennis the mesoscutum has distinct paramedian bands, which are absent in E. bifasciatus, and no specimens from Canada or the United States fitting such a description were seen.

Material studied

Type material. Primary: USA: Illinois: (lectotype ♂ [ANSP, catalog number: 2658]).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ADD5310. Specimens examined and sequenced.-Canada: Ontario: 1♀, 1♂ (PCYU).

USA: Florida: 1♂ (FSCA).

Non-barcoded material examined

Canada: Ontario: 5♀, 6♂ (CNC, DEBU, PCYU, ROM); 2 km N Shiloh (43.7400°N; 80.2675°W) (Wellington County), 08.viii.2004, M. Buck (4♀, DEBU); 6 km NW Saint Williams (42.7050°N; 80.4606°W) (Hard.Norfolk Reg., Manestar Tract), 14.vii.2006, S.M. Paiero (5♀, 1♂, DEBU); Rondeau Park (South Point Trail, Kent County), 29.vi.2002, M. Buck (4♀, 1♂, DEBU); Toronto, 04.viii.2005, A. Cosens (1♂, PCYU).

USA: Colorado: Hasty (Bent County), 03.vii.1975, H.E. Evans (1♂, CUM); Longmont (40.1627°N; 105.1441°W) (Boulder County), 17.viii.2012, V. Scott (1♂, CUM); Florida: 2♂ (AMNH, PCYU); Caverns State Park (Jackson County), 16.vi.1999, C. Porter and L. Stange (1♀, FSCA); Lake City (Columbia County), 23.vi.2011, S. Lenberger (1♂, FSCA); Lovers Key State Rec Area (Lee County), 12.v.2008, C. Porter and L. Stange (1♀, FSCA); San Felasco Hammock Preserve State Park (Alachua County), 09–12.v.1979, G.B. Fairchild (1♀, FSCA); St Augustine Beach (St. Johns County), 24.v.1992, F.J. Santana (1♂, FSCA); Georgia: Athens (Whitehall Preserve, Clarke County), 14–19.v.1979, R.H. Turnbow, Jr. (1♂, FSCA); Illinois: 2♀ (AMNH); Iowa: Ames, 18.viii.1934, H.A. Scullen (1♀, CUM); Kansas: Baldwin, vii.????, J.C. Bridwell (1♀, CUM); Maryland: 2♀ (AMNH, BIML); Michigan: 5 km N West Olive (42.9884°N; 86.1423°W) (Ottawa County), 24.viii.2014, J. Gibbs (1♀, JBWM); East Lansing (42.7540°N; 84.4860°W) (Ingham County), 25.viii.2013, J. Gibbs (1♂, JBWM); Near Saline, 26.vi.1954, U.N. Lanham (1♂, CUM); Missouri: Rolla (Phelps County), 26.viii.1962, B. Vogel (2♀, CUM); New York: 1♂ (BIML); North Carolina: 1♂ (AMNH); Ohio: West Jefferson, G. Salt (2♀, NHMUK); Pennsylvania: 1♂ (BIML); South Carolina: 1♂ (DEBU); South Dakota: Oacoma (1 km W Chamberlain, Lyman County), 08.viii.2005, R.E. Wrigley (1♀, JBWM); Texas: Bentsen-Rio Grande Valley State Park, 01–13.vi.1976, C.C. Porter (1♂, FSCA); McAllen Botanical Gardens (McAllen), 03.vi.1976, C.C. Porter (1♂, FSCA); Wisconsin: 1♀ (PCYU).

Epeolus brumleyi sp. n.

Figs 2B, 28, 29, 103B

Epeolus brevicornus Brumley, 1965. M.S. thesis, Utah State University, Logan 38 (♀) [nomen nudum].

Diagnosis

The following morphological features in combination can be used to tell E. brumleyi apart from all other North American Epeolus: the frontal carina is weakly convex, such that the supraclypeal area is barely protuberant in lateral view; the mesoscutum has distinct paramedian bands; the axilla is small to intermediate in size, not extending much beyond the midlength of the mesoscutellum (extending to <2/3 its length) but the free portion is at least 1/4 as long as (and less than 2/5) the entire medial length of the axilla, relatively straight along the medial margin, and ferruginous to some degree whereas the mesoscutellum is typically all black; the fore wing has three submarginal cells; the T1 basal and apical fasciae are subparallel; T2–T4 have complete fasciae; and the T2 fascia has a pair of anterolateral extensions of tomentum that are weakly convergent basally. Epeolus brumleyi most closely resembles E. australis, but in E. australis the frontal carina is strongly convex and the pygidial plate of the male is narrower (the medial length is ~1.5 × the basal width) than in E. brumleyi (the medial length ≈ the basal width).

Description

FEMALE: Length 7.6 mm; head length 1.9 mm; head width 2.7 mm; fore wing length 5.8 mm.

Integument coloration. Mostly black; notable exceptions as follows: partially to entirely ferruginous on mandible, labrum, antenna, pronotal lobe, tegula, axilla, legs, metasomal terga (including pygidial plate), and metasomal sterna. Mandible with apex darker than rest of mandible; preapical tooth slightly lighter than mandibular apex (difficult to see in holotype because mandible closed; described from paratypes). Antenna brown and orange in part. Pronotal lobe and tegula pale ferruginous to amber. Wing membrane subhyaline, apically dusky. Legs more extensively reddish orange than brown or black.

Pubescence. Face with tomentum densest around antennal socket. Clypeus, upper paraocular and frontal areas, and vertexal area mostly exposed. Dorsum of mesosoma and metasoma with bands of off-white to pale yellow short appressed setae. Mesoscutum with paramedian band. Mesopleuron densely hairy, except for two almost entirely bare patches (one beneath base of fore wing (hypoepimeral area), a larger circular patch occupying much of ventrolateral half of mesopleuron). Metanotum with tomentum rubbed off medially in holotype, but uninterrupted and uniformly off white in paratypes. T1 with discal patch elliptical and very wide, the basal and apical fasciae only narrowly joined laterally. T1 with basal fascia complete and apical fascia interrupted medially, T2–T4 with fasciae complete, T2 with fascia with anterolateral extensions of sparser tomentum. T5 with two large patches of pale tomentum lateral to and contacting pseudopygidial area. T5 with pseudopygidial area lunate, its apex more than twice as wide as medial length, indicated by silvery setae on impressed disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs not extending beyond apex of sternum by much more than 1/4 MOD.

Surface sculpture. Punctures dense. Labrum with areas of sparser punctures (i=1–2d) than clypeus (i<1d). Small impunctate shiny spot lateral to lateral ocellus. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula densely punctate mesally (i≤1d), less so laterally (i=1–2d). Mesopleuron with ventrolateral half densely punctate (i≤1d) to rugose, the interspaces shining; mesopleuron with punctures more or less equally dense throughout. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc.

Structure. Preapical tooth blunt and obtuse. Labrum with submedial pair of small denticles, apex edentate. Frontal keel not strongly raised. Scape with greatest length 1.8 × greatest width. F2 as long as wide (L/W ratio = 1.0). Preoccipital ridge not joining hypostomal carina, from which it is separated by no less than 1 MOD at its terminal (difficult to see in holotype; described from paratypes). Mesoscutellum moderately bigibbous. Axilla small to intermediate in size, its lateral margin (L) less than half as long as mesoscutellar width (W) (L/W ratio = 0.4) and tip not extending beyond midlength of mesoscutellum; axilla with tip visible, but unattached to mesoscutellum for less than 2/5 the medial length of axilla; axilla with lateral margin relatively straight and without carina. Fore wing with three submarginal cells. Pygidial plate apically truncate.

MALE: Description as for female except for usual secondary sexual characters and as follows: F2 shorter, nearly as long as wide (L/W ratio = 0.9); S4 and S5 with much longer coppery to silvery subapical hairs; pygidial plate apically rounded, with large deep punctures closely clustered.

Figure 28. 

Epeolus brumleyi A female paratype, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male paratype, lateral habitus (scale bar 3 mm), and D female paratype axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Etymology

This species is named after its discoverer, Richard L. Brumley, who recognized it and five other Epeolus formally described here (E. axillaris, E. chamaesarachae, E. diadematus, E. splendidus, and E. tessieris) as new species.

Distribution

Arizona to Texas and presumably Mexico, given the close proximity of some collection localities (e.g., Douglas, Arizona) to the Mexico–United States border (Fig. 29).

Figure 29. 

Approximate geographic range of E. brumleyi (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: Four representatives of this species were collected at a single site in southeast Arizona in the spring of 2016 (see Material studied), from or flying near patches of Chamaesaracha (A. Gray) Benth. (Solanaceae), which were visited by large numbers of Colletes (presumably the host species). Using Stephen’s (1954) key, collected females were identified as C. scopiventer Swenk (a species known only from females) whereas males were identified (based in part on examination of the terminalia, which were excised) as C. wickhami Timberlake (a species known only from males), and sequenced specimens of both sexes were assigned the same BIN (BOLD:AAJ7578).

FLORAL RECORDS: Labels of examined voucher specimens indicate floral associations with Chamaesaracha coniodes (Moric. ex Dunal) Britton and Physalis L. (Solanaceae).

Discussion

Epeolus brumleyi is a southwestern species that exhibits very little intraspecific morphological variation. Adults have been collected in every month from March to September, and barcoded specimens collected in early May, June, and late August were assigned the same BIN.

Material studied

Type material. Primary: USA: Texas: Davis Mountains, 10.vii.1942, E.C. Van Dyke (holotype ♀, CAS).

Secondary: USA: Arizona: 1 mi E Douglas (Cochise County), 08.v.1989, J.G. Rozen (paratype ♀ [CCDB-28315 G10], AMNH); 14 mi SW Apache (Cochise County), 14.v.1988, J.G. Rozen (paratype ♀, AMNH); 3 mi NE Portal (Cochise County), 18.viii.1970, J.G. Rozen (paratype ♂, AMNH); 3–7 mi S San Simon (Cochise County), 21.v.1988, J.G. Rozen (paratype ♀, AMNH); 9 mi E Douglas (Cochise County), 17.ix.1976, J.G. Rozen (paratype ♂, AMNH); Hwy 80 (31.4450°N; 109.4722°W) (~8 mi NE Douglas, Cochise County), 10.v.2016, T.M. Onuferko (allotype ♂, PCYU), 10.v.2016, T.M. Onuferko (paratypes 2♀ (1 barcoded [CCDB-24580 B11]), 1♂, PCYU); S Blue Sky Road (4 mi E Willcox, Cochise County), 30.viii.2015, J.S. Francis (paratype ♂ [CCDB-28238 A04], PCYU); New Mexico: 0.7 km E Longview Spring (32.1007°N; 104.6137°W) (Eddy County), 22.vi.2010, A. Druk and J.D. Herndon (paratype ♀, BBSL); 1 mi W Animas (Hidalgo County), 30.viii.1977, R.W. Brooks (paratype ♀, KUNHM); 1.1 km SW by W Oak Spring (32.1743°N; 104.4580°W) (Eddy County), 11.viii.2010, J.D. Herndon (paratype ♀, BBSL); 4 mi S Animas (Hidalgo County), 24.viii.1974, Rozen and Favreau (paratype ♂, AMNH); Loving (Eddy County), 28.v.1945, J.W. MacSwain (paratype ♂, BBSL); Walnut Canyon (32.1872°N; 104.3936°W) (2.6 km SE by S Cottonwood Spring, Eddy County), 03.vi.2010, A. Druk and J.D. Herndon (paratype ♀, BBSL); Texas: 18 km N Coleman (Coleman County), 01.vi.1989, B.N. Danforth (paratype ♀ [CCDB-28315 C09], KUNHM); 2 mi S Falfurrias (Brooks County), 13.iii.1999, J.L. Neff, A. Hook, and C. R. Riley (paratype ♂, CTMI); Davis Mountains, 28.vi.1942, E.C. Van Dyke (paratype ♂, BBSL), 17.iv.1954, R.H. Beamer (paratype ♂, BBSL); Sarita (Kenedy County), 15.iv.1976, J.E. Gillaspy (paratype ♀, BBSL).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ACZ9234. See Type material for specimens examined and sequenced (indicated by unique CCDB-plate and well number).

Epeolus canadensis Mitchell, 1962

Figs 30, 31, 102B

Epeolus canadensis Mitchell, 1962. N. C. Agric. Exp. Stn. Tech. Bull. 152: 444 (♀).

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. canadensis apart from all other North American Epeolus except E. compactus and E. ferrarii: in females, F2 is at least 1.2 × as long as wide; the mesoscutum has a small anteromedial patch of pale tomentum; the axilla is small to intermediate in size, not extending much beyond the midlength of the mesoscutellum (extending to <2/3 its length) but the free portion is more than 1/4 as long as the entire medial length of the axilla, and the axilla (except sometimes the tip) and mesoscutellum are black; the mesopleuron is closely (most i<1d) and evenly punctate; and the T2 fascia lacks lobe-like anterolateral extensions of tomentum, although it may be broader laterally. Epeolus canadensis differs from E. compactus and E. ferrarii in the shape of the T1 discal patch, which in E. canadensis is distinctly triangular or semicircular (the basal fascia is conspicuously arched and fully continuous with the longitudinal band) and its medial longitudinal extent is more than 1/3 the lateral extent. In E. compactus and E. ferrarii the shape of the T1 discal patch is variable but typically quadrangular with the basal and apical fasciae subparallel and separated by a distinct longitudinal band. In E. compactus, the medially-interrupted T1 basal and apical fasciae may be so broad laterally that they are joined, resulting in a diamond shape with concave sides. In E. ferrarii the discal patch may be trapezoidal or almost semicircular, but if at all semicircular its medial longitudinal extent is at most 1/3 the lateral extent and the basal fascia and longitudinal band are at least joined at somewhat of an angle.

Figure 30. 

Epeolus canadensis A female, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Redescription

This species was recently redescribed (Onuferko 2017).

Distribution

Atlantic Canada to southwestern United States (Fig. 31).

Figure 31. 

Approximate geographic range of E. canadensis (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: An association between Colletes kincaidii Cockerell and E. canadensis hypothesized earlier (Onuferko 2017) seems more likely now based on new knowledge that the two species have been collected in co-occurrence near Six Mile Creek (Ithaca), New York, USA (J. Ascher, personal communication, 2017) and personal collections of the two species in early July, 2017 on the side of a road in Navan (east of Ottawa), Ontario, Canada. Colletes kincaidii females and males were collected from staghorn sumac (Rhus typhina L. (Anacardiaceae)) on the same dates E. canadensis were collected from daisy-like flowers (Compositae) closer to the ground.

FLORAL RECORDS: See Onuferko (2017) for floral records. Floral associations are also indicated in Suppl. material 1, which includes a newly discovered association with Grindelia Willd. (Compositae) based on the label of one examined voucher specimen.

Discussion

Detailed morphological and taxonomic remarks about this species are given in Onuferko (2017).

Material studied

Type material. Primary: Canada: Nova Scotia: Ingonish (Cape Breton Island), 07.viii.1928, G. Fairchild (holotype ♀ [MCZ, catalog number: 32859]).

Secondary: USA: New York: 9-Mile Creek (Ithaca), 10.vii.1937, P.P. Babiy (allotype ♂ [CUIC, catalog number: 00015611]).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ADA0845. Specimens examined and sequenced.–Canada: Ontario: 1♀, 1♂ (DEBU); Navan (45.3982°N; 75.3623°W) (Caroltodd Dr & Whispering Willow Dr), 02.vii.2017, T.M. Onuferko (1♂, PCYU), 03.vii.2017, T.M. Onuferko (1♀, PCYU).

USA: Arizona: 1♂ (PCYU); Flagstaff (35.1737°N; 111.6756°W) (Coconino County), 01–03.vi.2017, T.M. Onuferko (1♀, PCYU); New Mexico: 2♂ (DEBU, PCYU).

Non-barcoded material examined

Canada: Nova Scotia: 3♀, 4♂ (CNC); Ontario: 10♀, 15♂ (CNC, DEBU, PCYU, ROM); Forks of the Credit Provincial Park, vii.2002?, J. Grixti (1♂, PCYU); Prince Edward Island: 1♀ (CNC); Quebec: 3♀ (CNC).

USA: Arizona: 5♀, 3♂ (AMNH, CNC, PCYU); Flagstaff (35.1737°N; 111.6756°W) (Coconino County), 01–03.vi.2017, T.M. Onuferko (1♀, PCYU); Huachuca Mountains, 14.ix.1938, R.H. Crandall (1♀, 1♂, LACM); Santa Catalina Mountains (Pima County), J.L. Neff (1♂, LACM); Arkansas: 1♀ (FSCA); Colorado: Boulder (Boulder County), 12.ix.1965, U.N. Lanham (1♀, CUM); Illinois: 1♀ (KUNHM); Kansas: 2♀ (KUNHM); Missouri: 1♀ (KUNHM); New Mexico: 5♀, 5♂ (AMNH, BBSL, CNC).

Epeolus carolinus Mitchell, 1962

Figs 3C, 32, 33, 92B

Epeolus carolinus Mitchell, 1962. N. C. Agric. Exp. Stn. Tech. Bull. 152: 445 (♂).

Diagnosis

The following morphological features in combination can be used to tell E. carolinus apart from all other North American Epeolus: the mandible has a blunt, obtuse preapical tooth; the axilla is elongate, extending well beyond the midlength of the mesoscutellum but not beyond its posterior margin, and the free portion is distinctly hooked; the mesopleuron is closely (most i<1d) and evenly punctate; and the metasomal fasciae are yellow to orange and interrupted medially. Epeolus carolinus resembles E. deyrupi in general appearance, but in E. deyrupi the axilla is larger, extending as far back as or beyond the posterior margin of the mesoscutellum, and dilated laterally but relatively straight along the medial margin, and the mesopleuron commonly has sparser punctures ventrolaterally (i≤2d) than that of E. carolinus, with the interspaces shining or somewhat dull due to tessellate surface microsculpture.

Redescription

MALE: Length 6.5 mm; head length 1.8 mm; head width 2.4 mm; fore wing length 5.7 mm.

Integument coloration. Mostly black; notable exceptions as follows: partially to entirely ferruginous on mandible, antenna, pronotal lobe, tegula, axilla, mesoscutum, mesoscutellum, legs, and pygidial plate. Mandible with apex darker than rest of mandible; preapical tooth slightly lighter than mandibular apex (difficult to see in holotype; described from paratype). Antenna brown except scape, pedicel, and F1 extensively orange. Pronotal lobe and tegula pale ferruginous to amber. Mesoscutum with orange spot anterolaterally between pronotal lobe and tegula. Wing membrane subhyaline, apically dusky. Legs more extensively reddish orange than brown or black.

Pubescence. Face with tomentum densest around antennal socket. Tomentum slightly sparser on clypeus; upper paraocular and frontal areas, and vertexal area mostly exposed. Dorsum of mesosoma and metasoma with bands of off-white and yellow short appressed setae. Mesoscutum with paramedian band. Mesopleuron densely hairy, except for two sparsely hairy circular patches (one behind pronotal lobe, a larger one occupying much of ventrolateral half of mesopleuron). Metanotum with tomentum sparser medially, uniformly off white. T1 with discal patch quadrangular and very wide, the basal and apical fasciae only narrowly joined laterally by few sparsely scattered pale hairs (not joined in paratype and multiple non-type specimens). T1–T5 with apical fasciae interrupted medially, those of T2–T4 somewhat broader laterally, T2 with fascia without anterolateral extensions of tomentum. T6 with fascia complete. S4 and S5 with long coppery to silvery subapical hairs.

Surface sculpture. Punctures dense. Labrum with larger punctures than clypeus, but punctures of both equally dense (i<1d). Impunctate spot lateral to lateral ocellus absent. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula very densely punctate mesally (i<1d), much less so laterally (i>2d). Mesopleuron with ventrolateral half densely punctate (i<1d) to rugose; mesopleuron with punctures more or less equally dense throughout. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc.

Structure. Preapical tooth inconspicuous, blunt and obtuse. Labrum with pair of small subapical denticles not preceded by carinae. Frontal keel not strongly raised. Scape with greatest length 1.8 × greatest width. F2 noticeably longer than wide (L/W ratio = 1.4). Preoccipital ridge not joining hypostomal carina, from which it is separated by less than 1 MOD at its terminal (difficult to see in holotype; described from non-type specimens). Mesoscutellum weakly bigibbous. Axilla large, its lateral margin (L) more than half as long as mesoscutellar width (W) (L/W ratio = 0.6) and tip extending well beyond midlength of mesoscutellum but not as far back as its posterior margin; axilla with tip conspicuously diverging from side of mesoscutellum, distinctly hooked, and axilla with free portion 2/5 its medial length; axilla with lateral margin arcuate and carinate. Fore wing with three submarginal cells. Pygidial plate apically rounded, with large deep punctures more or less evenly spaced throughout, with the interspaces shining.

FEMALE: Description as for male except for usual secondary sexual characters and as follows: F2 even longer than wide (L/W ratio = 1.7); T5 with pseudopygidial area lunate, its apex more than twice as wide as medial length, indicated by silvery setae on flat disc of apicomedial region elevated from rest of tergum; S4 and S5 with much shorter hairs (S5 with apical fimbria of coppery to silvery hairs not extending beyond apex of sternum by more than 1/4 MOD); pygidial plate apically truncate, with small, denser punctures.

Figure 32. 

Epeolus carolinus A female, lateral habitus (scale bar 3 mm) B female, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Distribution

South Atlantic states (Fig. 33).

Figure 33. 

Approximate geographic range of E. carolinus (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: The host species of E. carolinus is/are presently unknown.

FLORAL RECORDS: Mitchell (1962) indicated a floral association with Eupatorium L. (Compositae), and BugGuide (http://www.bugguide.net/) indicates an association with Solidago fistulosa Mill. Labels of examined voucher specimens further indicate associations with Euthamia graminifolia (L.) Nutt. (Compositae), Heterotheca subaxillaris (Lam.) Britton & Rusby (Compositae), and Spermacoce L. (Rubiaceae).

Discussion

This southeastern species is quite variable in terms of integument coloration and pubescence on the metasomal terga. The mesoscutellum and disc of T1 range from entirely black to entirely ferruginous. The axillae appear to be at least partially ferruginous. Whereas T1 and T2 have prominent yellow fasciae, the fasciae on the remaining terga range from prominent to reduced or even absent. Adults of Epeolus carolinus are active in September and October.

Material studied

Type material. Primary: USA: North Carolina: Kill Devil Hills, 12.ix.1956, T.B. Mitchell (holotype ♂ [USNM, catalog number: 534042]).

Secondary: USA: North Carolina: Kill Devil Hills, 13.ix.1956, T.B. Mitchell (paratype ♂, NHMUK); New River, 20–30.ix.1944, G.E. Bohart (paratype ♂, BBSL).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ACM5698. Specimens examined and sequenced.–USA: Florida: Timucuan Ecological & Historic Preserve (30.3842°N; 81.4857°W) (Duval County), 15.x.2012, C. Pontifet (1♂, BIML); South Carolina: Prince George Estates (E Hwy 17, Georgetown County), 09.x.2006, S. Paiero and S.A. Marshall (1♂, DEBU).

Non-barcoded material examined

USA: Florida: Archbold Biological Station (Highlands County), 11.x.1978, H.V. Weems, Jr. and S.J. Chance (2♀, LACM), 08.x.1964, P.H. Arnaud, Jr. (1♂, LACM); Cedar Key (Levy County), 27.x.1974, E.E. Grissell (3♀, 1♂, UCBME); Doyle Conner Bldg (Gainesville, Alachua County), 04.x.1995, C. Porter (1♂, FSCA), 12.x.1995, C. Porter (1♂, FSCA), 17.x.1995, C. Porter (2♂, FSCA); Gainesville (Alachua County), 13.x.??48 (1♀, LACM), 25.viii.1976, W.H. Pierce (1♂, UCBME); Mason Road (Melrose, Putnam County), 11.x.2009, J.S. Ascher and H.G. Hall (1♂, AMNH); Perry (Taylor County), 1983, L. Packer (1♀, PCYU); W Murdock (Charlotte County), 20.x.1983, L. Packer (2♀, 2♂, PCYU); South Carolina: Aiken Savannah River Site (33.3449°N; 81.6614°W), 17.x.2016, S. Breland (1♀, JBWM); Prince George Estates (E Hwy 17, Georgetown County), 09.x.2006, S. Paiero and S.A. Marshall (1♂, DEBU).

Epeolus chamaesarachae sp. n.

Figs 1, 34, 35, 91C, 92I

Epeolus lobus Brumley, 1965. M.S. thesis, Utah State University, Logan 51 (♀) [nomen nudum].

Diagnosis

Epeolus chamaesarachae does not closely resemble any other species of Epeolus except E. diadematus. Unique in the genus to both species are each of the following morphological features: the vertexal area has two pairs of shiny (usually impunctate) protrusions, the mesoscutum is distinctly ornamented with mostly separate patches of (but some intermixed) pale and ferruginous tomentum, and the T2 fascia has two pairs of anterolateral extensions of tomentum. The difference is that in E. chamaesarachae the mesopleuron has sparser punctures ventrolaterally (most i>1d) whereas in E. diadematus the mesopleuron has denser (most i≤1d) and more numerous punctures ventrolaterally.

Description

FEMALE: Length 7.0 mm; head length 2.0 mm; head width 2.6 mm; fore wing length 5.7 mm.

Integument coloration. Mostly black; notable exceptions as follows: partially to entirely ferruginous on mandible, antenna, pronotal collar, pronotal lobe, tegula, axilla, mesoscutellum, and legs. Mandible with apex darker than all but extreme base; preapical tooth lighter than mandibular apex (difficult to see in holotype; described from paratypes). Antenna dark brown except scape, pedicel, and F1 brownish orange in part. Pronotal lobe and tegula pale ferruginous to amber. Wing membrane subhyaline, apically dusky. Legs more extensively reddish orange than brown or black.

Pubescence. Face with tomentum densest around antennal socket. Vertexal area with tomentum mostly ferruginous. Dorsum of mesosoma with bands of off-white and ferruginous short appressed setae. Dorsum of metasoma with bands of off-white to pale yellow short appressed setae. Pronotal lobe entirely obscured by pale tomentum. Pronotal collar with tomentum black medially, pale and ferruginous laterally. Mesoscutum with paramedian band of pale tomentum; ferruginous and pale tomentum encircling black spots medially and laterally, respectively, on anterior margin; and ferruginous tomentum along medial mesoscutal line and parapsidal line. Mesopleuron with upper half densely hairy, although scrobe visible; ventrolateral half nearly bare. Metanotum with tomentum uninterrupted, off white laterally and black medially. T1 with median diamond-shaped black discal patch enclosed by pale tomentum, except for medial separation at apex. T1 with apical fascia with black spot posterolaterally. T2–T4 with fasciae interrupted medially, T2 with fascia with paired anterolateral extensions of tomentum. T3 and T4 with fasciae interrupted laterally, with medial portion on apical margin and lateral portion encircling black tomentum on apical margin. T5 with two large patches of pale tomentum lateral to and separate from pseudopygidial area. T5 with pseudopygidial area lunate, its apex more than twice as wide as medial length, indicated by silvery setae on disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs extending beyond apex of sternum by ~1/3 MOD.

Surface sculpture. Punctures dense, but those of head and mesosoma sparser in some areas, larger, deep, and distinct. Labrum with larger and sparser punctures (i=1–2d) than clypeus (i≤1d). Upper paraocular area and vertexal area with few punctures, the interspaces shining. Mesoscutum, mesoscutellum, and axilla coarsely and densely to sparsely punctate; the interspaces shining. Tegula densely punctate mesally (i≤1d), less so laterally (i=1–2d). Mesopleuron with denser (i≤1d) punctures in upper half than ventrolateral half, and ventrolateral half with most interspaces large (i>1d); the interspaces shining. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc.

Structure. Labral apex with pair of small denticles (preceded by submedial pair of small denticles) separated by shallow concavity and between second pair of apical lobes. Frontal keel strongly raised. Vertexal area with two pairs of impunctate shiny protrusions. Scape with greatest length 1.6 × greatest width. F2 as long as wide (L/W ratio = 1.0). Preoccipital ridge not joining hypostomal carina, from which it is separated by about 1.5 MOD at its terminal. Mesoscutellum strongly bigibbous. Axilla intermediate in size, its lateral margin (L) nearly half as long as mesoscutellar width (W) (L/W ratio = 0.4–0.5) and tip not extending beyond midlength of mesoscutellum; axilla with tip conspicuously diverging from side of mesoscutellum, distinctly hooked, but unattached to mesoscutellum for less than 1/3 the medial length of axilla; axilla with lateral margin relatively straight and without carina. Fore wing with three submarginal cells. Pygidial plate apically truncate.

MALE: Description as for female except for usual secondary sexual characters and as follows: F2 shorter, nearly as long as wide (L/W ratio = 0.8); S4 and S5 with much longer coppery to silvery subapical hairs; pygidial plate apically rounded, with large deep punctures closely clustered apically and sparser basally, with the interspaces shining.

Figure 34. 

Epeolus chamaesarachae A female paratype, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male paratype, lateral habitus (scale bar 3 mm), and D female paratype axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Etymology

The name is in reference to the genus of flowers (Chamaesaracha) on which the holotype was collected.

Distribution

Northwestern Mexico and southwestern United States (Fig. 35).

Figure 35. 

Approximate geographic range of E. chamaesarachae (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: The female PCYU paratype collected by H.T. Ngo (see Material studied) is labelled with the same collection information as three Colletes specimens (2♀, 1♂) of the presumed host species, which were barcoded and all share the same BIN (BOLD:AAJ7578). Using Stephen’s (1954) key, the two females were identified as C. scopiventer (a species known only from females) whereas the male was identified (based in part on examination of the terminalia, which were excised) as C. wickhami (a species known only from males).

FLORAL RECORDS: Labels of examined voucher specimens indicate floral associations with Baccharis L. (Compositae), Chamaesaracha, Kallstroemia grandiflora Torr. ex A. Gray (Zygophyllaceae), Margaranthus solanaceous Schltdl. (Solanaceae), Sphaeralcea angustifolia (Cav.) G. Don, and Tidestromia lanuginosa (Nutt.) Standl. (Amaranthaceae).

Discussion

This species and the very similar E. diadematus are unusual among Epeolus in that the vertexal area has two pairs of shiny (usually impunctate) protrusions, and dorsally the mesosoma and metasoma have unique patterns of ferruginous (mesosoma only) and off-white to pale yellow short appressed setae. Epeolus chamaesarachae occurs in the Southwestern United States, and its flight season, based on material examined, is late summer.

Material studied

Type material. Primary: USA: Arizona: Douglas Model Plane Airport (31.3433°N; 109.4980°W) (Cochise County), 24.viii.2010, T.L. Griswold (holotype ♀ [CCDB-28239 F07], BBSL).

Secondary: Mexico: Durango: Durango, 14.viii.1947, D. Rockefeller Exp. Michener (paratype ♂, AMNH); San Juan del Río, 30.vii.1947, D. Rockefeller Exp. Michener (paratype ♀, AMNH).

USA: Arizona: 1 mi E Douglas (Cochise County), 16.viii.1962, M. Statham (paratype ♂, AMNH), 27.viii.2007, H.T. Ngo (paratype ♀ [CCDB-22013 G05], PCYU); 1 mi E Douglas (31.3356°N; 109.4950°W) (Cochise County), 23.viii.2003, J.G. Rozen (paratype ♀, AMNH); 12 mi NW Douglas (Cochise County), 30.viii.1989, J.G. and B.L. Rozen and R. Foster (paratype ♀, AMNH); 14 mi SW Apache (Cochise County), 04.viii.1961, J.G. Rozen (paratype ♀, AMNH), 21.viii.2008, J.S. Ascher, J.G. Rozen, and M.A. Rozen (paratype ♂ [CCDB-22791 A09], AMNH); 25 mi SE Sanders (Apache County), 14.viii.1972, J.G. Rozen and R. McGinley (paratype ♂, AMNH); 8 mi NE Portal (Cochise County), 14.viii.1990, J.G. Rozen and J. Krieger (paratype ♀, AMNH); Douglas Model Plane Airport (31.3433°N; 109.4980°W) (Cochise County), 24.viii.2010, T.L. Griswold (allotype ♂ [CCDB-28239 F09], BBSL), 24.viii.2010, T.L. Griswold (paratype ♂, BBSL); Geronimo Trail at Sycamore Creek (31.4432°N; 109.1390°W) (Cochise County), 28.viii.2016, L. Packer (paratype ♀, PCYU); Tombstone (Cochise County), 17.viii.1975, J.G. Rozen (paratype ♀, AMNH); New Mexico: 16 mi S Animas (31.7211°N; 108.8224°W) (Hidalgo County), 03.ix.2011, J.G. Rozen and E.S. Wyman (paratype ♀ [CCDB-22791 A07], AMNH); 2.6 mi E Animas (31.9542°N; 108.7630°W) (NM Hwy 9, 2.6 mi E NM Hwy 338), 11.viii.1972, T.J. Zavortink (paratypes 1♀, 2♂, UCBME); 5.5 mi E Animas (31.9558°N; 108.7142°W) (Hidalgo County), 18–25.viii.2002, E. Elle (paratype ♂, AMNH).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ACP9403. See Type material for specimens examined and sequenced (indicated by unique CCDB-plate and well number).

Epeolus compactus Cresson, 1878

Figs 3F, 36, 37, 38

Epeolus compactus Cresson, 1878. Trans. Am. Entomol. Soc. 7: 89 (♀, ♂); Cresson, 1916. Mem. Am. Entomol. Soc. 1: 115 (♀) [lectotype designation].

Epeolus crucis Cockerell, 1904. Ann. Mag. Nat. Hist. 13: 39 (♀), syn. n.

Epeolus hitei Cockerell, 1908. Entomologist 41: 60 (♀).

Triepeolus gabrielis Cockerell, 1909. Ann. Mag. Nat. Hist. 5: 26 (♂).

Epeolus geminatus Cockerell and Sandhouse, 1924. Proc. Calif. Acad. Sci. (4) 13: 315 (♀).

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. compactus apart from all other North American Epeolus except E. canadensis and E. ferrarii: in females, F2 is at least 1.2 × as long as wide; the mesoscutum has a small anteromedial patch of pale tomentum; the axilla is small to intermediate in size, not extending much beyond the midlength of the mesoscutellum (extending to <2/3 its length) but the free portion is more than 1/4 as long as the entire medial length of the axilla, and the axilla (except sometimes the tip) and mesoscutellum are black; the mesopleuron is closely (most i<1d) and evenly punctate; and the T2 fascia lacks lobe-like anterolateral extensions of tomentum, although it may be broader laterally. Epeolus compactus is most similar to E. ferrarii, and in both species the T1 discal patch is typically quadrangular with the basal and apical fasciae subparallel and separated by a distinct longitudinal band, but in E. ferrarii the T2–T4 fasciae are not broadened medially into rounded lobes (as in E. compactus) but evenly broad or tapering until separated medially. Epeolus canadensis differs from both species in that the T1 discal patch is distinctly triangular or semicircular (the basal fascia is conspicuously arched and fully continuous with the longitudinal band) and its medial longitudinal extent is more than 1/3 the lateral extent. In E. compactus, the medially-interrupted T1 basal and apical fasciae may be so broad laterally that they are joined, resulting in a diamond shape but with concave sides; in E. canadensis the lateral sides are straight or convex.

Figure 36. 

Epeolus compactus A female, lateral habitus (scale bar 3 mm) B female lectotype, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Redescription

This species was recently redescribed (Onuferko 2017).

Distribution

Western North America (Fig. 37).

Figure 37. 

Approximate geographic range of E. compactus (orange) based on occurrence records known to the author (yellow circles).

Ecology

See Onuferko (2017) for host and floral records. Floral associations are also indicated in Suppl. material 1.

Discussion

Epeolus compactus is a commonly collected species, widespread in Western North America. It is most similar to E. canadensis and E. ferrarii. In the original description of E. crucis Cockerell, the holotype was said to have been initially identified as E. compactus by W.J. Fox, but Cockerell (1904) considered it to be distinct, mainly because of differences in coloration and pubescence. The specimen (unusually) has abundant pale tomentum on the discs of the metasomal terga (Fig. 38A), but representatives of several species (e.g., E. ainsliei, E. minimus, and E. novomexicanus) exhibiting atypical abundance of pale tomentum on the mesosoma and metasoma were also observed. Despite the presence of pale tomentum, the discal patch is quadrangular/diamond-shaped (Fig. 38A) as is typical for E. compactus (Fig. 38B), and the fascia of T2 is separated medially into rounded lobes. In the E. crucis holotype, the axillae and mesoscutellum are (unusually) ferruginous, but it is not unprecedented for species of the genus to have representatives displaying atypical integument coloration. Interestingly, Brumley (1965) treated E. crucis as distinct, but the features listed as unique for that species are evident only in the holotype of E. rufulus. In fact, his key does not work for the holotypes of E. crucis and E. novomexicanus, which Brumley believed to be the same species. Unlike in E. rufulus, in the E. crucis holotype the axillae do not extend beyond the midlength of the mesoscutellum, and the axilla is not conspicuously diverging from the side of the mesoscutellum – the free portion is less than 1/3 as long as the entire medial length of the axilla. As a result of Brumley’s work, specimens of what are actually E. rufulus housed at various entomological institutions have been identified (or rather misidentified) as E. crucis.

Figure 38. 

A E. crucis female holotype (herein synonymized under E. compactus), dorsal habitus, and B E. compactus typical female, dorsal habitus, in which the axilla, mesoscutellum, and discs of the metasomal terga (in terms of integument coloration and pubescence) are black or nearly black. Scale bars 3 mm.

Material studied

Type material. Primary: USA: California: Mill Creek Canyon (San Bernardino County), 12.ix.1923, E.P. Van Duzee (E. geminatus holotype ♀ [CAS, catalog number: 01610]); San Gabriel Mountains (near Pasadena), 15.vii.1909, F. Grinnell, Jr. (T. gabrielis holotype ♂ [USNM, catalog number: 534044]); Colorado: Copeland Park (Boulder County), 06.ix.1907, G.M. Hite (E. hitei holotype ♀ [USNM, catalog number: 534045]); New Mexico: Las Cruces, C.H. Townsend (E. crucis holotype ♀ [USNM, catalog number: 534043]); Texas: G.W. Belfrage (E. compactus lectotype ♀ [ANSP, catalog number: 2227]).

Secondary: USA: Colorado: (E. compactus paralectotype ♀, AMNH).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ACU6228. Specimens examined and sequenced.–Canada: Manitoba: Birds Hill Provincial Park (50.0114°N; 96.9028°W) (Division 12), 15.vii.2017, J. Gibbs (1♂, JBWM).

USA: California: 1♀ (PCYU); Oregon: 3♂ (PCYU); Washington: 1♀ (PCYU).

Non-barcoded material examined

Canada: Alberta: 1♀ (KUNHM); British Columbia: 2♀, 1♂ (CNC); McIntyre Road (Oliver), 29.v.1958, H. and A. Howden (1♂, CNC); Saskatchewan: 1♂ (CNC).

Mexico: Baja California: 1 mi W San Borja, 12–13.vi.1967, E.L. Sleeper and E.M. Fisher (1♀, LACM); Baja California Sur: 6 km E Insurgentes, 24.iv.1975, E.M. Fisher (1♀, LACM); La Paz and vicinity, 11–14.vi.1975, H. Evans, W. Rubink, and D. Gwynne (1♀, CUM); Durango: Durango, 13.viii.1962, A.E. Michelbacher (1♀, EMEC); Sonora: 16 mi NW Puerto Peñasco, 29.iii.1965, C.J. McCoy (1♂, CUM).

USA: Arizona: 2♀, 1♂ (AMNH, PCYU); 15 mi S Bullhead City (Mohave County), 07.iv.1977, L. Bezark (1♀, UCBME); Oak Creek Valley Road (Yavapai County), 16.vi.1978, R.C. Miller (1♀, UCBME); California: 1♀, 3♂ (AMNH, FSCA); Andreas Canyon (Riverside County), 30.iii.1977, R.M. Bohart (1♂, UCBME); Arroyo Seco Campground (Monterey County), 19.v.1964, F.D. Parker (1♀, UCBME); 19.v.1964, R.M. Bohart (1♂, UCBME), 23.vii.1967, R.F. Denno (1♀, UCBME); Charlton Flats (San Gabriel Mountains), 08.ix.1977, A.S. Menke (1♀, UCBME); Felton Springs (Santa Cruz County), 16.vi.1973, R.M. Bohart (1♂, UCBME); Granite Mountains (San Bernardino County), 10.x.1977, N.J. Smith (1♀, UCBME), 10.x.1977, R.M. Bohart (1♀, UCBME); Mojave (Kern County), 23.v.1978, R.P. Meyer (2♂, UCBME); Peña Spring (San Diego County) (1♀, BBSL); Thousand Palms (Riverside County), 11.iv.1970, E.E. Grissell (1♀, UCBME); Colorado: 3♀ (AMNH, PCYU); Nevada: Kings Canyon (5 mi W Carson City), 07.viii.1975, B. Villegas (1♂, UCBME); New Mexico: 8♂ (AMNH, PCYU); Granite Gap (18 mi N Rodeo, Hidalgo County), 07.ix.1976, R.M. Bohart (1♀, UCBME); Oklahoma: 1♀ (FSCA); Oregon: 1♂ (PCYU); Texas: 7.6 mi S Van Horn (Culberson County), 27.iv.1979, R.R. Snelling (1♀, LACM); Rd 1108 (4–8 mi SE 652, Culberson County), 14.vi.2005, J.L. Neff and A. Hook (1♂, CTMI); Z H Canyon (30.0920°N; 104.6620°W) (Presidio County), 19.v.2005, J.L. Neff and A. Hook (1♀, CTMI); Washington: 1♀ (PCYU); Wyoming: 1♀, 2♂ (AMNH).

Epeolus deyrupi sp. n.

Figs 39, 40, 92C

Diagnosis

The following morphological features in combination can be used to tell E. deyrupi apart from all other North American Epeolus: the axilla is large, with the tip extending well beyond the midlength of the mesoscutellum, dilated laterally, and like the mesoscutellum ferruginous; the mesopleuron commonly has sparser punctures ventrolaterally (i≤2d) than in upper half, with the interspaces shining or somewhat dull due to tessellate surface microsculpture; and the T1–T3 apical fasciae are interrupted and (to varying degrees) brownish orange medially and off white laterally. Epeolus deyrupi resembles E. andriyi, E. floridensis, E. howardi, and E. packeri in that the axilla is large, with the lateral margin arcuate, and like the mesoscutellum ferruginous, and that the T1–T3 apical fasciae are interrupted medially. However, in E. deyrupi the pseudopygidial area of the female is wider (the apex >2 × the medial length) than in E. andriyi, E. floridensis, or E. howardi (the apex <2 × the medial length), and the T1 basal fascia is absent or reduced to a pair of small patches of pale tomentum whereas in E. andriyi, E. floridensis, and E. howardi T1 has a distinct, although often medially-interrupted, basal fascia. Epeolus deyrupi closely resembles E. packeri, but in E. packeri the mesopleuron has denser punctures ventrolaterally (most i<1d) than that of E. deyrupi and the metasomal terga have pale but not brownish orange pubescence.

Description

FEMALE: Length 8.8 mm; head length 2.2 mm; head width 2.9 mm; fore wing length 6.1 mm.

Integument coloration. Mostly black; notable exceptions as follows: at least partially ferruginous on mandible, labrum, clypeus, antenna, pronotal collar, pronotal lobe, tegula, axilla, mesoscutum, mesoscutellum, metanotum, mesopleuron, metapleuron, propodeum, and legs. Mandible with apex darker than rest of mandible; preapical tooth lighter than mandibular apex (difficult to see in holotype because mandible closed; described from paratypes). Antenna brown and orange in part. Pronotal lobe and tegula pale ferruginous to amber. Mesoscutum reddish brown laterally and posteriorly. Wing membrane subhyaline, apically dusky. Legs more extensively reddish orange than brown or black.

Pubescence. Face with tomentum densest around antennal socket. Dorsum of mesosoma and metasoma with bands of off-white and brownish orange short appressed setae. Mesoscutum with paramedian band. Mesopleuron mostly bare, but tomentum moderately dense ventrally as well as between two almost entirely bare patches (one beneath base of fore wing (hypoepimeral area), a larger circular patch occupying much of ventrolateral half of mesopleuron). Metanotum with tomentum uninterrupted except for median bare patch in posterior half (also bare along posterior margin), uniformly off white. T1 with basal fascia reduced to pair of small patches of off-white tomentum; T1–T4 with apical fasciae brownish orange medially and off white laterally, and medially interrupted and removed from apical margin; T2 with fascia without anterolateral extensions of tomentum. T4 with fascia interrupted laterally. T5 with two patches of pale tomentum bordering and separate from pseudopygidial area. T5 with pseudopygidial area lunate, its apex more than twice as wide as medial length, indicated by silvery setae on disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs not extending beyond apex of sternum by more than 1/4 MOD.

Surface sculpture. Punctures dense. Labrum with larger and sparser punctures (i=1–2d) than clypeus (i<1d). Small impunctate shiny spot lateral to lateral ocellus. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula densely punctate mesally (i≤1d), less so laterally (i=1–2d). Mesopleuron with denser (i≤1d) punctures in upper half than ventrolateral half (i≤2d), the interspaces somewhat dull due to tessellate surface microsculpture. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc.

Structure. Preapical tooth obtuse. Labral apex with pair of small denticles, each preceded by longitudinal carina. Frontal keel not strongly raised. Scape with greatest length 1.7 × greatest width. F2 noticeably longer than wide (L/W ratio = 1.2). Preoccipital ridge not joining hypostomal carina, from which it is separated by less than 1 MOD at its terminal. Mesoscutellum moderately bigibbous. Axilla large, its lateral margin (L) more than half as long as mesoscutellar width (W) (L/W ratio = 0.6) and tip nearly extending as far back as apex of horizontal dorsal portion of mesoscutellum; axilla with tip clearly visible, but unattached to mesoscutellum for less than 2/5 the medial length of axilla; axilla with lateral margin arcuate. Fore wing with three submarginal cells. Pygidial plate apically truncate.

MALE: Description as for female except for usual secondary sexual characters and as follows: F2 shorter, not noticeably longer than wide (L/W ratio = 1.1); S4 and S5 with much longer coppery to silvery subapical hairs; pygidial plate with apex slightly concave and large deep punctures closely clustered basally and sparser apically, with the interspaces shining.

Figure 39. 

Epeolus deyrupi A female holotype, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male allotype, lateral habitus (scale bar 3 mm), and D female paratype axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Etymology

This species is named after its discoverer, Dr. Mark A. Deyrup, who recognized it as a new species and brought his discovery to my attention.

Distribution

Florida and coastal Georgia (Fig. 40).

Figure 40. 

Occurrence records of E. deyrupi known to the author (yellow circles).

Ecology

HOST RECORDS: The host species of E. deyrupi is/are presently unknown.

FLORAL RECORDS: Labels of examined voucher specimens indicate a floral association with Sideroxylon tenax L. (Sapotaceae).

Discussion

Epeolus deyrupi is a southeastern species that exhibits very little intraspecific morphological variation. Most of the known specimens of this species were collected in Highlands County, Florida. Based on known records, adults of E. deyrupi are active in spring.

Material studied

Type material. Primary: USA: Florida: Flamingo Villas Preserve (27.4423°N; 81.3782°W) (Highlands County), 26.v.2009, M. Deyrup, A. May, and H. Otte (holotype ♀ [CCDB-24583 F06], FSCA).

Secondary: USA: Florida: Allen David Broussard Catfish Creek Preserve State Park (27.8503°N; 81.4954°W) (Polk County), 08.vi.2009, M. Deyrup, A. May, and H. Otte (paratype ♂, ABS); Archbold Biological Station (27.1239°N; 81.3661°W) (Highlands County), 21.vi.2010, M. and L. Deyrup (paratype ♀, ABS); Archbold Biological Station (Highlands County), 29.v.1979, H.V. Weems, Jr. and S. Halkin (paratype ♀, LACM), 14.vi.2010, M. and L. Deyrup (paratype ♀, ABS); Flamingo Villas Preserve (27.4423°N; 81.3782°W) (Highlands County), 25.v.2009, M. Deyrup, A. May, and H. Otte (paratype ♀, ABS); Flamingo Villas Preserve (27.4487°N; 81.3767°W) (Highlands County), 01.vi.2009, M. Deyrup, A. May, and H. Otte (paratype ♀, ABS); Gould Rd Preserve (27.1336°N; 81.3256°W), 25.v.2009, M. Deyrup, A. May, and H. Otte (paratype ♀, PCYU), 26.v.2009, M. Deyrup, A. May, and H. Otte (paratype ♀, ABS); Lake Placid (Archbold Biological Station, Highlands County), 12.vi.1983, M. Deyrup (paratype ♀, ABS), 11.vi.1986, M. Deyrup (paratype ♀, ABS); The Nature Conservancy Tiger Creek Preserve (27.8077°N; 81.4816°W) (Polk County), 04.vi.2010, J. Dunlap, M. and N. Deyrup, and K. Dearborn (paratype ♀ [CCDB-24583 H04], PCYU); Tiger Creek Preserve (27.8133°N; 81.4868°W) (Polk County), 12.vi.2010, J. Dunlap, M. and N. Deyrup, and K. Dearborn (paratype ♀ [CCDB-24583 H02], USNM); Georgia: St Catherines Island (Liberty County), 24–27.vi.1989, Rozen, Quinter, and Eickwort (allotype ♂, AMNH), 27.vi.1974, R.O. Schuster and E.C. Teftner (paratype ♂, UCBME).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ADF0241. See Type material for specimens examined and sequenced (indicated by unique CCDB-plate and well number).

Epeolus diadematus sp. n.

Figs 41, 42, 92J

Epeolus torus Brumley, 1965. M.S. thesis, Utah State University, Logan 71 (♀) [nomen nudum].

Diagnosis

Epeolus diadematus does not closely resemble any other species of Epeolus except E. chamaesarachae. Unique in the genus to both species are each of the following morphological features: the vertexal area has two pairs of shiny (usually impunctate) protrusions, the mesoscutum is distinctly ornamented with mostly separate patches of (but some intermixed) pale and ferruginous tomentum, and the T2 fascia has two pairs of anterolateral extensions of tomentum. The difference is that in E. diadematus the mesopleuron has denser punctures ventrolaterally (most i≤1d) whereas in E. chamaesarachae the mesopleuron has sparser (most i>1d) and fewer punctures ventrolaterally.

Description

FEMALE: Length 6.9 mm; head length 2.0 mm; head width 2.6 mm; fore wing length 6.0 mm.

Integument coloration. Mostly black; notable exceptions as follows: partially to entirely ferruginous on mandible, antenna, pronotal collar, pronotal lobe, tegula, axilla, mesoscutellum, and legs. Mandible with apex darker than all but extreme base; preapical tooth lighter than mandibular apex. Antenna dark brown except scape, pedicel, and F1 brownish orange in part. Pronotal lobe and tegula pale ferruginous to amber. Wing membrane subhyaline, apically dusky. Legs more extensively reddish orange than brown or black.

Pubescence. Face with tomentum densest around antennal socket. Vertexal area with tomentum mostly ferruginous. Dorsum of mesosoma with bands of off-white and ferruginous short appressed setae. Dorsum of metasoma with bands of off-white to pale yellow short appressed setae. Pronotal collar with tomentum black medially, pale and ferruginous laterally. Mesoscutum with paramedian band of pale tomentum; ferruginous and pale tomentum encircling black spots medially and laterally, respectively, on anterior margin; and ferruginous tomentum along medial mesoscutal line and parapsidal line. Mesopleuron with upper half densely hairy, although scrobe visible; ventrolateral half nearly bare. Metanotum with tomentum uninterrupted, off white laterally and black medially. T1 with median diamond-shaped black discal patch enclosed by pale tomentum, except for medial separation at apex. T1 with apical fascia with black spot posterolaterally. T2–T4 with fasciae interrupted medially, T2 with fascia with paired anterolateral extensions of tomentum. T3 and T4 with fasciae interrupted laterally, with medial portion on apical margin and lateral portion encircling black tomentum on apical margin. T5 with two large patches of pale tomentum lateral to and separate from pseudopygidial area. T5 with pseudopygidial area lunate, its apex more than twice as wide as medial length, indicated by silvery setae on disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs extending beyond apex of sternum by ~1/3 MOD.

Surface sculpture. Punctures dense, but those of head and mesosoma sparser in some areas, larger, deep, and distinct. Labrum mostly with larger and sparser punctures (i=1–2d) than clypeus (i≤1d). Upper paraocular area and vertexal area sparsely punctate (i=1–2d), the interspaces shining. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate; the interspaces shining. Tegula densely punctate mesally (i=1–2d), much less so laterally (i>2d). Mesopleuron with denser (i<1d) punctures in upper half than ventrolateral half, although ventrolateral half with most interspaces small (i≤1d); the interspaces shining. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc.

Structure. Labral apex with two pairs of small denticles (the middlemost pair preceded by submedial pair of small denticles and separated by shallow concavity). Frontal keel strongly raised. Vertexal area with two pairs of nearly impunctate shiny protrusions. Scape with greatest length 1.6 × greatest width. F2 as long as wide (L/W ratio = 1.0). Preoccipital ridge not joining hypostomal carina, from which it is separated by no less than 1 MOD at its terminal. Mesoscutellum strongly bigibbous. Axilla intermediate in size, its lateral margin (L) nearly half as long as mesoscutellar width (W) (L/W ratio = 0.4–0.5) and tip not extending much beyond midlength of mesoscutellum (extending to <2/3 its length); axilla with tip conspicuously diverging from side of mesoscutellum, distinctly hooked, but unattached to mesoscutellum for less than 1/3 the medial length of axilla; axilla with lateral margin somewhat arcuate. Fore wing with three submarginal cells. Pygidial plate mostly hidden in holotype, but apically truncate in paratypes.

MALE: Description as for female except for usual secondary sexual characters and as follows: F2 shorter, nearly as long as wide (L/W ratio = 0.8); S4 and S5 with much longer coppery to silvery subapical hairs; pygidial plate apically rounded, with large deep punctures closely clustered apically and sparser basally, with the interspaces shining.

Figure 41. 

Epeolus diadematus A female holotype, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male paratype, lateral habitus (scale bar 3 mm), and D female paratype axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Etymology

The name is in reference to the four shiny, usually impunctate, tubercles on the vertexal area of the head of this species. From the Latin, “diadema” (royal headband).

Distribution

Texas and presumably Mexico, given the close proximity of some collection localities (e.g., Southmost, Texas) to the Mexico–United States border (Fig. 42).

Figure 42. 

Approximate geographic range of E. diadematus (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: The host species of E. diadematus is/are presently unknown.

FLORAL RECORDS: The label of one examined voucher specimen indicates a floral association with Engelmannia pinnatifida A.Gray ex Nutt. (Compositae). This species has also been collected from Aphanostephus riddellii Torr. & A. Gray (Compositae) (J. Neff, personal communication, 2016).

Discussion

This species and E. chamaesarachae are very similar in terms of integument coloration, pubescence, and structure, and are presumably sister species. Specimens of E. diadematus are distinct from those designated as E. chamaesarachae in that the mesopleuron has much denser punctation. The status of E. diadematus as a separate species is further supported by a separate BIN and large barcode sequence divergence (3.2%) from its nearest neighbor, E. chamaesarachae (Suppl. material 2). The ranges and flight seasons of these species also differ. With one exception, examined specimens of E. diadematus were collected in spring, and all are from Coastal or South Texas. By contrast, E. chamaesarachae occurs further west in the United States, and adults are active in late summer.

Material studied

Type material. Primary: USA: Texas: McAllen Botanical Gardens (McAllen), 21.xi.1982, C. Porter (holotype ♀, FSCA).

Secondary: USA: Texas: 5 mi SE Realitos (27.3980°N; 98.5490°W) (Duval County), 22.iv.2005, J.L. Neff and A. Hook (paratype ♂, CTMI); Ben Bolt (Jim Wells County), 12.v.1952, M. Cazier, W. Gertsch, and R. Schrammel (paratype ♀, AMNH); Brackenridge Field Laboratory (Austin, Travis County), 28.iv.1989, A. Hook (paratype ♂, CTMI); Chaparral Wildlife Management Area (Dimmit County), 06.iv.2007, J.L. Neff and A. Hook (paratype ♂, CTMI), 11.iv.2003, J.L. Neff and A. Hook (paratype ♂, CTMI); Dallas, 22.v.??06, W.D. Pierce (paratypes 2♂, USNM); Galveston?, L. Packer (paratype ♀ [CCDB-30383 F06], PCYU); Southmost (Cameron County), 13.vi.1953, Univ. Kans. Mex. Expedition (allotype ♂, KUNHM).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ADJ9659. See Type material for specimens examined and sequenced (indicated by unique CCDB-plate and well number).

Epeolus erigeronis Mitchell, 1962

Figs 43, 44, 92E

Epeolus erigeronis Mitchell, 1962. N. C. Agric. Exp. Stn. Tech. Bull. 152: 445 (♀).

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. erigeronis apart from all other North American Epeolus except E. ilicis and E. inornatus: the mandible is simple; the axilla does not attain the midlength of the mesoscutellum but the free portion is distinctly hooked, with the tip unattached to the mesoscutellum for more than 1/3 of the entire medial length of the axilla; the pronotal collar and metasomal terga are black; the metasomal terga have rather fine punctures; and the pseudopygidial area of the female is distinctly campanulate with the apex <2 × the medial length and not in contact with two large patches of pale tomentum (one on each side) throughout its length (in contact only at apex, diverging basally). Although in all three species the mesopleuron is closely and evenly punctate, in E. erigeronis the punctures are more variable in size, with many smaller punctures among large ones, and most interspaces are narrower such that the surface appears to be very coarsely and densely rugose-punctate. By contrast, in E. ilicis and E. inornatus the mesopleuron has punctures that are similar in size and shiny interspaces that are commonly equal to the puncture diameters.

Redescription

FEMALE: Length 8.6 mm; head length 2.2 mm; head width 3.0 mm; fore wing length 6.3 mm.

Integument coloration. Mostly black; notable exceptions as follows: partially to entirely ferruginous on mandible, labrum, antenna, pronotal lobe, tegula, and legs. Mandible with apex darker than all but extreme base. Antenna brown except scape, pedicel, and F1 orange in part. Pronotal lobe and tegula pale ferruginous to amber. Wing membrane subhyaline, apically dusky. Legs more extensively reddish orange than brown or black.

Pubescence. Face with tomentum densest around antennal socket. Tomentum slightly sparser on clypeus; upper paraocular and frontal areas, and vertexal area mostly exposed. Dorsum of mesosoma and metasoma with bands of off-white to pale yellow short appressed setae. Mesoscutum with paramedian band. Mesopleuron with upper half hairy, except beneath base of fore wing (hypoepimeral area); ventrolateral half nearly bare. Metanotum with tomentum uninterrupted except for median bare patch in posterior half, uniformly off white. T1 with discal patch quadrangular and very wide, the basal and apical fasciae only narrowly joined laterally. T1 and T2 with apical fasciae interrupted medially, those of T2 and T3 somewhat broader laterally, T2 with fascia with faint anterolateral extensions of sparser tomentum. T3 and T4 with fasciae complete. T5 with two large patches of pale tomentum lateral to and separate from pseudopygidial area. T5 with pseudopygidial area campanulate, its apex less than twice as wide as medial length, indicated by silvery setae on impressed disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs extending beyond apex of sternum by 1/3 MOD.

Surface sculpture. Punctures dense. Labrum with larger punctures than clypeus, but punctures of both equally dense (i<1d). Small impunctate matte spot lateral to lateral ocellus. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula very densely punctate mesally (i<1d), less so laterally (i=1–2d). Mesopleuron with ventrolateral half coarsely and densely rugose-punctate (i<1d), the interspaces somewhat dull due to surface microsculpture; mesopleuron with many smaller punctures among large ones, punctures more or less equally dense throughout. Metasomal terga with punctures very fine, dense (i=1–2d), evenly distributed on disc; the interspaces shining somewhat.

Structure. Mandible without preapical tooth. Labrum with pair of small subapical denticles not preceded by carinae. Frontal keel not strongly raised. Scape with greatest length 1.8 × greatest width. F2 noticeably longer than wide (L/W ratio = 1.6). Preoccipital ridge not joining hypostomal carina, from which it is separated by no less than 1 MOD at its terminal. Mesoscutellum weakly bigibbous. Axilla intermediate in size, its lateral margin (L) nearly half as long as mesoscutellar width (W) (L/W ratio = 0.4–0.5) and tip attaining midlength of mesoscutellum; axilla with tip conspicuously diverging from side of mesoscutellum, distinctly hooked, and axilla with free portion 2/5 its medial length; axilla with lateral margin arcuate and carinate. Fore wing with three submarginal cells. Pygidial plate apically truncate.

MALE: Description as for female except for usual secondary sexual characters and as follows: F2 shorter, but still longer than wide (L/W ratio = 1.3); S4 and S5 with much longer coppery to silvery subapical hairs; pygidial plate apically rounded, with large deep punctures closely clustered basomedially and sparser apically and laterally, with the interspaces shining.

Figure 43. 

Epeolus erigeronis A female, lateral habitus (scale bar 3 mm) B female, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Distribution

South Atlantic states (Fig. 44).

Figure 44. 

Approximate geographic range of E. erigeronis (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: The host species of E. erigeronis is/are presently unknown.

FLORAL RECORDS: Mitchell (1962) indicated floral associations with Erigeron quercifolius Lam. (Compositae), Hypericum L. (Hypericaceae), and Melilotus albus Medik. (Leguminosae). Labels of examined voucher specimens further indicate associations with Clinopodium ashei (Weath.) Small (Lamiaceae), Ilex glabra (L.) A. Gray (Aquifoliaceae), and Vaccinium darrowii Camp (Ericaceae).

Discussion

Epeolus erigeronis exhibits very little intraspecific morphological variation. However, in some specimens the axillae are partially ferruginous whereas in others they and the mesoscutellum are entirely black. Based on examined records, adults of E. erigeronis are active throughout spring.

Although BIN-compliant sequences are presently not available for E. erigeronis, four partial sequences (three 422 bp and one 394 bp in length) are available for specimens from North and South Florida, and these sequences form a distinct cluster that does not include any sequences from other Epeolus species in a NJ tree (Suppl. material 2).

Material studied

Type material. Primary: USA: Florida: Levy County, 13.iv.1955, H.V. Weems, Jr. (holotype ♀, FSCA).

Secondary: USA: Florida: Alachua County, 15.iv.1955, R.A. Morse (paratype ♀, FSCA); Levy County, 13.iv.1955, H.V. Weems, Jr. (allotype ♂, FSCA); North Carolina: Southport, 24.vi.1928, T.B. Mitchell (paratype ♀, NHMUK).

DNA barcoded material with BIN-compliant sequences

Unavailable.

Non-barcoded material examined

USA: Florida: 5 mi S Paynes Prairie (SE Gainesville, Alachua County), 05–12.v.1996, B.D. Sutton (1♀, FSCA); Apalachicola National Forest (30.3292°N; 84.5052°W) (Wakulla County), 08–15.v.2005, Ronquist lab (1♀, PCYU); Archbold Biological Station (Highlands County), 10.v.1979, H.V. Weems, Jr. and S. Halkin (1♀, BBSL), 17–23.iv.2007, S.M. Paiero (1♂, DEBU), 17.v.2005, M. Deyrup (1♀, ABS), 08.iv.1980, H.V. Weems, Jr. and F.E. Lohrer (1♀, FSCA), 24.iii.1980, H.V. Weems, Jr. and F.E. Lohrer (1♂, FSCA); Archbold Biological Station (27.1838°N; 81.3532°W) (Highlands County), 23.v.2010, M. Deyrup (1♀, ABS), 28.v.2010, M. Deyrup (1♀, ABS); Austin Cary Forest (Gainesville, Alachua County), 10.vi.1976 (1♂, UCBME), 16.x.1977, G.B. Fairchild (1♀, UCBME), 17.v.1991, L.R. Davis, Jr. (1♀, FSCA), 20.vi.1978, G.B. Fairchild and H.V. Weems, Jr. (1♀, UCBME); Brighton, 07.iv.1937, H.I. Scudder (1♀, CAS); Flamingo Villas Preserve (27.4487°N; 81.3767°W) (Highlands County), 01.vi.2009, M. Deyrup, A. May, and H. Otte (1♀, ABS); Flamingo Villas Preserve (27.4515°N; 81.3854°W) (Highlands County), 05.v.2010, M. Deyrup and J. Dunlap (1♀, ABS); Highlands Hammock State Park, 14.iv.1968, H.V. Weems, Jr. (2♀, FSCA); Kincaid Road (SE Gainesville, Alachua County), 03.iv.1999, B.D. Sutton (1♀, FSCA); Lake Placid (27.2195°N; 81.3803°W) (Highlands County), 14.iv.2010, M. Deyrup and J. Dunlap (1♀, ABS); New Smyrna Beach, 14.iii.1943, R.L. Usinger (1♂, EMEC); Osceola National Forest (Baker County and Columbia County line), 13–26.iv.1977, J.R. Wiley (1♂, FSCA), 01.v.2011, S. Lenberger (1♀, FSCA); San Felasco State Hammock Preserve, 16.v.1977, G.B. Fairchild and H.V. Weems, Jr. (1♀, UCBME).

Epeolus ferrarii sp. n.

Figs 45, 46

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. ferrarii apart from all other North American Epeolus except E. canadensis and E. compactus: in females, F2 is at least 1.2 × as long as wide; the mesoscutum has a small anteromedial patch of pale tomentum; the axilla is small to intermediate in size, not extending much beyond the midlength of the mesoscutellum (extending to <2/3 its length) but the free portion is more than 1/4 as long as the entire medial length of the axilla, and the axilla (except sometimes the tip) and mesoscutellum are black; the mesopleuron is closely (most i<1d) and evenly punctate; and the T2 fascia lacks lobe-like anterolateral extensions of tomentum, although it is broader laterally. Epeolus ferrarii is most similar to E. compactus, and in both species the T1 discal patch is typically quadrangular with the basal and apical fasciae subparallel and separated by a distinct longitudinal band, but in E. compactus the T2–T4 fasciae are not evenly broad or tapering until separated medially (as in E. ferrarii) but broadened medially into rounded lobes, which may be joined or separated. Epeolus canadensis differs from both species in that the T1 discal patch is distinctly triangular or semicircular (the basal fascia is conspicuously arched and fully continuous with the longitudinal band) and its medial longitudinal extent is more than 1/3 the lateral extent. In E. ferrarii the discal patch may be trapezoidal or almost semicircular, but if at all semicircular its medial longitudinal extent is at most 1/3 the lateral extent and the basal fascia and longitudinal band are at least joined at somewhat of an angle.

Description

MALE: Length 7.1 mm; head length 1.9 mm; head width 2.6 mm; fore wing length 6.0 mm.

Integument coloration. Mostly black; notable exceptions as follows: partially to entirely ferruginous on mandible, antenna, pronotal lobe, tegula, legs, and pygidial plate. Mandible with apex and preapical tooth darker than all but basal quarter. Antenna brown except F1 extensively orange. Pronotal lobe and tegula pale ferruginous to amber. Wing membrane subhyaline, apically dusky. Legs from tibia to tarsus extensively reddish orange. Pygidial plate orange along apical margin, otherwise dark brown.

Pubescence. Face with tomentum densest around antennal socket. Tomentum slightly sparser on clypeus; upper paraocular and frontal areas, and vertexal area mostly exposed. Dorsum of mesosoma and metasoma with bands of off-white to pale yellow short appressed setae. Mesoscutum with anteromedial horseshoe-shaped patch of pale tomentum. Mesopleuron densely hairy, except for two sparsely hairy circular patches (one behind pronotal lobe, a larger one occupying much of ventrolateral half of mesopleuron). Metanotum with tomentum uninterrupted, pale yellow laterally and black medially. T1 with median elliptical verging on semicircular discal patch. T1–T3 with apical fasciae medially interrupted, narrowed (broader laterally), and removed from apical margin; T2 with fascia without anterolateral extensions of tomentum. T4–T6 with fasciae complete, those of T4 and T5 somewhat narrowed medially. S4 and S5 with long coppery to silvery subapical hairs, which individually are often darker apically.

Surface sculpture. Punctures dense. Labrum with larger punctures than clypeus, but punctures of both equally dense (i≤1d). Small impunctate shiny spot lateral to lateral ocellus. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula very densely punctate mesally (i<1d), less so laterally (i=1–2d). Mesopleuron with ventrolateral half coarsely and densely punctate (i<1d) to rugose; mesopleuron with punctures more or less equally dense throughout (only few i=1d ventrolaterally). Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc.

Structure. Labral apex with pair of small denticles, each preceded by longitudinal carina. Frontal keel not strongly raised. Scape with greatest length 1.8 × greatest width. F2 as long as wide (L/W ratio = 1.0). Preoccipital ridge not joining hypostomal carina, from which it is separated by about 1.5 MOD at its terminal (difficult to see in holotype; described from paratypes). Mesoscutellum weakly bigibbous. Axilla small to intermediate in size, its lateral margin (L) less than half as long as mesoscutellar width (W) (L/W ratio = 0.4) and tip not extending much beyond midlength of mesoscutellum (extending to <2/3 its length); axilla with tip clearly visible, but unattached to mesoscutellum for less than 2/5 the medial length of axilla; axilla with lateral margin relatively straight and without carina. Fore wing with three submarginal cells. Pygidial plate apically rounded, with large deep punctures closely clustered medially and sparser laterally, with the interspaces shining.

FEMALE: Description as for male except for usual secondary sexual characters and as follows: F2 slightly but not noticeably longer than wide (L/W ratio = 1.1); T5 with large, nearly continuous patch of pale tomentum bordering and separate from pseudopygidial area present only in female; T5 with pseudopygidial area lunate, its apex more than twice as wide as medial length, indicated by silvery setae on impressed disc of apicomedial region elevated from rest of tergum; S4 and S5 with much shorter hairs (S5 with apical fimbria of coppery to silvery hairs extending beyond apex of sternum by ~1/3 MOD); pygidial plate apically truncate, with small, denser punctures.

Figure 45. 

Epeolus ferrarii A female allotype, lateral habitus (scale bar 3 mm) B female allotype, dorsal habitus (scale bar 3 mm) C male holotype, lateral habitus (scale bar 3 mm), and D female paratype axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Etymology

This species is named in honor of my colleague, Rafael Ferrari, with whom I collected this species in Southwestern New Mexico, USA.

Distribution

Arizona and New Mexico to southeastern Mexico (Fig. 46).

Figure 46. 

Approximate geographic range of E. ferrarii (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: The host species of E. ferrarii is/are presently unknown.

FLORAL RECORDS: Labels of examined voucher specimens indicate a floral association with Melilotus albus.

Discussion

Epeolus ferrarii is a cryptic species that most closely resembles E. canadensis and E. compactus, and can only be differentiated from these two species on the basis of very subtle differences in the patterns of pubescence on the metasomal terga. Its status as a separate species is supported by a separate BIN, but unusually its nearest neighbor is E. splendidus (a very different species, although presumably in the same species group), from which E. ferrarii exhibits a large barcode sequence divergence (3.9%). Although most species of Epeolus were described from a female name-bearing type, a male specimen is designated as the holotype of E. ferrarii because a barcode-compliant sequence is associated with it and because the collection locality is more precise than for the available female specimens, one of which is herein designated as the allotype.

Material studied

Type material. Primary: USA: New Mexico: 47 km S Animas (31.5438°N; 108.8757°W) (Co Rd C001), 30.viii.2015, R. Ferrari and T.M. Onuferko (holotype ♂ [CCDB-24583 H08], PCYU).

Secondary: Guatemala: Zacapa: San Lorenzo, xi.1986, M. Sharkey (paratype ♂, CNC).

Mexico: Chiapas: Yerbabuena (20 mi N Bochil), 21.v.1969, W.R.M. Mason (paratype ♂, CNC); Hidalgo: 2 mi N Pachuca, 24.viii.1962, M.G. Naumann (paratype ♀, KUNHM); Nuevo León: Cola de Caballo, 20.vi.1976, D. Weems (paratype ♂, FSCA); Puebla: 5 mi NE Teziutlán, 20.vi.1961, Univ. Kans. Mex. Expedition (paratype ♀, KUNHM); Veracruz: 10 km N Coscomatepec, 09.vii.1974, J.A. Chemsak, E. and J. Linsley, and J. Powell (paratype ♀, EMEC).

USA: Arizona: Southwestern Research Station (5 mi W Portal, Cochise County), 01.viii.1956, C. and M. Cazier (paratype ♀, AMNH), 02.viii.1956, C. and M. Cazier (paratype ♀, AMNH); New Mexico: 47 km S Animas (31.5438°N; 108.8757°W) (Co Rd C001), 30.viii.2015, R. Ferrari and T.M. Onuferko (paratypes 2♂ (1 barcoded [CCDB-24580 G07]), PCYU), 30.viii.2015, C. Parsons (paratype ♂, PCYU); 5 mi N Alamogordo (Otero County), 24.iv.1965, O.W. Richards (paratype ♀, NHMUK); Granite Gap (18 mi N Rodeo, Hidalgo County), 07.ix.1976, R.M. Bohart (allotype ♀, UCBME), 07.ix.1976, R.M. Bohart (paratypes 1♀, 1♂, UCBME); Texas: 23 mi W Fort Davis, 01.vi.1959, W.R.M. Mason (paratype ♀, CNC); Big Bend National Park, 04.vi.1970, C.W. O’Brien (paratype ♀, LACM); Grapevine Spring (Big Bend National Park), 20.v.1959, W.R.M. Mason (paratype ♀, CNC); Dugout Wells (Big Bend National Park), 22.v.1959, J.F. McAlpine (paratypes 3♀, CNC); Sanderson, 28–29.iv.1959, W.R.M. Mason (paratype ♀, CNC).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ADD6263. See Type material for specimens examined and sequenced (indicated by unique CCDB-plate and well number).

Epeolus flavofasciatus Smith, 1879

Figs 2C, 47, 48

Epeolus flavofasciatus Smith, 1879. Descr. New Species Hymen.: 103 (♀, ♂), new lectotype designation.

Triepeolus flavofasciatus Cockerell 1904. Ann. Mag. Nat. Hist. 13: 36.

Triepeolus agaricifer Cockerell, 1907c. Ann. Mag. Nat. Hist. 20: 60 (♂).

Diagnosis

The following morphological features in combination can be used to tell E. flavofasciatus apart from all other North American Epeolus: the dorsum of the mesosoma and metasoma have bright or pale yellow pubescence, the mesoscutum has distinct paramedian bands, the axilla does not attain the midlength of the mesoscutellum, and T1 has a median triangular or semicircular discal patch. Epeolus canadensis resembles E. flavofasciatus in that the integument is mostly black, the axilla does not attain the midlength of the mesoscutellum, and T1 has a median triangular or semicircular discal patch, but in E. canadensis the mesoscutum has a distinct anteromedial patch of pale tomentum instead of paramedian bands. Epeolus flavofasciatus is quite large for Epeolus (≥9 mm in length), and the pygidial plate of the male is narrower than that in most species, so males may be confused with Triepeolus. However, in E. flavofasciatus the mandible has a blunt, obtuse preapical tooth, whereas in all Triepeolus the mandible is simple.

Redescription

FEMALE: Length 9.6 mm; head length 2.4 mm; head width 3.3 mm; fore wing length 8.5 mm.

Integument coloration. Mostly black; notable exceptions as follows: partially to entirely ferruginous on mandible, antenna, pronotal lobe, tegula, axilla, legs, and pygidial plate. Mandible with apex darker than all but extreme base; preapical tooth lighter than mandibular apex (difficult to see in the E. flavofasciatus lectotype because mandible closed; described from non-type specimens). Antenna brown except scape, pedicel, and F1 extensively orange. F2 with orange spot basally. Pronotal lobe and tegula pale ferruginous to amber. Wing membrane dusky subhyaline, slightly darker at apex. Legs more extensively reddish orange than brown or black.

Pubescence. Face with tomentum densest around antennal socket. Tomentum slightly sparser on clypeus; upper paraocular and frontal areas, and vertexal area mostly exposed. Dorsum of mesosoma and metasoma with bands of off-white and bright to pale yellow short appressed setae. Mesoscutum with paramedian band. Mesopleuron sparsely hairy except mesally with densely hairy sigmoid patch and ventrally. Metanotum with tomentum uninterrupted, uniformly black (uniformly pale yellow in the E. agaricifer holotype and multiple non-type specimens, uniformly black or to varying degrees bright or pale yellow laterally and black medially in other non-type specimens). T1 with median semicircular black discal patch enclosed by pale tomentum (basal fascia widely separated medially and with much tomentum rubbed off in the E. flavofasciatus lectotype, but conspicuously arched and narrowly interrupted medially in non-type specimens). T2–T4 with fasciae complete, T2 with fascia without anterolateral extensions of tomentum. T5 with two large patches of pale tomentum lateral to and separate from pseudopygidial area. T5 with pseudopygidial area lunate, its apex more than twice as wide as medial length, indicated by silvery setae on disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs not extending beyond apex of sternum by much more than 1/4 MOD.

Surface sculpture. Punctures dense. Labrum with larger punctures than clypeus, but punctures of both equally dense (i<1d). Small impunctate matte spot lateral to lateral ocellus. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula very densely punctate mesally (i<1d), less so laterally (i=1–2d). Mesopleuron with ventrolateral half densely punctate (i<1d); mesopleuron with punctures more or less equally dense throughout. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc.

Structure. Labral apex with pair of small denticles preceded by submedial pair of small denticles and separated by shallow concavity. Frontal keel not strongly raised. Scape with greatest length 1.7 × greatest width. F2 noticeably longer than wide (L/W ratio = 1.4). Preoccipital ridge not joining hypostomal carina, from which it is separated by about 1.5–2 MOD at its terminal (difficult to see in the E. flavofasciatus lectotype; described from non-type specimens). Mesoscutellum moderately bigibbous. Axilla small to intermediate in size, its lateral margin (L) less than half as long as mesoscutellar width (W) (L/W ratio = 0.4) and tip not extending beyond midlength of mesoscutellum; axilla with tip clearly visible, but unattached to mesoscutellum for less than 1/3 the medial length of axilla; axilla with lateral margin relatively straight and without carina. Fore wing with three submarginal cells. Pygidial plate apically truncate.

MALE: Description as for female except for usual secondary sexual characters and as follows: F2 shorter, but still longer than wide (L/W ratio = 1.2); S3–S5 with much longer coppery to silvery subapical hairs, which individually are often darker apically; pygidial plate unusually narrow (Triepeolus-like) and apically rounded, with large deep punctures closely clustered.

Figure 47. 

Epeolus flavofasciatus A female, lateral habitus (scale bar 3 mm) B female, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Distribution

Mexico, excluding the Baja California Peninsula, and southwestern United States to central America (Fig. 48).

Figure 48. 

Approximate geographic range of E. flavofasciatus (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: The host species of E. flavofasciatus is/are presently unknown.

FLORAL RECORDS: Labels of examined voucher specimens indicate floral associations with Heterotheca subaxillaris and Vicia L. (Leguminosae).

Discussion

Smith (1879) described E. flavofasciatus from both sexes, represented by two syntypes (one female and one male) deposited at the NHMUK. Both specimens were examined, and the female is herein designated as the lectotype because it is in better condition, because most Epeolus spp. are represented by female name-bearing types, and because Smith (1879) provided a more complete description of the female. The male syntype at the NHMUK is herein designated as the lectoallotype. Cockerell (1907) described this species under the name Triepeolus agaricifer, which Rightmyer (2008) synonymized under E. flavofasciatus. I have examined the male holotype specimen of T. agaricifer, and agree with Rightmyer’s treatment. Two specimens (both males) were barcoded, one of which is from Southeast Arizona, USA (nearer the type locality of T. agaricifer: Beulah, New Mexico, USA) and the other is from Jalisco, Mexico (nearer the type locality of E. flavofasciatus: Oaxaca, Mexico), and both were assigned the same BIN. Brumley also described this species under the manuscript name Epeolus artus [nomen nudum] in 1965.

There is some intraspecific variation in the pubescence on the metanotum, which ranges from entirely yellow to medially or mostly black, and T1, in which the apical fascia is either complete or interrupted medially, with differences not conforming to any discernable geographic pattern. Based on examined records, the range of this species appears to be quite continuous from the American Southwest to Central America.

Among the examined specimens of this species is one that appears to be the first known example of bilateral gynandromorphism in Epeolus (see Material studied). Descriptions and images of the aberrant features exhibited by the specimen are published separately (Onuferko 2018).

Material studied

Type material. Primary: Mexico: Oaxaca: (E. flavofasciatus lectotype ♀ [NHMUK, catalog number: 010812212]).

USA: New Mexico: Beulah, viii.????, T.D. Cockerell (T. agaricifer holotype ♂ [USNM, catalog number: 534034]).

Secondary: Mexico: Oaxaca: (E. flavofasciatus lectoallotype ♂ [NHMUK, catalog number: 010812250]).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ACZ9233. Specimens examined and sequenced.–Mexico: Jalisco: 8 km N Atemajac de Brizuela, 08.x.2008, L. Packer (1♂, PCYU).

USA: Arizona: vic. Hannagan Meadow (33.6300°N; 109.3200°W) (Greenlee County), 19–20.vii.1998, B. Harris (1♀, LACM).

Non-barcoded material examined

Guatemala: Escuintla: Volcán Pacaya, 30.xi.1975, S.W.T. Batra (1♀, USNM).

Mexico: Chiapas: San Cristóbal de las Casas, 29.v.1969, W.R.M. Mason (1♀, CNC); Durango: Coyotes (Durango Dist.), 08.viii.1947, D. Rockefeller Exp. Michener (1♀, BBSL);

Navíos (26 mi E El Salto), 02.viii.1964, L.A. Kelton (1♀, CNC); Michoacán: 17 mi N Hidalgo, 29.vii.1962, Univ. Kans. Mex. Expedition (2♀, KUNHM); Hidalgo, 12.vii.1963, F.D. Parker and L.A. Stange (1♂, UCBME); Morelos: 10 mi N Cuernavaca, 15.viii.1954, Univ. Kans. Mex. Expedition (1♀, KUNHM); Sinaloa: Las Palmitas, 13.ix.1977, E.I. Schlinger (2♀, EMEC); Tlaxcala: 8 mi WNW Apizaco, 18.vi.1961, Univ. Kans. Mex. Expedition (1♀, KUNHM).

USA: Arizona: Catalina Mountains (19 HkHy), 25.vii.1954, G.D. Butler (1♂, KUNHM); Catalina Mountains (24 HkHy), 26.vii.1954, G.D. Butler (1♂, KUNHM); Catalina Mountains (25 HkHy), 14.viii.1954, G. Bohart and G. Butler (1♂, KUNHM); Catalina Mountains (26 HkHy), 14.viii.1954, G. Bohart and G. Butler (1♂, KUNHM), 25.viii.1954, G.D. Butler (1♀, BBSL), 25.viii.1954, G.D. Butler (1♀, KUNHM); Flagstaff (Coconino County), 25.vii.1952, M. Cazier, W. Gertsch, and R. Schrammel (1 chimera, AMNH); Grand Canyon, 19.viii.??39 (1♀, BBSL); Mount Graham (Graham County), 29.viii.1995, J.G. Rozen and S.A. Budick (1♀, AMNH); Pinaleno Mountains (Graham County), 22.viii.1989, Rozen, Foster, and Brewster (1♀, AMNH); Ramsey Canyon (Huachuca Mountains, Cochise County), 1954, W.M. Mann (2♂, USNM); Rose Peak (30 mi N Clifton, Greenlee County), 16.viii.1964, C.D. Michener (1♂, KUNHM); San Francisco Mountains (Flagstaff, Coconino County), 15.viii.1934, E.L. Bell (1♀, AMNH); Santa Catalina Mountains (Pima County), J.L. Neff (1♂, LACM); New Mexico: Sapello Canyon (San Miguel County), 26.vii.??02 (1♂, USNM), 27.vii.??02 (1♀, USNM), 31.vii.-01.viii.1963, T.C. Emmel (1♀, LACM); Texas: Big Bend National Park (Brewster County), 14.viii.1976, R.T. Ross (1♂, UCBME).

Epeolus floridensis Mitchell, 1962

Figs 49, 50, 97B

Epeolus floridensis Mitchell, 1962. N. C. Agric. Exp. Stn. Tech. Bull. 152: 446 (♀).

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. floridensis apart from all other North American Epeolus: the axilla is large, with the tip extending as far back as or beyond the posterior margin of the mesoscutellum, dilated laterally, and like the mesoscutellum ferruginous; the mesopleuron is closely (i≤1d) and evenly punctate; T1 is (with few exceptions) ferruginous and with a distinct, although sometimes medially-interrupted, basal fascia; the mesoscutum and metasomal terga have bands of pale gray to white short appressed setae; at least the T1–T3 apical fasciae are distinctly interrupted medially; and the pseudopygidial area of the female is lunate with the apex <2 × the medial length. Epeolus floridensis is similar to E. howardi, but in E. howardi the mesoscutum and metasomal terga have bands of bright or pale yellow short appressed setae and the metasomal terga (including T1) are black. Epeolus floridensis is also similar to E. packeri, but in E. packeri the T1 basal fascia is absent or reduced to a pair of small patches of pale tomentum, the metasomal terga (including T1) are black, and the pseudopygidial area of the female is lunate with the apex >2 × the medial length.

Redescription

FEMALE: Length 7.5 mm; head length 2.1 mm; head width 2.7 mm; fore wing length 5.5 mm.

Integument coloration. Black in part, at least partially ferruginous on mandible, labrum, clypeus, antenna, pronotal collar, pronotal lobe, tegula, axilla, mesoscutum, mesoscutellum, metanotum, mesopleuron, metapleuron, propodeum, legs, T1, T5, pygidial plate, and metasomal sterna. Mandible with apex darker than rest of mandible; preapical tooth slightly lighter than mandibular apex. Antenna brown and orange in part. Pronotal lobe and tegula pale ferruginous to amber. Mesoscutum almost entirely reddish brown. Wing membrane subhyaline, apically dusky. Legs more extensively reddish orange than brown or black.

Pubescence. Face with tomentum densest around antennal socket. Tomentum slightly sparser on clypeus; upper paraocular and frontal areas, and vertexal area mostly exposed. Dorsum of mesosoma and metasoma with bands of off-white to pale gray short appressed setae. Mesoscutum with paramedian band. Mesopleuron sparsely hairy, but tomentum moderately dense along margins. Metanotum with tomentum uninterrupted, uniformly off white. T1 with discal patch quadrangular and very wide, the basal and apical fasciae only narrowly joined laterally by few sparsely scattered pale hairs. T1–T4 with apical fasciae interrupted medially and somewhat broader laterally, T2 with fascia without anterolateral extensions of tomentum. T5 with two patches of pale tomentum lateral to and contacting pseudopygidial area. T5 with pseudopygidial area lunate, its apex less than twice as wide as medial length, indicated by silvery setae on impressed disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs not extending beyond apex of sternum by more than 1/4 MOD.

Surface sculpture. Punctures dense. Labrum with larger and sparser punctures (i=1–2d) than clypeus (i<1d). Upper paraocular and frontal areas, and vertexal area with punctures equally dense. Impunctate spot lateral to lateral ocellus absent in holotype, but shiny spot present in non-type specimens. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula densely punctate mesally (i≤1d), less so laterally (i=1–2d). Mesopleuron with ventrolateral half densely punctate (i≤1d), the interspaces shining; mesopleuron with punctures more or less equally dense throughout. Metasomal terga with punctures very fine, dense (i=1–2d), evenly distributed on disc; the interspaces shining somewhat.

Structure. Preapical tooth inconspicuous, blunt and obtuse. Labrum with pair of small subapical denticles not preceded by carinae. Frontal keel not strongly raised. Scape with greatest length 1.8 × greatest width. F2 noticeably longer than wide (L/W ratio = 1.6). Preoccipital ridge not joining hypostomal carina, from which it is separated by about 1.5 MOD at its terminal (difficult to see in holotype; described from non-type specimens). Mesoscutellum weakly bigibbous. Axilla large, its lateral margin (L) more than half as long as mesoscutellar width (W) (L/W ratio = 0.6) and tip extending as far back as apex of horizontal dorsal portion of mesoscutellum; axilla with tip clearly visible, but unattached to mesoscutellum for less than 1/3 the medial length of axilla; axilla with lateral margin arcuate. Fore wing with three submarginal cells. Pygidial plate apically truncate.

MALE: Description as for female except for usual secondary sexual characters and as follows: upper paraocular area very finely and sparsely punctate in part, the interspaces shining; F2 shorter, but still longer than wide (L/W ratio = 1.3); S4 and S5 with much longer coppery to silvery subapical hairs; pygidial plate apically rounded, with large deep punctures closely clustered basomedially and sparser apically and laterally, with the interspaces shining.

Figure 49. 

Epeolus floridensis A female, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Distribution

Florida peninsula (Fig. 50).

Figure 50. 

Occurrence records of E. floridensis known to the author (yellow circles).

Ecology

HOST RECORDS: The host species of E. floridensis is/are presently unknown.

FLORAL RECORDS: Mitchell (1962) indicated a floral association with Eriogonum tomentosum Michx. (Polygonaceae). Labels of examined voucher specimens further indicate associations with Licania michauxii Prance (Chrysobalanaceae), Ptilimnium capillaceum (Michx.) Raf. (Apiaceae), and Sabal etonia Swingle ex Nash (Arecaceae).

Discussion

Epeolus floridensis exhibits very little intraspecific morphological variation. However, one specimen was observed in which T1 is as dark as the remaining terga rather than bright ferruginous, the usual state. Also, in some males the upper paraocular area has comparatively fewer punctures than in females while in other specimens punctures are similarly dense between the sexes. Based on examined records, adults of E. floridensis appear to be most active in spring, although Mitchell (1962) lists some paratypes that were collected in mid-July.

Material studied

Type material. Primary: USA: Florida: Arcadia (DeSoto County), 27.iv.1955, H.E. and M.A. Evans (holotype ♀ [CUIC, catalog number: 00015349]).

Secondary: USA: Florida: Arcadia (DeSoto County), 27.iv.1955, H.E. and M.A. Evans (allotype ♂ [CUIC, catalog number: 00015348]), 27.iv.1955, H.E. and M.A. Evans (paratypes 1♀, 1♂, NCSU).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ACZ9059. Specimens examined and sequenced.–USA: Florida: Archbold Biological Station (Highlands County), 28.iv.-18.v.2008, S.M. Paiero (1♀, 1♂, DEBU); Lake Placid (Highlands County), 17.v.2014, S. Lenberger (1♀, FSCA).

Non-barcoded material examined

USA: Florida: Archbold Biological Station (27.1838°N; 81.3532°W) (Highlands County), 28.v.2010, M. Deyrup (1♀, ABS); Lake Wales Ridge State Forest (27.6611°N; 81.3964°W) (Polk County), 30.iv.2009, M. Deyrup, A. May, and H. Otte (1♀, ABS); Lake Wales Ridge State Forest (27.6933°N; 81.4279°W) (Polk County), 30.iv.2009, M. Deyrup, A. May, and H. Otte (1♂, ABS); Lake Wales Ridge State Forest (27.6915°N; 81.4282°W) (Polk County), 06.v.2009, M. Deyrup, A. May, and H. Otte (1♀, ABS); N FWC Carter Creek (27.5313°N; 81.4104°W) (Highlands County), 15.v.2010, J. Dunlap, M. and N. Deyrup, and K. Dearborn (1♂, ABS); Saddle Blanket Lakes (27.6696°N; 81.5758°W) (Polk County), 07.v.2009, M. Deyrup (1♂, ABS); Saddle Blanket Lakes (27.6716°N; 81.5759°W) (Polk County), 08.v.2009, M. Deyrup, A. May, and H. Otte (1♀, ABS); Walk-In-The-Water State Forest (27.7613°N; 81.4877°W) (Polk County), 29.v.2010, M. Deyrup (1♀, ABS).

Epeolus gibbsi sp. n.

Figs 3D, 51, 52, 96C, 97F

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. gibbsi apart from all other North American Epeolus: the mandible has a blunt, obtuse preapical tooth; in females, F2 is less than 1.2 × as long as wide; the axilla does not attain the midlength of the mesoscutellum but the free portion is distinctly hooked, with the tip unattached to the mesoscutellum for more than 1/3 of the entire medial length of the axilla; the mesopleuron is closely and evenly punctate (i≤1d), with the interspaces shining and punctures similar in size; the legs are usually darker, at least from the metacoxa to metatibia; the metasomal terga have rather fine punctures; S4 and S5 of the male have long curved coppery to silvery subapical hairs; and the pseudopygidial area of the female is distinctly campanulate with the apex <2 × the medial length and in contact with two large patches of pale tomentum (one on each side [the two are parallel to each other]) throughout its length. Epeolus gibbsi most closely resembles E. ilicis and E. inornatus, but in males of the latter S4 and S5 have short straight subapical hairs and in both E. ilicis and E. inornatus the mandible is simple, and in females of both species F2 is more than 1.2 × as long as wide and the pseudopygidial area is not in contact with two large patches of pale tomentum (one on each side) throughout its length (in contact only at apex, diverging basally).

Description

FEMALE: Length 7.3 mm; head length 1.9 mm; head width 2.5 mm; fore wing length 5.8 mm.

Integument coloration. Mostly black; notable exceptions as follows: partially to entirely ferruginous on mandible, antenna, pronotal lobe, tegula, and legs. Mandible with apex darker than all but extreme base; preapical tooth lighter than mandibular apex (difficult to see in holotype; described from paratype). Antenna dark brown except scape and F1 reddish brown in part. Pronotal lobe dark brown to black. Tegula pale ferruginous to amber. Wing membrane subhyaline, apically dusky. Legs more extensively reddish orange than brown or black.

Pubescence. Face with tomentum densest around antennal socket. Tomentum slightly sparser on clypeus; upper paraocular and frontal areas, and vertexal area mostly exposed. Dorsum of mesosoma and metasoma with bands of off-white to pale yellow short appressed setae. Mesoscutum with paramedian band. Mesopleuron densely hairy, except for two sparsely hairy circular patches (one behind pronotal lobe, a larger one occupying much of ventrolateral half of mesopleuron). Metanotum with tomentum uninterrupted except for median bare patch in posterior half, uniformly off white. T1 with median elliptical verging on semicircular discal patch. T1 and T2 with apical fasciae interrupted medially, those of T2 and T3 somewhat broader laterally, T2 with fascia with anterolateral extensions of sparser tomentum. T3 and T4 with fasciae complete. T5 with two large patches of pale tomentum parallel to and contacting pseudopygidial area throughout its length. T5 with pseudopygidial area campanulate, its apex less than twice as wide as medial length, indicated by silvery setae on impressed disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs extending beyond apex of sternum by ~1/3 MOD.

Surface sculpture. Punctures dense. Labrum with larger punctures than clypeus, but punctures of both equally dense (i<1d). Impunctate spot lateral to lateral ocellus absent in holotype, but shiny spot present in some paratypes. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula densely punctate (i≤2d). Mesopleuron with ventrolateral half densely punctate (i≤1d), the interspaces shining; mesopleuron with punctures similar in size and more or less equally dense throughout. Metasomal terga with punctures very fine, dense (i=1–2d), evenly distributed on disc; the interspaces shining somewhat.

Structure. Preapical tooth blunt and obtuse. Labrum with pair of small subapical denticles not preceded by carinae. Frontal keel not strongly raised. Scape with greatest length 1.8 × greatest width. F2 not noticeably longer than wide (L/W ratio = 1.1). Preoccipital ridge not joining hypostomal carina, from which it is separated by about 1 MOD at its terminal (difficult to see in holotype; described from paratype). Mesoscutellum strongly bigibbous. Axilla small to intermediate in size, its lateral margin (L) less than half as long as mesoscutellar width (W) (L/W ratio = 0.4) and tip attaining midlength of mesoscutellum; axilla with tip conspicuously diverging from side of mesoscutellum, distinctly hooked, and axilla with free portion 2/5 its medial length; axilla with lateral margin relatively straight and without carina. Fore wing with three submarginal cells. Pygidial plate apically truncate.

MALE: Description as for female except for usual secondary sexual characters and as follows: F2 shorter, as long as wide (L/W ratio = 1.0); S4 and S5 with much longer coppery to silvery subapical hairs; pygidial plate apically rounded, with large deep punctures closely clustered.

Figure 51. 

Epeolus gibbsi A female holotype, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male allotype, lateral habitus (scale bar 3 mm), and D female holotype axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Etymology

This species is named after its discoverer, Prof. Jason Gibbs, who collected the specimen herein designated as the holotype, recognized it as an unusual find, and brought his discovery to my attention.

Distribution

Upper midwest and adjacent Canada (Fig. 52).

Figure 52. 

Occurrence records of E. gibbsi known to the author (yellow circles).

Ecology

HOST RECORDS: The holotype of E. gibbsi was collected in an area where Colletes brevicornis and C. kincaidii were in abundance, the latter of which is likely associated with E. minimus, which was also present at the site, as was E. ainsliei and its tentative host C. susannae (J. Gibbs, personal communication, 2017).

FLORAL RECORDS: Unknown.

Discussion

What Romankova (2004) identified as E. ilicis, which constituted a new record of that species in Canada, might actually be E. gibbsi and/or E. inornatus. Unfortunately, the vouchered material from that study (three specimens from Ontario) cannot be traced, so the presence of E. ilicis in Canada has not been confirmed in the present study. Epeolus ilicis has been reported from the New England states, though the only examined specimen from that region (a male from Massachusetts) that has been identified as E. ilicis (by Richard L. Brumley) appears to actually be E. inornatus based on the very short straight subapical hairs on S4 and S5. In Canada, E. gibbsi is only confirmed from southern Manitoba, so the specimens from southern Ontario studied by Romankova could represent any of the three species. The key presented in Onuferko (2017) still works for E. ilicis, but can also lead to E. gibbsi and E. inornatus with the modifications presented in Suppl. material 3 starting at couplet 4. Presently, only a single 422 bp sequence is available for E. ilicis (a male specimen from Florida, USA), which clusters with sequences of E. zonatus (Suppl. material 2), and all were assigned the same BIN. In addition to the diagnostic morphological features that separate E. gibbsi from other similar species (notably E. erigeronis, E. ilicis, and E. inornatus, for which only partial sequences 394 to 422 bp in length are available), the status of E. gibbsi as a separate species is supported by a separate BIN and large barcode sequence divergence (4.7%) from its nearest neighbor, E. glabratus. Based on the few known records, adults of E. gibbsi appear to be active in late spring/early summer.

Material studied

Type material. Primary: Canada: Manitoba: Spruce Woods Provincial Park (49.6630°N; 99.2790°W) (Spirit Sands, Division 7), 07.vii.2017, J. Gibbs and Nozoe (holotype ♀ [CCDB-30345 D02], JBWM).

Secondary: USA: Wisconsin: Two Rivers, 26.vi.1911 (allotype ♂, CUM), 26.vi.1911 (paratypes 1♀, 6♂, CUM).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ADI6791. See Type material for specimens examined and sequenced (indicated by unique CCDB-plate and well number).

Epeolus glabratus Cresson, 1878

Figs 53, 54, 93B

Epeolus glabratus Cresson, 1878. Trans. Am. Entomol. Soc. 7: 90 (♂).

Pyrrhomelecta glabrata Ashmead, 1899. Trans. Am. Entomol. Soc. 26: 66.

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. glabratus apart from all other North American Epeolus except E. lectoides: the axilla is elongate, extending well beyond the midlength of the mesoscutellum but not as far back as its posterior margin, and the free portion is distinctly hooked; the mesopleuron has sparser punctures ventrolaterally (most i>1d) than in upper half, with the interspaces shining; the metasomal terga have minute, shallow punctures; T2–T4 are medially bare; and the pseudopygidial area of the female is distinctly campanulate with the apex <2 × the medial length. Whereas in E. lectoides the pronotal collar is black, as are sometimes the axilla and mesoscutellum, and the metasomal terga are black and fasciate, in E. glabratus the pronotal collar, axilla, mesoscutellum, and discs of T1 and T2 are ferruginous and the pale pubescence on the metasomal terga are commonly reduced to discrete lateral patches.

Redescription

MALE: Length 8.4 mm; head length 1.8 mm; head width 2.5 mm; fore wing length 7.9 mm.

Integument coloration. Black in part, at least partially ferruginous on mandible, labrum, clypeus, antenna, pronotal collar, pronotal lobe, tegula, axilla, mesoscutum, mesoscutellum, mesopleuron, metapleuron, legs, T1, T2, pygidial plate, and metasomal sterna. Mandible with apex darker than rest of mandible; preapical tooth lighter than mandibular apex (difficult to see in holotype because mandible closed; described from non-type specimens). Antenna brown except scape, pedicel, and F1 extensively orange. F2 with orange spot basally. Pronotal lobe and tegula pale ferruginous to amber. Wing membrane dusky subhyaline, slightly darker at apex. Legs more extensively reddish orange than brown or black.

Pubescence. Face with tomentum densest around antennal socket. Tomentum slightly sparser on clypeus; upper paraocular and frontal areas, and vertexal area mostly exposed. Dorsum of mesosoma and metasoma with bands of off-white to pale yellow short appressed setae. Mesoscutum with paramedian band. Mesopleuron sparsely hairy, but tomentum dense ventrally as well as between two sparsely hairy patches (one beneath base of fore wing (hypoepimeral area), a larger circular patch occupying much of ventrolateral half of mesopleuron). Metanotum with tomentum uninterrupted except for median bare patch in posterior half, uniformly off white. T1 with discal patch quadrangular and very wide, the basal and apical fasciae only narrowly joined laterally. T1 with basal and apical fasciae and T2–T4 with apical fasciae widely separated medially, the apical fasciae reduced to pairs of small patches somewhat broader laterally; T2 with fascia without anterolateral extensions of tomentum, although few sparsely scattered pale hairs present. Remaining metasomal terga mostly hidden in holotype, but T5 and T6 with complete or narrowly interrupted fasciae in non-type specimens. S4 and S5 with long coppery to silvery subapical hairs.

Surface sculpture. Punctures dense, but those of head and mesosoma sparser in some areas, larger, deep, and distinct. Labrum with larger punctures than clypeus, but punctures of both equally dense (i<1d). Small impunctate shiny spot lateral to lateral ocellus. Mesoscutum, mesoscutellum, and axilla very coarsely and densely punctate; the interspaces shining. Tegula densely punctate (i≤2d). Mesopleuron mostly with denser (i≤1d) punctures in upper half than ventrolateral half (i>1d), the interspaces shining. Metasomal terga with punctures very fine, dense (i≥1d), somewhat evenly distributed on disc; the interspaces shining somewhat.

Structure. Preapical tooth blunt and obtuse. Labrum with pair of small subapical denticles, each preceded by small discrete longitudinal ridge. Frontal keel not strongly raised. Scape with greatest length 1.7 × greatest width. F2 noticeably longer than wide (L/W ratio = 1.3). Preoccipital ridge not joining hypostomal carina, from which it is separated by less than 1 MOD at its terminal (difficult to see in holotype; described from non-type specimens). Mesoscutellum moderately bigibbous. Axilla large, its lateral margin (L) more than half as long as mesoscutellar width (W) (L/W ratio = 0.6) and tip extending well beyond midlength of mesoscutellum but not as far back as its posterior margin; axilla with tip conspicuously diverging from side of mesoscutellum, distinctly hooked, and axilla with free portion 2/5 its medial length; axilla with lateral margin relatively straight and with tip carinate. Fore wing with three submarginal cells. Pygidial plate mostly hidden in holotype, but apically rounded, with large deep punctures more or less evenly spaced throughout with the interspaces shining in non-type specimens.

FEMALE: Description as for male except for usual secondary sexual characters and as follows: F2 even longer than wide (L/W ratio = 1.5); T5 with two large patches of pale tomentum lateral to and separate from pseudopygidial area present only in female; T5 with pseudopygidial area campanulate, its apex less than twice as wide as medial length, indicated by silvery setae on impressed disc of apicomedial region elevated from rest of tergum; S4 and S5 with much shorter hairs (S5 with apical fimbria of coppery to silvery hairs extending beyond apex of sternum by ~1/3 MOD); pygidial plate apically truncate, with small, denser punctures.

Figure 53. 

Epeolus glabratus A female, lateral habitus (scale bar 3 mm) B female, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Distribution

Florida and coastal Georgia (Fig. 54).

Figure 54. 

Approximate geographic range of E. glabratus (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: The host species of E. glabratus is/are presently unknown.

FLORAL RECORDS: Mitchell (1962) indicated a floral association with Vicia. Labels of examined voucher specimens further indicate associations with Coreopsis L., Hyptis mutabilis (Rich.) Briq. (Lamiaceae), Ilex glabra, Pluchea odorata (L.) Cass. (Compositae), Polygonella myriophylla (Small) Horton (Polygonaceae), Richardia brasiliensis Gomes (Rubiaceae), Serenoa repens (W. Bartram) Small (Arecaceae), Spermacoce verticillata L., and Verbena brasiliensis Vell.

Discussion

Sequenced specimens of E. glabratus share the same BIN as those of E. lectoides. There is virtually no divergence (<1%) between the barcode sequences of the two species, but the morphological differences are pronounced. Structurally, E. glabratus and E. lectoides are identical, but in E. glabratus the pronotal collar, axilla, mesoscutellum, and discs of T1 and T2 are ferruginous, whereas in E. lectoides at least the pronotal collar and metasomal terga are entirely black. Epeolus glabratus appears to be restricted to Florida and parts of Georgia, and the prevalence of red integument coloration among Florida Hymenoptera is a well-known unexplained phenomenon (Deyrup and Eisner 2003). Except in some examined specimens from Georgia, in E. glabratus the metasomal fasciae are lacking; the pale pubescence is instead reduced to discrete lateral patches. By contrast, in E. lectoides the metasomal terga are always fasciate. Although both species inhabit Florida, E. glabratus (with red coloration and reduced pubescence on the metasomal terga) appears to be present only on the peninsula whereas E. lectoides (with fasciae and black metasomal terga) appears to be restricted to the Florida panhandle. Since the marked abundance of red coloration is coupled with a general loss of pubescence in E. glabratus, and since these are features restricted to specimens from a particular geographical region, I have opted to treat E. glabratus and E. lectoides as heterospecific, despite the lack of evidence of genetic divergence.

Material studied

Type material. Primary: USA: Georgia: H.K. Morrison (holotype ♂ [ANSP, catalog number: 2230]).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:AAF2273. Specimens examined and sequenced.–USA: Florida: Archbold Biological Station (27.1711°N; 81.3483°W) (Highlands County), 21–26.iv.2011, R.J. Pivar (1♂, DEBU); Archbold Biological Station (Highlands County), 07–13.v.1995, C. Darling (1♀, PCYU); N FWC Carter Creek (27.5313°N; 81.4104°W) (Highlands County), 11.v.2010, J. Dunlap, M. and N. Deyrup, and K. Dearborn (1♂, ABS).

Non-barcoded material examined

USA: Florida: Archbold Biological Station (Highlands County), 14.iv.1963, J.G and B.L. Rozen (1♀, AMNH); Doyle Conner Bldg (Gainesville, Alachua County), 12.vi.1996, C. Porter (1♀, FSCA), 18.vi.1996, C. Porter (2♀, FSCA), 26.vi.1996, C. Porter (1♂, FSCA); Gainesville (Alachua County), 03–17.vii.1987, BRC Hymenoptera Team (1♀, PCYU), 07.vi.1976, W.H. Pierce (1♀, UCBME), 16.vi.1991, F.J. Santana (1♀, FSCA), 17.vi.1976, W.H. Pierce (1♀, 1♂, UCBME); Lake Alice (29.6442°N; 82.3630°W) (University of Florida, Gainesville, Alachua County), 05.vi.2007, J.S. Ascher and G. Hall (2♀, AMNH); Lake Placid (Highlands County), 17.v.2014, S. Lenberger (1♂, FSCA); Lake Wales Ridge State Forest (27.6611°N; 81.3964°W) (Polk County), 06.v.2009, M. Deyrup, A. May, and H. Otte (1♀, ABS); Naples (Golden Gate Estates Subdivision, Collier County), 25.v.2013, S. Lenberger (1♂, FSCA); Near Wilcox (Gilchrist County), 27.v.1981, C. Porter, L. Stange, and H. Greenbaum (1♀, FSCA); Newberry (Alachua County), 15.vii.1973, E.E. Grissell (1♂, UCBME); Royal Palm Park, 12–18.iv.1923 (1♂, AMNH); San Felasco State Hammock Preserve, 20.v.1977, G.B. Fairchild and H.V. Weems, Jr. (1♂, UCBME); Sarasota (Sarasota County), 31.v.1993, F.J. Santana (2♀, FSCA); U.S. Highway 41 S Lake City (Columbia County), 19.vi.2014, S. Lenberger (2♀, FSCA); Georgia: St Catherines Island (Liberty County), 10–15.v.1991, E. Quinter and A. Sharkov (1♂, AMNH); St Catherines Island (South Beach, Liberty County), 27.vi.1974, R.O. Schuster and E.C. Teftner (1♀, UCBME).

Epeolus howardi Mitchell, 1962

Figs 55, 56

Epeolus howardi Mitchell, 1962. N. C. Agric. Exp. Stn. Tech. Bull. 152: 447 (♀).

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. howardi apart from all other North American Epeolus: the axilla is large, with the tip extending as far back as or beyond the posterior margin of the mesoscutellum, dilated laterally, and like the mesoscutellum ferruginous; the mesopleuron is closely (i≤1d) and evenly punctate; the metasomal terga are black; T1 has a distinct, although sometimes medially-interrupted, basal fascia; the mesoscutum and metasomal terga have bands of bright or pale yellow short appressed setae; at least the T1–T3 apical fasciae are distinctly interrupted medially; and the pseudopygidial area of the female is lunate with the apex <2 × the medial length. Epeolus howardi most closely resembles E. andriyi and E. floridensis, but in E. andriyi the axillae are shorter, not extending as far back as the posterior margin of the mesoscutellum, and in E. floridensis the mesoscutum and metasomal terga have bands of pale gray to white short appressed setae and T1 is (with few exceptions) ferruginous. Epeolus howardi is also similar to E. scutellaris, but in E. scutellaris the T1–T3 apical fasciae are complete or only very narrowly interrupted medially, and the pseudopygidial area of the female is lunate with the apex >2 × the medial length.

Redescription

FEMALE: Length 8.6 mm; head length 2.2 mm; head width 2.9 mm; fore wing length 6.0 mm.

Integument coloration. Black in part, at least partially ferruginous on mandible, labrum, clypeus, antenna, pronotal collar, pronotal lobe, tegula, axilla, mesoscutum, mesoscutellum, metanotum, mesopleuron, legs, T1, pygidial plate, and metasomal sterna. Mandible with apex darker than rest of mandible; preapical tooth slightly lighter than mandibular apex. Antenna brown and orange in part. Pronotal lobe and tegula pale ferruginous to amber. Mesoscutum reddish brown along lateral margin and with pair of reddish-brown markings near posterior margin between midline and parapsidal line. Wing membrane dusky subhyaline, slightly darker at apex. Legs more extensively reddish orange than brown or black. T1 dark in general, not contrasting strongly with remaining metasomal terga, but reddish brown laterally.

Pubescence. Face with tomentum densest around antennal socket. Clypeus, upper paraocular and frontal areas, and vertexal area mostly exposed. Dorsum of mesosoma and metasoma with bands of off-white to pale yellow short appressed setae. Mesoscutum with paramedian band. Mesopleuron sparsely hairy, but tomentum moderately dense along margins. Metanotum with tomentum uninterrupted, uniformly off white. T1 with discal patch quadrangular and very wide, the basal and apical fasciae only narrowly joined laterally by few sparsely scattered pale hairs. T1–T4 with apical fasciae interrupted medially and narrowed before becoming somewhat broader laterally, T2 with fascia without anterolateral extensions of tomentum. T5 with two patches of pale tomentum lateral to and contacting pseudopygidial area. T5 with pseudopygidial area lunate, its apex less than twice as wide as medial length, indicated by silvery setae on impressed disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs not extending beyond apex of sternum by more than 1/4 MOD.

Surface sculpture. Punctures dense. Labrum with larger punctures than clypeus, but punctures of both equally dense (i<1d). Upper paraocular area sparsely punctate in part, the interspaces shining. Small impunctate shiny spot lateral to lateral ocellus. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula densely punctate mesally (i≤1d), less so laterally (i=1–2d). Mesopleuron with denser (i≤1d) punctures in upper half than ventrolateral half (i≤2d), the interspaces shining. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc; the interspaces shining somewhat.

Structure. Preapical tooth inconspicuous, blunt and obtuse. Labrum with pair of small subapical denticles, each preceded by small discrete longitudinal ridge. Frontal keel not strongly raised. Scape with greatest length 1.8 × greatest width. F2 noticeably longer than wide (L/W ratio = 1.7). Preoccipital ridge not joining hypostomal carina, from which it is separated by less than 1 MOD at its terminal. Mesoscutellum weakly bigibbous. Axilla large, its lateral margin (L) more than half as long as mesoscutellar width (W) (L/W ratio = 0.7) and tip extending beyond apex of horizontal dorsal portion of mesoscutellum; axilla with tip clearly visible, but unattached to mesoscutellum for less than 2/5 the medial length of axilla; axilla with lateral margin arcuate. Fore wing with three submarginal cells. Pygidial plate apically truncate.

MALE: Description as for female except for usual secondary sexual characters and as follows: F2 shorter, but still longer than wide (L/W ratio = 1.3); S4 and S5 with much longer coppery to silvery subapical hairs; pygidial plate apically rounded, with large deep punctures more or less evenly spaced throughout, with the interspaces shining.

Figure 55. 

Epeolus howardi A female, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Distribution

Mid-Atlantic states to Texas (Fig. 56).

Figure 56. 

Approximate geographic range of E. howardi (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: According to Mitchell (1962), Colletes howardi Swenk is the suspected host of E. howardi.

FLORAL RECORDS: Mitchell (1962) indicated a floral association with Dalea pinnata (J.F.Gmel.) Barneby. Labels of examined voucher specimens further indicate associations with Heterotheca subaxillaris ssp. latifolia (Buckley) Semple, Symphyotrichum drummondii var. texanum (E.S. Burgess) G.L. Nesom, and Xanthisma texanum DC. (Compositae).

Discussion

Epeolus howardi is a southeastern species that appears to be uncommon, or at least uncommonly collected. In general, there is little morphological variation among examined specimens except in integument coloration; the mesoscutum and mesopleuron range from varying degrees of ferruginous to entirely black, with differences not conforming to any discernable geographic pattern. Based on known records, adults of E. howardi are active in late summer and much of autumn.

Material studied

Type material. Primary: USA: North Carolina: Southern Pines, 30.ix.1951, T.B. Mitchell (holotype ♀ [USNM, catalog number: 534046]).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ADK0941. Specimens examined and sequenced.–USA: Maryland: Denton (38.9196°N; 75.8273°W) (Caroline County), 19.viii.2012, S. Westre (1♂, BIML).

Non-barcoded material examined

USA: Texas: Austin (Travis County), 27.x.1981, J.L. Neff (1♀, CTMI); Brackenridge Field Laboratory (Austin, Travis County), 02.xi.1992, J.L. Neff (1♀, CTMI); Brazos County, 24.x.1960, A.H. Alex (1♀, USNM); Dallas, 15.x.??05, F.C. Bishopp (1♀, USNM); Sayersville (Bastrop County), 20.ix.1998, J.L. Neff (1♀, CTMI).

Epeolus ilicis Mitchell, 1962

Figs 3E, 57, 58, 92F, 97G, 100A

Epeolus ilicis Mitchell, 1962. N. C. Agric. Exp. Stn. Tech. Bull. 152: 448 (♀).

Epeolus vernalis Mitchell, 1962. N. C. Agric. Exp. Stn. Tech. Bull. 152: 455 (♀), syn. n.

Epeolus weemsi Mitchell, 1962. N. C. Agric. Exp. Stn. Tech. Bull. 152: 455 (♂), syn. n.

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. ilicis apart from all other North American Epeolus except E. erigeronis and E. inornatus: the mandible is simple; the axilla does not attain the midlength of the mesoscutellum but the free portion is distinctly hooked, with the tip unattached to the mesoscutellum for more than 1/3 of the entire medial length of the axilla; the pronotal collar and metasomal terga are black; the metasomal terga have rather fine punctures; and the pseudopygidial area of the female is distinctly campanulate with the apex <2 × the medial length and not in contact with two large patches of pale tomentum (one on each side) throughout its length (in contact only at apex, diverging basally). Epeolus ilicis is most similar to E. inornatus, and in both species the mesopleuron has punctures that are similar in size and shiny interspaces that are commonly equal to the puncture diameters. By contrast, in E. erigeronis the punctures are more variable in size, with many smaller punctures among large ones, and most interspaces are narrower such that the surface appears to be very coarsely and densely rugose-punctate. Whereas in E. inornatus the legs (and sometimes the pronotal lobe and tegula) are usually darker, at least from the metacoxa to metatibia, the dorsum of the mesosoma and metasoma have gray short appressed setae, and S4 and S5 of the male have short straight subapical hairs, in E. ilicis the pronotal lobe and legs are more extensively reddish orange than brown or black (at least the anterior surface of the metatibia and metatarsus are the same reddish orange color), the dorsum of the mesosoma and metasoma have gray but also usually some pale yellow short appressed setae, and S4 and S5 of the male have long curved coppery to silvery subapical hairs. Epeolus ilicis is also similar to E. gibbsi, but in E. gibbsi the mandible has a blunt, obtuse preapical tooth; in females F2 is less than 1.2 × as long as wide (it is more than 1.2 × as long as wide in female E. ilicis); and the pseudopygidial area of the female is in contact with two large patches of pale tomentum (one on each side [the two are parallel to each other]) throughout its length.

Figure 57. 

Epeolus ilicis A female, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male paratype, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum (photo of E. vernalis holotype [herein synonymized under E. ilicis]), dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Redescription

This species was recently redescribed (Onuferko 2017).

Distribution

Southeastern United States (Fig. 58).

Figure 58. 

Approximate geographic range of E. ilicis (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: Rozen (1989) described first instar E. ilicis based on two larvae recovered from the nest of Colletes brimleyi Mitchell on St. Catherines Island in Georgia, USA, from where conspecifics of the former have been recorded (see Material studied).

FLORAL RECORDS: Onuferko (2017) lists associations with five plant genera based on Mitchell (1962) and a record on Discover Life (Ascher and Pickering 2017 [then 2016]). Since the discovery of E. inornatus, a cryptic species very similar to E. ilicis whose name applies to at least one of Mitchell’s paratypes of E. ilicis (see Material studied under E. inornatus), my taxon concept of E. ilicis has changed. As a result, I have only been able to determine that records of Ilex glabra and Prunus angustifolia Marshall (Rosaceae), taken from the collection labels of the holotypes of E. ilicis and E. weemsi respectively, are associated with what is here understood to be the true E. ilicis.

Discussion

Both the holotype of E. ilicis and the holotype of E. vernalis were examined, and the two appear to be the same species. In Mitchell’s (1962) key, the two species were differentiated on the basis of whether or not (and if so to what degree) the metasomal fasciae are interrupted medially, but the T1–T3 apical fasciae are interrupted medially (those of T1 and T2 are somewhat more widely separated medially) in both holotype specimens and the T4 fascia is complete in the E. ilicis holotype and only very narrowly interrupted in the E. vernalis holotype. Moreover, the type locality is the same for both (Holly Shelter [Pender County], North Carolina, USA), and the two specimens were collected only 12 days apart.

Presently, only a single 422 bp sequence is available for E. ilicis (a male specimen from Florida, USA), which clusters with sequences of E. zonatus (Suppl. material 2), and all were assigned the same BIN. The Florida specimen is most similar to the holotype of E. weemsi, which Mitchell (1962) described before noting that it might be the male of E. vernalis. In both the sequenced specimen and E. weemsi holotype, S4 and S5 have long curved coppery to silvery subapical hairs, which are absent in the very similar E. inornatus but present in all other North American male Epeolus. Whereas I have opted to treat E. ilicis and E. zonatus as heterospecific based on remarkably consistent differences in integument coloration coupled with a general loss of pubescence in E. zonatus, despite the apparent lack of evidence of genetic divergence, the extremely subtle differences in integument coloration and pubescence among the holotypes of E. ilicis, E. vernalis, and E. weemsi seem to fall within the range of intraspecific variation, and therefore E. vernalis and E. weemsi are herein synonymized under E. ilicis. Although the three names were published simultaneously, priority of the name should be given to E. ilicis because the holotype is in the best condition (those of E. vernalis and E. weemsi have broken antennae and in the latter much of the pubescence is discolored or rubbed off), it is female and most Epeolus spp. were described from female name-bearing types (the holotype of E. weemsi is male), and because an allotype and paratypes were designated for E. ilicis but not E. vernalis or E. weemsi. This species appears to be quite common in the Southeastern United States, where it may be confused with E. erigeronis or E. inornatus.

Material studied

Type material. Primary: USA: Florida: Alachua County, 23.ii.1957, H.V. Weems, Jr. (E. weemsi holotype ♂, FSCA); North Carolina: Holly Shelter (Pender County), 30.v.1950, T.B. Mitchell (E. ilicis holotype ♀ [USNM, catalog number: 534048]), 18.v.1950, T.B. Mitchell (E. vernalis holotype ♀ [USNM, catalog number: 534607]).

Secondary: USA: Georgia: Fort Gordon (Richmond County), 25.iv.1959, R.R. Snelling (paratype ♂, NCSU); South Carolina: McClellanville, 12.v.??44, H.K. Townes (paratype ♂, NCSU), 19.v.??44, H. and G. Townes (paratype ♂, NCSU).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ACM5887. Specimens examined and sequenced.–USA: Florida: Apalachicola National Forest (30.3291°N; 84.5052°W) (Forest Rd 366, Leon County), 15–20.v.2005, A. Deans, S. Joshi, and D. Murray (1♂, AMNH).

Non-barcoded material examined

USA: Alabama: Bon Secour National Wildlife Refuge (Baldwin County), 05–07.v.1994, S.A. Marshall (1♀, DEBU); Florida: A.T. Slosson (1♀, AMNH); 3 mi NW Sopchoppy (near Sopchoppy River, Wakulla County), 19.iv.1979, G.B. Fairchild (3♀, FSCA); Blackwater River State Forest (4 mi N Munson, Santa Rosa County), 12.vi.1988, L. Stange and J. Wiley (1♀, FSCA); Destin (Okaloosa County), 17.v.1969, H.V. Weems, Jr. (1♀, FSCA); St. Andrews State Park (Bay County), 05–07.v.1987, L. Stange and J. Wiley (2♀, FSCA), 06–07.v.1987, L. Stange and J. Wiley (1♀, 1♂, FSCA); Suwannee River State Park, 13–25.iv.1977, J.R. Wiley (1♂, FSCA); Torreya State Park (Liberty County), 18.v.1970, H.V. Weems, Jr. (1♀, FSCA); Georgia: St. Catherines Island (Liberty County), 24–28.iv.1972, Thompson and Picchi (1♂, AMNH), 10–14.iv.1991, J.G. Rozen, E. Quinter, and A. Sharkov (1♀, AMNH); South Carolina: Hunting Island State Park (Beaufort County), 08.iv.1963, J.G. and B.L. Rozen (1♂, AMNH).

Epeolus inornatus sp. n.

Figs 59, 60, 92G, 93C, 96D, 100B

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. inornatus apart from all other North American Epeolus except E. erigeronis and E. ilicis: the mandible is simple; the axilla does not attain the midlength of the mesoscutellum but the free portion is distinctly hooked, with the tip unattached to the mesoscutellum for more than 1/3 of the entire medial length of the axilla; the pronotal collar and metasomal terga are black; the metasomal terga have rather fine punctures; and the pseudopygidial area of the female is distinctly campanulate with the apex <2 × the medial length and not in contact with two large patches of pale tomentum (one on each side) throughout its length (in contact only at apex, diverging basally). Epeolus inornatus is most similar to E. ilicis, and in both species the mesopleuron has punctures that are similar in size and shiny interspaces that are commonly equal to the puncture diameters. By contrast, in E. erigeronis the punctures are more variable in size, with many smaller punctures among large ones, and most interspaces are narrower such that the surface appears to be very coarsely and densely rugose-punctate. Whereas in E. ilicis the pronotal lobe and legs are more extensively reddish orange than brown or black (at least the anterior surface of the metatibia and metatarsus are the same reddish orange color), the dorsum of the mesosoma and metasoma have gray but also usually some pale yellow short appressed setae, and S4 and S5 of the male have long curved coppery to silvery subapical hairs, in E. inornatus the legs (and sometimes the pronotal lobe and tegula) are usually darker, at least from the metacoxa to metatibia, the dorsum of the mesosoma and metasoma have gray short appressed setae, and S4 and S5 of the male have short straight subapical hairs. Epeolus inornatus is also similar to E. gibbsi, but in E. gibbsi the mandible has a blunt, obtuse preapical tooth; in males S4 and S5 have long curved coppery to silvery subapical hairs, as in E. ilicis and all other Nearctic Epeolus; in females F2 is less than 1.2 × as long as wide (it is more than 1.2 × as long as wide in female E. inornatus); and the pseudopygidial area of the female is in contact with two large patches of pale tomentum (one on each side [the two are parallel to each other]) throughout its length.

Description

FEMALE: Length 8.2 mm; head length 1.9 mm; head width 2.6 mm; fore wing length 5.7 mm.

Integument coloration. Mostly black; notable exceptions as follows: partially to entirely ferruginous on mandible, antenna, pronotal lobe, tegula, and legs. Mandible with apex darker than all but extreme base. Antenna dark brown except F1 reddish brown in part. Pronotal lobe dark brown to black. Tegula pale ferruginous to amber. Wing membrane subhyaline, apically dusky. Legs with brown or black more extensive than reddish orange.

Pubescence. Face with tomentum densest around antennal socket. Tomentum slightly sparser on clypeus; upper paraocular and frontal areas, and vertexal area mostly exposed. Dorsum of mesosoma and metasoma with bands of off-white to pale gray short appressed setae. Mesoscutum with paramedian band. Mesopleuron with upper half hairy, ventrolateral half nearly bare. Metanotum with tomentum uninterrupted except for median bare patch in posterior half, uniformly off white. T1 with median quadrangular black discal patch enclosed by pale tomentum, except for medial separation at apex. T2 with fascia interrupted medially and with faint anterolateral extensions of sparser tomentum. T3 and T4 with fasciae complete. T5 with two large patches of pale tomentum lateral to and contacting pseudopygidial area at apex, diverging from pseudopygidial area basally. T5 with pseudopygidial area campanulate, its apex less than twice as wide as medial length, indicated by silvery setae on impressed disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs not extending beyond apex of sternum by more than 1/4 MOD.

Surface sculpture. Punctures dense. Labrum with larger and sparser punctures (i=1–2d) than clypeus (i<1d). Small impunctate shiny spot lateral to lateral ocellus. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula very densely punctate mesally (i<1d), less so laterally (i=1–2d). Mesopleuron with ventrolateral half densely punctate (i≤1d), the interspaces shining; mesopleuron with punctures similar in size and more or less equally dense throughout. Metasomal terga with punctures very fine, dense (i=1–2d), evenly distributed on disc; the interspaces shining somewhat.

Structure. Mandible without preapical tooth. Labrum with pair of small subapical denticles not preceded by carinae. Frontal keel not strongly raised. Scape with greatest length 1.9 × greatest width. F2 noticeably longer than wide (L/W ratio = 1.4). Preoccipital ridge not joining hypostomal carina, from which it is separated by no less than 1 MOD at its terminal. Mesoscutellum moderately bigibbous. Axilla small to intermediate in size, its lateral margin (L) less than half as long as mesoscutellar width (W) (L/W ratio = 0.4) and tip not extending beyond midlength of mesoscutellum; axilla with tip conspicuously diverging from side of mesoscutellum, distinctly hooked, and axilla with free portion 2/5 its medial length; axilla with lateral margin relatively straight and carinate. Fore wing with three submarginal cells. Pygidial plate apically truncate.

MALE: Description as for female except for usual secondary sexual characters and as follows: F2 shorter, not noticeably longer than wide (L/W ratio = 1.1); pygidial plate apically rounded, with large deep punctures closely clustered.

Figure 59. 

Epeolus inornatus A female holotype, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male allotype, lateral habitus (scale bar 3 mm), and D female paratype axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Etymology

The name is in reference to the grayish pubescence and largely monochromatic dark brown or black integument of this species. From the Latin, “inornatus” (unadorned).

Distribution

Mid-Atlantic states to Texas (Fig. 60).

Figure 60. 

Approximate geographic range of E. inornatus (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: The host species of E. inornatus is/are presently unknown.

FLORAL RECORDS: Labels of examined voucher specimens indicate floral associations with Quercus laevis Walter (Fagaceae) and Vaccinium arboreum Marshall.

Discussion

The specimens from Texas, USA that Brumley (1965) identified as E. ilicis are probably E. inornatus. Although BIN-compliant sequences are presently not available for E. inornatus, a single 421 bp sequence is available for a female specimen (the holotype) from East Texas, which does not cluster with the single sequence (422 bp in length) available for what is herein considered to be the true E. ilicis (a male specimen from Florida, USA) based on its greater resemblance to the holotype of that species (Suppl. material 2). Instead, the sequence from the Florida specimen clusters with sequences of E. zonatus, which is a visibly different bee, and all were assigned the same BIN. Whereas male E. inornatus are unique among Epeolus in having very short straight subapical hairs on S4 and S5 instead of the usual long curved coppery to silvery subapical hairs, females are practically indistinguishable from E. ilicis in terms of surface sculpture and structure. Although consistent, the features (differences in integument coloration and pubescence) that in combination may be used to distinguish female E. inornatus from E. ilicis are subtle. Based on known records, adults of E. inornatus appear to be most active in spring, the same time of year when adults of E. ilicis and E. zonatus are active.

Material studied

Type material. Primary: USA: Texas: Lick Creek Park (College Station, Brazos County), 05–09.iv.2000, M. Buck (holotype ♀ [DEBU, catalog number: 00106728]).

Secondary: USA: Arkansas: Magazine Mountain (Logan County), 23.v.1991, J. Powell (paratype ♀, EMEC); Florida: Liberty County, 24.iv.1961, H.V. Weems, Jr. (paratype ♂, BBSL); Torreya State Park (Liberty County), 12.v.1968, H.V. Weems, Jr. (paratype ♂, FSCA); Georgia: 2 mi SE Blue Ridge (Fannin County), 29.vi.1982, J.B. Whitfield (paratype ♂, EMEC); Rabun Bald (Rabun County), 14.vii.1957, J.G. Chillcott (paratype ♀, CNC); Satolah (Rabun County), 01.vii.1957, J.R. Vockeroth (paratype ♀, CNC), 04.vii.1957, W.R.M. Mason (paratype ♂, CNC); Massachusetts: Amherst, spring 1929, L.A. Carruth (paratype ♂, USNM); North Carolina: Chestnut Bald (Pisgah National Forest, Haywood County), 02.viii.1957, J.G. Chillcott (paratype ♀, CNC); Highlands, 27.vi.1957, W.R.M. Mason (paratype ♀, CNC), 27.vi.1957, J.R. Vockeroth (paratypes 3♂, CNC), 29.vi.1957, J.R. Vockeroth (paratype ♀, CNC), 25.vi.1957, W.R.M. Mason (paratype ♂, CNC); Horse Cove (Highlands), 27.vi.1957, J.R. Vockeroth (paratype ♂, CNC); Wayah Bald (Macon County), 06.vii.1957, W.R.M. Mason (paratype ♀, CNC); Whiteside Mountain (Highlands), 29.vi.1957, W.R.M. Mason (paratype ♀, CNC); South Carolina: Mountain Rest, 14.vi.1957, W.R.M. Mason (paratype ♂, CNC); Texas: 2.5 mi S Delhi (29.7730°N; 97.4020°W) (Caldwell County), 19.iv.2007, J.L. Neff and A. Hook (paratype ♀, CTMI); 8 km SE Elkhart (Anderson County), 27.iv.1985, C.D. Michener (paratype ♂, KUNHM); Brackenridge Field Laboratory (Austin, Travis County), 13.v.1988, A. Hook (paratype ♂, CTMI); Lick Creek Park (College Station, Brazos County), 05–09.iv.2000, M. Buck (allotype ♂ [DEBU, catalog number: 00106727]); Stengl Lost Pines Biological Research Station (30.0800°N; 97.1830°W) (Bastrop County), 13.iv.2006, J.L. Neff (paratype ♀, CTMI).

DNA barcoded material with BIN-compliant sequences

Unavailable.

Non-barcoded material examined

USA: North Carolina: Whiteside Mountain (Macon County), 11.vii.1937, T.B. Mitchell (E. ilicis paratype ♂, NCSU).

Epeolus interruptus Robertson, 1900

Figs 61, 62

Epeolus interruptus Robertson, 1900. Trans. Acad. Sci. St. Louis 10: 55 (♀).

Diagnosis

Unique to E. interruptus among North American species of Epeolus are each of the following morphological features: the metanotum has a blunt median process and T1 has a wide triangular discal patch with concave lateral sides. Epeolus interruptus most closely resembles E. tessieris in that the mesoscutum has short paramedian bands; the axilla does not attain the midlength of the mesoscutellum and like the mesoscutellum is ferruginous (although both are occasionally black in E. interruptus); the mesopleuron commonly has sparser punctures ventrolaterally than in upper half, with the interspaces shining; and T1–T4 have medially-interrupted metasomal fasciae. However, in E. tessieris the metanotum is flat and T1 has a trapezoidal to nearly semicircular discal patch.

Figure 61. 

Epeolus interruptus A female, lateral habitus (scale bar 3 mm) B female, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue arrow indicates blunt median process of metanotum; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Redescription

This species was recently redescribed (Onuferko 2017).

Distribution

Central and western Canada, east of the Rocky Mountains, to northern Mexico (Fig. 62).

Figure 62. 

Approximate geographic range of E. interruptus (orange) based on occurrence records known to the author (yellow circles).

Ecology

See Onuferko (2017) for host and floral records. Floral associations are also indicated in Suppl. material 1, which includes a newly discovered association with Heterotheca villosa (Pursh) Shinners based on the label of one examined voucher specimen.

Discussion

Detailed morphological and taxonomic remarks about this species are given in Onuferko (2017).

Material studied

Type material. Primary: USA: Illinois: Carlinville (Macoupin County), C.A. Robertson (holotype ♀ [INHS, catalog number: 44384]).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ACZ9058. Specimens examined and sequenced.–USA: Arizona: 3♀, 2♂ (PCYU); Utah: 1♂ (BBSL); Virginia: 1♀ (CTMI).

Non-barcoded material examined

Canada: Manitoba: 6♀ (CNC); Ontario: 1♀ (CNC).

Mexico: Baja California: 1♂ (EMEC); San Vicente, 08.vii.1963, J.D. Birchim (2♀, CAS); Baja California Sur: vic.Est. Microondas “Ligüí” (48 km S Loreto), 07.ix.1977, R.R. Snelling (1♀, 1♂, LACM); Nuevo León: Cola de Caballo, 18.vi.1975, H.V. Weems, Jr. (1♂, FSCA).

USA: Arizona: 2♀, 4♂ (AMNH, PCYU); 4.7 mi SE Portal (Cochise County), 03.ix.1978, R.E. Coville (1♀, EMEC); California: Colton, 26–28.v.1917, E.P. Van Duzee (1♂, CAS); Colorado: Boulder, 20.vii.1908, S.A. Rohwer (1♂, CAS); Eldorado Springs, 08.vii.1962, U.N. Lanham (1♂, CUM); Roxborough State Park (39.4356°N; 105.0760°W), 12.vi.2000, A.L. Hicks and V. Scott (1♂, CUM); Idaho: 5 mi E Harvard, 21.vii.1971, R.M. Bohart (1♂, UCBME); Iowa: 1♀ (AMNH); Louisiana: 1♂ (USNM); Michigan: G. H. Gordon Biological Station (44.0470°N; 85.6670°W) (Lake County), 28.vi.2015, J. Gibbs (1♂, JBWM); Nebraska: 1♀ (AMNH); New Mexico: 2♂ (BBSL, FMNH); Texas: 3♀ (AMNH, CTMI); 30 mi N Uvalde (Uvalde County), 21.vi.1983, W.J. Pulawski (1♂, CAS); McAllen Botanical Gardens (McAllen), 1973, C.C. Porter (1♂, FSCA), 20.iii.1976, C.C. Porter (1♀, FSCA); Utah: 1♂ (BBSL).

Epeolus lectoides Robertson, 1901

Figs 63, 64

Epeolus lectoides Robertson, 1901. Can. Entomol. 33: 231 (♀).

Epeolus semilectus Cockerell, 1907a. Entomologist 40: 136 (♂).

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. lectoides apart from all other North American Epeolus except E. glabratus: the axilla is elongate, extending well beyond the midlength of the mesoscutellum but not as far back as its posterior margin, and the free portion is distinctly hooked; the mesopleuron has sparser punctures ventrolaterally (most i>1d) than in upper half, with the interspaces shining; the metasomal terga have minute, shallow punctures; the T2–T4 fasciae are conspicuously narrowed or interrupted medially; and the pseudopygidial area of the female is distinctly campanulate with the apex <2 × the medial length. Whereas in E. glabratus the pronotal collar, axilla, mesoscutellum, and discs of T1 and T2 are ferruginous and the pale pubescence on the metasomal terga are commonly reduced to discrete lateral patches, in E. lectoides the pronotal collar is black, as are sometimes the axilla and mesoscutellum, and the metasomal terga are black and fasciate. Epeolus lectoides is also similar to E. lectus, but in E. lectus the metasomal terga have coarse, deep punctures and the T2–T4 fasciae are complete and evenly broad.

Figure 63. 

Epeolus lectoides A female, lateral habitus (scale bar 3 mm) B female, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Redescription

This species was recently redescribed (Onuferko 2017).

Distribution

Eastern North America (Fig. 64).

Figure 64. 

Approximate geographic range of E. lectoides (orange) based on occurrence records known to the author (yellow circles).

Ecology

See Onuferko (2017) for host and floral records. Floral associations are also indicated in Suppl. material 1, which includes newly discovered associations with Aralia spinosa L. (Araliaceae), Castanea pumila (L.) Mill. (Fagaceae), Helenium amarum (Raf.) H. Rock (Compositae), Helianthella Torr. & A. Gray (Compositae), Helianthus L. (Compositae), Ligustrum L. (Oleaceae), Rudbeckia hirta L. (Compositae), and Vitex L. (Lamiaceae) based on labels of examined voucher specimens.

Discussion

Detailed morphological and taxonomic remarks about this species are given in Onuferko (2017).

Material studied

Type material. Primary: USA: Illinois: Carlinville (Macoupin County), C.A. Robertson (E. lectoides holotype ♀ [INHS, catalog number: 44383]); Virginia: Falls Church, 04.vii.????, N. Banks (E. semilectus holotype ♂ [USNM, catalog number: 534053]).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:AAF2273. Specimens examined and sequenced.–Canada: Ontario: 2♂ (DEBU).

USA: Alabama: Tuskegee National Forest (32.4800°N; 85.6028°W) (Macon County), 24.vii.2016, C.H. Ray (1♀, 1♂, AUMNH); Nebraska: 1♂ (BIML); South Carolina: 1♀, 2♂ (PCYU).

Non-barcoded material examined

Canada: Ontario: 15♀, 23♂ (DEBU, PCYU, ROM); Rondeau Provincial Park (42.2814°N; 81.8427°W) (Beach Access #10, near Visitor Centre), 08.viii.2017, R. Ferrari (1♀, 1♂, PCYU).

USA: Alabama: Auburn University Ornamental Horticulture Research Center (30.7018°N; 88.1454°W), 09.v.2016, Ray, Clem, and Chowdhury (2♂, AUMNH); Auburn (32.5701°N; 85.4603°W) (Lee County), 20.vi.2015, C.H. Ray (2♂, AUMNH); Autauga County (32.4757°N; 86.8597°W), 12.vi.2016, Ray and Chowdhury (2♂, AUMNH); Autauga County (32.3988°N; 86.7918°W), 12.vi.2016, Ray and Chowdhury (1♂, AUMNH); Grand Bay (30.4763°N; 88.3422°W) (Mobile County), 26.v.2010, S. Martin (1♀, AUMNH); Louise Kreher Forest Ecology Preserve (32.6654°N; 85.4845°W), 02.vii.2016, C.H. Ray (1♀, AUMNH); Randolph County (33.1164°N; 85.5435°W), 22.v.2016, C.H. Ray (1♀, AUMNH); Tuskegee National Forest (32.4788°N; 85.5639°W) (Macon County), 28.v.2016, C.H. Ray (2♀, 2♂, AUMNH); Tuskegee National Forest (32.4816°N; 85.6129°W) (Macon County), 13.viii.2016, C.H. Ray (1♀, AUMNH); Tuskegee National Forest (32.4701°N; 85.5840°W) (Macon County), 24.vii.2016, C.H. Ray (1♀, AUMNH); Tuskegee National Forest (32.4800°N; 85.6028°W) (Macon County), 24.vii.2016, C.H. Ray (1♀, 3♂, AUMNH); Florida: Greensboro (Gadsden County), 05.vi.2006, S. Lenberger (1♂, FSCA); Liberty County, 06.vi.2006, S. Lenberger (1♂, FSCA); Shalimar (Okaloosa County), 14.vi.2015, F.W. Eliand, II (1♀, AUMNH); Suwannee River State Park, 24.vi.-14.vii.1977, J.R. Wiley (1♂, FSCA); Torreya State Park (Liberty County), 16.v.1964, H.V. Weems, Jr. (1♀, FSCA); Kansas: 2♂ (USNM); Maryland: 1♀, 1♂ (BIML, DEBU); Michigan: Rose Lake State Wildlife Research Area (42.8075°N; 84.3630°W) (Shiawassee County), 04.vii.2014, J. Gibbs (1♂, JBWM), 13.vii.2014, J. Gibbs (1♂, JBWM); Warren Dunes State Park (41.9030°N; 86.6040°W) (Berrien County), 06.vii.2014, J. Gibbs (1♀, JBWM); New Jersey: 1♀ (BIML); New York: 1♀, 2♂ (AMNH); North Carolina: 2♀ (AMNH); North Dakota: 1♀ (AMNH); 11 mi W Walcott (Richland County), 12.vii.1990, J.R. Powers (1♀, EMEC); 7 mi SE Sheldon (Ransom County), 02.vii.1988, J.R. Powers (1♀, EMEC); South Carolina: 1♀ (BIML); Virginia: 1♀, 2♂ (BIML).

Epeolus lectus Cresson, 1878

Figs 65, 66, 91B, 92A, 93A

Epeolus lectus Cresson, 1878. Trans. Am. Entomol. Soc. 7: 88 (♀).

Epeolus agnatus Cresson, 1878. Trans. Am. Entomol. Soc. 7: 89 (♂).

Diagnosis

The following morphological features in combination can be used to tell E. lectus apart from all other North American Epeolus: the mesopleuron has sparser punctures ventrolaterally (most i>1d) than in upper half, with the interspaces shining; the metasomal terga have coarse, deep punctures; and T2–T4 have complete and evenly broad fasciae. Epeolus lectus is most similar to E. lectoides, and in both species the free portion of the axilla is distinctly hooked and the pseudopygidial area of the female is distinctly campanulate with the apex <2 × the medial length, but in E. lectoides the metasomal terga have minute, shallow punctures and the T2–T4 fasciae are conspicuously narrowed or interrupted medially.

Redescription

FEMALE: Length 9.2 mm; head length 2.3 mm; head width 3.1 mm; fore wing length 7.2 mm.

Integument coloration. Mostly black; notable exceptions as follows: partially to entirely ferruginous on mandible, antenna, tegula, axilla, mesoscutellum, legs, and metasomal sterna. Mandible with apex darker than all but extreme base; preapical tooth lighter than mandibular apex (difficult to see in the E. lectus holotype; described from non-type specimens). Flagellum brown and (except F1) slightly lighter than partially dark brown (otherwise orange) scape and F1 and entirely dark brown pedicel, primarily due to extensive pilosity on flagellum. F2 with orange spot basally. Wing membrane dusky subhyaline, slightly darker at apex. Legs from trochanter to tarsus extensively reddish orange, coxae brown.

Pubescence. Face with tomentum densest around antennal socket. Tomentum slightly sparser on clypeus; upper paraocular and frontal areas, and vertexal area mostly exposed. Dorsum of mesosoma and metasoma with bands of off-white to pale yellow short appressed setae. Mesoscutum with paramedian band. Mesopleuron with upper half sparsely hairy, ventrolateral half nearly bare. Metanotum with tomentum sparser medially, uniformly off white. T1 with discal patch elliptical and very wide, the basal and apical fasciae only narrowly joined laterally. T1 with basal and apical fasciae and T2–T3 with apical fasciae complete (T4 entirely retracted in the E. lectus holotype, but with complete fascia in non-type specimens), T2 with fascia with faint anterolateral extensions of sparser tomentum. T5 with two large patches of pale tomentum lateral to and contacting pseudopygidial area at apex. T5 with pseudopygidial area campanulate, its apex less than twice as wide as medial length, indicated by silvery setae on impressed disc of apicomedial region elevated from rest of tergum. S5 with apical fimbria of coppery to silvery hairs extending beyond apex of sternum by 1/3 MOD.

Surface sculpture. Punctures dense, but those of head and mesosoma sparser in some areas, larger, deep, and distinct. Labrum with larger punctures than clypeus, but punctures of both equally dense (i<1d). Small impunctate shiny spot lateral to lateral ocellus. Mesoscutum, mesoscutellum, and axilla very coarsely and densely punctate; the interspaces shining. Tegula very densely punctate mesally (i≤1d), less so laterally (i=1–2d). Upper half of mesopleuron and anterior margin with denser (i≤1d) punctures than rest of mesopleuron (i>1d), the interspaces shining. Metasomal terga with punctures coarse, dense (i≈1d), evenly distributed on disc; the interspaces shining somewhat.

Structure. Preapical tooth blunt and obtuse. Labrum with pair of small subapical denticles, each preceded by small discrete longitudinal ridge. Frontal keel not strongly raised. Scape with greatest length 1.7 × greatest width. F2 noticeably longer than wide (L/W ratio = 1.5). Preoccipital ridge not joining hypostomal carina, from which it is separated by no less than 1 MOD at its terminal (difficult to see in the E. lectus holotype; described from non-type specimens). Mesoscutellum moderately bigibbous. Axilla large, its lateral margin (L) half as long as mesoscutellar width (W) (L/W ratio = 0.5) and tip not extending much beyond midlength of mesoscutellum (extending to <2/3 its length in the E. lectus holotype and all examined non-type specimens; extending to ~2/3 its length in the E. agnathus holotype); axilla with tip conspicuously diverging from side of mesoscutellum, distinctly hooked, and axilla with free portion approximately half its medial length; axilla with lateral margin relatively straight and carinate. Fore wing with three submarginal cells. Pygidial plate apically truncate.

MALE: Description as for female except for usual secondary sexual characters and as follows: F2 shorter, but still longer than wide (L/W ratio = 1.2); S4 and S5 with much longer coppery to silvery subapical hairs; pygidial plate apically rounded, with large deep punctures closely clustered medially and sparser laterally, with the interspaces shining.

Figure 65. 

Epeolus lectus A female, lateral habitus (scale bar 3 mm) B female holotype, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Distribution

Great Plains and Mountain states east of the Continental Divide (Fig. 66).

Figure 66. 

Approximate geographic range of E. lectus (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: In late July 2015, I collected several specimens of this species near the Poudre River in the Roosevelt National Forest, Colorado, USA, where large numbers of Colletes females were collected and observed foraging on purple Dalea flowers. Using Stephen’s (1954) key, collected specimens were identified as being either C. robertsonii Dalla Torre or C. timberlakei Stephen, the females of which cannot be reliably distinguished morphologically, although the short triangular mesosomal spines and fine punctation on the tegulae of examined specimens coupled with their collection locality suggest they are C. timberlakei.

FLORAL RECORDS: The label of one examined voucher specimen indicates a floral association with Cryptantha cinerea var. jamesii (Torr.) Cronquist (Boraginaceae).

Discussion

The names Epeolus agnathus and E. lectus were published simultaneously, although Cresson (1878) remarked that E. agnathus may be the male of E. lectus as the two specimens are structurally similar. Robertson (1902) synonymized E. agnathus under E. lectus, and separated both specimens from E. lectoides based on differences in metasomal pubescence and punctation (see diagnosis). I have examined the holotype specimens of E. lectus and E. agnathus, and agree with Robertson’s treatment. Although Robertson (1902) did not provide any justification for selecting the name E. lectus over E. agnathus, the holotype of the former is in better condition (that of E. agnathus is missing an antenna) and is female, the sex upon which most Epeolus species descriptions have been based. While Cresson’s Epeolus types include remarkably little collection data, the type locality of E. agnathus (Dakota Territory) is even more vague than that of E. lectus (Kansas).

In contrast to the similar and presumably closely related E. lectoides, E. lectus has a much more restricted range and is rare in collections. Both species are known from the Great Plains, although the range of E. lectus extends further west. In E. lectus, the metasoma has much coarser punctures than that of any other North American species in the genus, including E. lectoides, in which the metasoma has much finer and sparser punctures. In addition to this and other clear morphological differences (see diagnosis), the distinction between E. lectus and E. lectoides is supported by separate BINs for the two species.

Material studied

Type material. Primary: USA: Dakota: H. Ulke (E. agnathus holotype ♂ [ANSP, catalog number: 2226]); Kansas: Wilson (E. lectus holotype ♀ [ANSP, catalog number: 2225]).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ACZ8246. Specimens examined and sequenced.–USA: Colorado: Bellvue (40.6882°N; 105.3070°W) (N Cache La Poudre River and E Gordon Creek, Larimer County), 28.vii.2015, A.T. and T.M. Onuferko (2♀, PCYU).

Non-barcoded material examined

USA: Colorado: Bellvue (40.6882°N; 105.3070°W) (N Cache La Poudre River and E Gordon Creek, Larimer County), 28.vii.2015, A.T. and T.M. Onuferko (3♀, PCYU); Kansas: 4 mi NW Coldwater (Comanche County), 12.vi.2002, G.A. Salsbury (1♀, KUNHM); South Dakota: Chamberlain (Brule County), 15.vi.1928, H.C. Severin (1♂, USNM).

Epeolus mesillae (Cockerell, 1895)

Figs 67, 68, 91D

Phileremus mesillae Cockerell, 1895. Psyche (suppl.) 7: 10 (♂), new neotype designation.

Epeolus mesillae Cockerell, 1934. Am. Mus. Novit. 697: 12.

Epeolus mesillae palmarum Linsley, 1939. Pan-Pac. Entomol. 15: 2 (♀), syn. n.

Diagnosis

The following morphological features in combination can be used to tell E. mesillae apart from all other North American Epeolus: the axilla does not attain the midlength of the mesoscutellum and like the mesoscutellum is black, the fore wing has two submarginal cells, and T1–T4 have complete fasciae. Only in E. americanus and E. asperatus is the fore wing commonly with two submarginal cells, but in both species at least the T1 and T2 apical fasciae are interrupted or at least greatly narrowed medially. Epeolus brumleyi is similar to E. mesillae in axillar structure; in that in females F2 is shorter, as long as wide; and in that T1–T4 have complete fasciae. However, in E. brumleyi the axilla is commonly ferruginous in part and the fore wing has three submarginal cells.

Redescription

MALE: Length 6.6 mm; head length 1.7 mm; head width 2.4 mm; fore wing length 4.9 mm.

Integument coloration. Mostly black; notable exceptions as follows: at least partially ferruginous on mandible, antenna, pronotal lobe, tegula, and legs. Mandible orange between dark brown base and reddish-brown apex; preapical tooth slightly lighter than mandibular apex (difficult to see in the P. mesillae neotype because mandible closed; described from non-type specimens). Flagellum brown, except F1 extensively orange, and slightly lighter than dark brown scape and pedicel. Pronotal lobe reddish brown. Tegula pale ferruginous to amber. Wing membrane hyaline throughout. Legs, except tarsi, with brown or black more extensive than reddish orange.

Pubescence. Face with tomentum densest on clypeus and around antennal socket, sparser on upper paraocular area and vertexal area. Dorsum of mesosoma and metasoma with bands of off-white to pale yellow short appressed setae. Mesoscutum with paramedian band partly obscured by surrounding pale tomentum. Mesopleuron almost entirely obscured by white tomentum, except where rubbed off in the P. mesillae neotype. Metanotum with tomentum uninterrupted, uniformly off white. T1 with discal patch elliptical, narrow, and short. T2–T6 each with complete fascia, those of T2 and T3 somewhat broader laterally, T2 with fascia with anterolateral extensions of sparser tomentum. S3–S5 with long coppery to silvery subapical hairs.

Surface sculpture. Punctures dense. Labrum and clypeus with punctures equally dense (i<1d). Small impunctate spot lateral to lateral ocellus. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula densely punctate mesally (i≤1d), less so laterally (i=1–2d). Mesopleuron with ventrolateral half densely punctate (i<1d) to rugose; mesopleuron with punctures more or less equally dense throughout. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc.

Structure. Labrum with pair of small subapical denticles, each preceded by small discrete longitudinal ridge. Frontal keel not strongly raised. Scape with greatest length 1.7 × greatest width. F2 nearly as long as wide (L/W ratio = 0.9). Preoccipital ridge not joining hypostomal carina, from which it is separated by about 1.5–2 MOD at its terminal (difficult to see in the P. mesillae neotype; described from non-type specimens). Mesoscutellum moderately bigibbous. Axilla small to intermediate in size, its lateral margin (L) less than half as long as mesoscutellar width (W) (L/W ratio = 0.3) and tip not extending beyond midlength of mesoscutellum; axilla with tip visible, but unattached to mesoscutellum for less than 1/3 the medial length of axilla; axilla with lateral margin relatively straight and without carina. Fore wing with two submarginal cells. Pygidial plate apically rounded, with large deep punctures closely clustered.

FEMALE: Description as for male except for usual secondary sexual characters and as follows: F2 slightly longer, as long as wide (L/W ratio = 1.0); wing membrane subhyaline, apically dusky; T5 with large, continuous patch of pale tomentum bordering and separate from pseudopygidial area present only in female; T5 with pseudopygidial area lunate, its apex more than twice as wide as medial length, indicated by silvery setae on impressed disc of apicomedial region elevated from rest of tergum; S3–S5 with much shorter hairs (S5 with apical fimbria of coppery to silvery hairs extending beyond apex of sternum by ~2/5 MOD); pygidial plate apically truncate, with small, denser punctures.

Figure 67. 

Epeolus mesillae A female, lateral habitus (scale bar 3 mm) B female, dorsal habitus (scale bar 3 mm) C male (photo of P. mesillae neotype), lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Distribution

Known to occur in all major North American deserts (Fig. 68).

Figure 68. 

Approximate geographic range of E. mesillae (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: Colletes clypeonitens Swenk is the presumed host of E. mesillae (Hurd and Linsley 1975). Personal observations support such an association. In Whitewater, California, USA, I have collected large numbers of female E. mesillae and male C. clypeonitens in an area dominated by creosote bush (Larrea tridentata (Sessé & Moc. ex DC.) Coville (Zygophyllaceae)) in late March 2016. Only one specimen (a female) of a different species of Colletes (C. larreae Timberlake) was taken at the same locality.

FLORAL RECORDS: Collection records from data contributors to Discover Life (Ascher and Pickering 2017) compiled by J. Pickering indicate the following floral associations: Cryptantha flavoculata (A. Nelson) Payson, Erigeron canus A. Gray, Heterotheca villosa, Larrea tridentata, and Potentilla hippiana Lehm. (Rosaceae). Labels of examined voucher specimens further indicate associations with Baileya pleniradiata Harv. & A. Gray ex A. Gray (Compositae), Chaenactis stevioides Hook. & Arn. (Compositae), Dimorphocarpa wislizeni (Engelm.) Rollins (Brassicaceae), L. glutinosa Engelm., Melilotus Mill., Psoralea lanceolata Pursh (Leguminosae), Prosopis velutina Wooton (Leguminosae), and Tamarix gallica L. (Tamaricaceae).

Discussion

Epeolus mesillae was originally described under the now defunct genus Phileremus because the fore wing in this species has two rather than three submarginal cells, the typical state for most Epeolus species. Among North American Epeolus, E. mesillae exhibits unusual sexual dimorphism in that in females the fore wing and (to a lesser extent) hind wing are apically dusky whereas in males the wings are hyaline throughout. There is some variability in the pubescence on the metasomal terga among specimens, with some exhibiting more grayish-white than yellowish fasciae. Linsley (1939) recognized specimens from southern California as a distinct subspecies (E. mesillae palmarum) based on a larger body size and the presence of pale tomentum interspersed with darker tomentum on the discs of the metasomal terga, especially laterally. Specimens from across the range of this species exhibiting these features have been examined, as well as specimens from southern California in which the metasomal fasciae are clearly distinct from the all-dark discs. Specimens from near the type locality of E. mesillae palmarum were barcoded, and their sequences cluster closely with those from specimens from Southeast Arizona and adjacent Sonora, nearer the type locality (Las Cruces, New Mexico) of E. mesillae mesillae. Hence, I do not consider these to be distinct subspecies, and herein synonymize E. mesillae palmarum under E. mesillae, a change in taxonomic status first proposed by Brumley (1965).

I have not seen the male holotype of P. mesillae and do not know where it is housed, despite personally searching through the entomological collections where T.D. Cockerell deposited the types of other Epeolus species he described. In Brumley (1965), no reference was made to Cockerell’s holotype of P. mesillae, suggesting Brumley too was unable to find it. Moreover, no references in the literature to Cockerell’s type since the species’ original description could be found. In the same publication, another species was described under PhileremusP. verbesinae (now Neolarra verbesinae (Cockerell)) –, which was redescribed by Michener (1939) who indicated that the type was in the T.D.A. Cockerell Collection. It is unclear if either specimen has since ended up in an institution that maintains a research collection, but that the holotype of E. mesillae has not been referenced since its original description strongly suggests it is unlikely to turn up in the future and to all intents and purposes has been lost. In my search for the holotype at the CUM, a male specimen of E. mesillae (labelled as Phileremus mesillae Ckll.) from Mesilla Park (the original type locality) collected by Cockerell from Dimorphocarpa wislizeni on May 7th was discovered. The specimen, which is the property of the CUM, agrees with the original description, and was used to write the present redescription and diagnosis. Given that a synonymy under E. mesillae is proposed herein, it is sensible to have a neotype to serve as a point of reference for any future comparisons. Aside from the collection date, the specimen selected as the neotype of Phileremus mesillae fits the description of the original, which can no longer be traced. Hence, in this particular case the qualifying conditions for designating a neotype as listed under Article 75.3 of the International Commission on Zoological Nomenclature (ICZN) Code (http://iczn.org/iczn/index.jsp) seem to have been met.

Material studied

Type material. Primary: USA: California: Edom (Riverside County), 28.iii.1936, E.G. Linsley (E. mesillae palmarum holotype ♀ [CAS, catalog number: 04789]); New Mexico: Mesilla Park, 07.v.????, T.D. Cockerell (P. mesillae neotype ♂, CUM).

Secondary: USA: California: 1 mi W Edom (Riverside County), 28.iii.1936, E.G. Linsley (E. mesillae palmarum allotype ♂ [CAS, catalog number: 04790]).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:AAF0161. Specimens examined and sequenced.–Mexico: Sonora: 30 km E Agua Prieta (31.3333°N; 109.2403°W), 25.iv.2006, R.L. Minckley (3♀, 1♂, PCYU), 03.v.2005, R.L. Minckley (2♂, PCYU).

USA: Arizona: Douglas R/C Flying Field (31.3430°N; 109.4980°W) (Cochise County), 28.iv.2016, T.M. Onuferko (1♀, PCYU); California: 31 km N Lucerne Valley (34.6840°N; 116.9605°W) (San Bernardino County), 27.iv.2013, Z.M. Portman (1♂, BBSL); Kelso Dunes (34.8940°N; 115.7020°W) (Baker, San Bernardino County), 30.iv.2013, A. Ruttan (1♂, PCYU); Tipton Road (33.9079°N; 116.6510°W) (~1.4 mi SW Whitewater, Riverside County), 26.iii.2016, T.M. Onuferko (1♀, PCYU).

Non-barcoded material examined

Mexico: Baja California: Near La Zapopita Valle de Trinidad, 09–14.iv.1961, F.S. Truxal (2♂, LACM); Baja California Sur: 19 mi SW S. Miguel Comondu, 23.vi.1967, E.L. Sleeper and E.M. Fisher (1♂, LACM); Sonora: 30 km E Agua Prieta (31.3333°N; 109.2403°W), 03.v.2005, R.L. Minckley (1♀, 5♂, PCYU).

USA: Arizona: 11 mi NW Wickenberg, 18.iv.1993, J.G. Rozen (2♀, AMNH); 2 Km W Pima (32.9833°N; 110.2833°W) (Graham County), 25.iv.1996, R.L. Minckley (2♂, PCYU); 2 mi S Willcox (Cochise County), 07.v.1956, E. Ordway (1♀, AMNH); 2.5 mi S Willcox (Cochise County), 24.v.1956, E. Ordway (1♂, AMNH), 07.vi.1956, E. Ordway (1♂, AMNH); 4 mi E Willcox (Cochise County), 08.v.1986, J.G. Rozen (3♀, AMNH), 15.v.1986, J.G. Rozen (1♀, AMNH), 16.v.1986, J.G. Rozen (1♀, AMNH), 17.v.1986, J.G. Rozen (2♀, AMNH); 5 mi NE Douglas (Cochise County), 13.v.1987, J.G. Rozen (1♀, AMNH); Douglas R/C Flying Field (31.3430°N; 109.4980°W) (Cochise County), 23.iv.2016, T.M. Onuferko (2♀, PCYU), 28.iv.2016, T.M. Onuferko (1♀, PCYU); Beaver Dam (36.9028°N; 113.9145°W) (1.7 mi ENE Beaver Dam Wash, Mohave County), 10.v.2014, M.C. Orr (1♀, 1♂, BBSL); Skeleton Canyon Road (Cochise County), 12.v.1977, J.G. Rozen (1♂, AMNH); Southwestern Research Station (5 mi W Portal), 23.iv.1956, E. Ordway (1♀, AMNH); Willcox (Cochise County), 16.v.1985, J.G. Rozen (1♂, AMNH); California: 1 mi W Searchlight Junction (San Bernardino County), 21.iii.1971, R.F. Denno and R.W. Rust (1♂, UCBME); 18 mi W Blythe (Riverside County), 22.iv.1978, R.M. Bohart (1♂, UCBME); 25 mi E Twentynine Palms (34.0806°N; 115.5667°W) (Riverside County), 16.iv.2005, L. Packer (1♂, PCYU); 31 km N Lucerne Valley (34.6840°N; 116.9605°W) (San Bernardino County), 27.iv.2013, Z.M. Portman (1♂, BBSL); Borrego Springs (San Diego County), 31.iii.1973, C. Goodpasture (3♂, UCBME); Borrego Valley (San Diego County), 02.iv.1973, R.M. Bohart (1♂, UCBME); Darwin Falls (Inyo County), 12.v.1974, R.M. Bohart (1♀, UCBME); Goffs (San Bernardino County), 24.iv.1993, J.G. and B.L. Rozen (3♀, AMNH), 06.v.1993, J.G. and B.L. Rozen (1♀, AMNH); Morongo Valley (San Bernardino County), 27.iv.1962, O.C. La France (2♂, AMNH); Thousand Palms (Riverside County), 02.iv.1966, R.O. Schuster (1♂, UCBME); Tipton Road (33.9079°N; 116.6510°W) (~1.4 mi SW Whitewater, Riverside County), 26.iii.2016, T.M. Onuferko (6♀, PCYU); Colorado: Foster Ranch (El Paso County), 21.vi.1978, F.M. Brown (1♂, CUM); Nevada: 1 mi N Crystal (Nye County), 25.v.1999, L. Packer (1♀, PCYU); 2.8 mi E Wadsworth (Washoe County), 30.vi.1963, G.I. Stage (1♀, AMNH); E Las Vegas (36.0983°N; 115.0025°W) (Clark County), 29.iv.2001, A.L. Hicks and V. Scott (1♀, CUM); Overton (Clark County), 09.v.1958, R.C. Bechtel (1♀, AMNH); Sams Camp Wash (Lincoln County), 10.v.-11.vi.1984, R.C. Bechtel and J.B. Knight (1♀, BBSL); New Mexico: 10 mi S Animas (Hidalgo County), 15.v.2013, J.G. Rozen (1♂, AMNH); 15 mi E Animas (Hidalgo County), 15.v.2013, J.G. Rozen (3♂, AMNH); Carlsbad (Eddy County), 20.v.1969, Brothers, Krueger, and Michener (1♂, KUNHM); Road Forks (Hidalgo County), 16.v.2013, J.G. Rozen (2♂, AMNH); Texas: 20 km S Kent (Jeff Davis County), 30.iv.2003, L. Packer and G. Fraser (1♀, PCYU); 7.6 mi S Van Horn (Culberson County), 27.iv.1979, R.R. Snelling (1♂, LACM); Chihuahuan Desert Research Institute (Jeff Davis County), 29.iv.2003, L. Packer and G. Fraser (2♀, PCYU); Utah: Dry Fork (Kane County), 22.v.2000, O. Messinger (1♀, BBSL).

Epeolus minimus (Robertson, 1902)

Figs 69, 70, 101

Triepeolus minimus Robertson, 1902. Entomol. News 13: 81 (♀).

Argyroselenis minima Robertson, 1903. Can. Entomol. 35: 284.

Epeolus beulahensis Cockerell, 1904. Ann. Mag. Nat. Hist. 13: 40 (♀).

Epeolus lutzi Cockerell, 1921. Am. Mus. Novit. 23: 16 (♂).

Epeolus lutzi dimissus Cockerell, 1921. Am. Mus. Novit. 23: 16 (♀).

Epeolus arciferus Cockerell (in Cockerell and Sandhouse, 1924). Proc. Calif. Acad. Sci. (4) 13: 319 (♀).

Epeolus pilatei Cockerell (in Cockerell and Sandhouse, 1924). Proc. Calif. Acad. Sci. (4) 13: 320 (♀).

Epeolus eastwoodae Cockerell, 1937. Pan-Pac. Entomol. 13: 149 (♂).

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. minimus apart from all other North American Epeolus except E. banksi and E. olympiellus: in females, F2 is at least 1.2 × as long as wide; the mesoscutum has distinct, evenly broad paramedian bands that may be joined posteriorly; the axilla is small to intermediate in size, not extending much beyond the midlength of the mesoscutellum (extending to <2/3 its length) but the free portion is more than 1/4 as long as the entire medial length of the axilla, and the axilla (except sometimes the tip) and mesoscutellum are black; the mesopleuron is closely (most i<1d) and evenly punctate; T1 has a quadrangular discal patch, in dorsal view the longitudinal band is at least half as wide as the breadth of the apical fascia; and the T2 fascia has lobe-like anterolateral extensions of tomentum. Whereas in E. banksi the mesoscutum and metasomal terga have bands of gray short appressed setae, in E. minimus the mesoscutum and metasomal terga have bands of off-white to pale yellow short appressed setae. In this respect, E. minimus more closely resembles E. olympiellus, but in E. olympiellus the T3 and T4 fasciae are broken or at least narrowed laterally, as well as medially, whereas in E. minimus the T3 and T4 fasciae are not broken laterally, and are complete or narrowly interrupted medially. Epeolus minimus is also similar to E. axillaris, but in E. axillaris the metanotum has a distinct posteromedial depression (as opposed to being flat) and the axilla is more elongate, extending well beyond the midlength of the mesoscutellum but not as far back as its posterior margin.

Figure 69. 

Epeolus minimus A female, lateral habitus (scale bar 3 mm) B female, dorsal habitus (scale bar 3 mm) C male, lateral habitus (scale bar 3 mm), and D female axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Redescription

This species was recently redescribed (Onuferko 2017).

Distribution

Widely distributed across Canada and the United States, although apparently more common in the west; not known to occur in parts of northeastern North America or the high arctic (Fig. 70). Also, the single (perhaps mislabelled) examined specimen from Florida is an extreme outlier, and given the lack of other examined material from the Southern United States the record should be treated with some skepticism.

Figure 70. 

Approximate geographic range of E. minimus (orange) based on occurrence records known to the author (yellow circles).

Ecology

HOST RECORDS: Graenicher (1906) associated E. minimus (as A. minima) with C. eulophi Robertson based on detailed observations of a female of the former inspecting and entering the nest of a female of the latter in Lake Woods, Wisconsin, USA. However, according to Stephen (1954) Graenicher’s record of C. eulophi in Wisconsin is based on observations of C. kincaidii. Epeolus minimus has been collected with C. kincaidii in Birds Hill Provincial Park and Spruce Woods Provincial Park, Manitoba, Canada where no C. eulophi were collected or observed (J. Gibbs, personal communication, 2017), so the association between E. minimus and C. kincaidii seems likely.

FLORAL RECORDS: See Onuferko (2017). Floral associations are also indicated in Suppl. material 1, which includes newly discovered associations with Ericameria nauseosa var. nauseosa and Medicago L. (Leguminosae) based on labels of examined voucher specimens.

Discussion

In Onuferko (2017), E. minimus is said to be similar to a Californian species yet to be formally recognized, which herein is formally described under the name Epeolus axillaris. Detailed morphological and taxonomic remarks about this species are given in Onuferko (2017). Epeolus minimus is among the most widespread and commonly collected Epeolus species in North America.

Material studied

Type material. Primary: USA: California: Cuyler’s Cove (San Miguel Island), 27.vii.1937, T.D. Cockerell (E. eastwoodae holotype ♂ [CAS, catalog number: 04651]); Pacific Grove (Monterey County), ix.1920, F.E. Blaisdell (E. arciferus holotype ♀ [CAS, catalog number: 01614]); San Pedro, 25.x.1909, G.R. Pilate (E. pilatei holotype ♀ [CAS, catalog number: 01615]); Colorado: Leadville, 03–05.viii.1919 (E. lutzi dimissus holotype ♀ [AMNH, catalog number: 25099]); Walsenburg, 14.vi.1919 (E. lutzi holotype ♂ [AMNH, catalog number: 25098]); Illinois: Carlinville (Macoupin County), C.A. Robertson (T. minimus holotype ♀ [INHS, catalog number: 62276]); New Mexico: Beulah, 11.vii.????, T.D. Cockerell (E. beulahensis holotype ♀ [USNM, catalog number: 534040]).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:AAD3554. Specimens examined and sequenced.–Canada: Alberta: 2♂ (PCYU); British Columbia: Haynes’ Lease Ecological Reserve (49.0930°N; 119.5200°W), 29.vi.-01.vii.2011, G.A. Gielens (1♀, RSKM); Manitoba: Spruce Woods Provincial Park (49.6630°N; 99.2790°W) (Spirit Sands, Division 7), 07.vii.2017, J. Gibbs and Nozoe (1♀, 1♂, JBWM); Ontario: 1♀, 1♂ (PCYU); Saskatchewan: 3♂ (PCYU); Saskatchewan Landing Provincial Park (50.6950°N; 107.9030°W), 03.vii.2013, A. Fortney and M. Anderson (1♂, RSKM); Yukon: 1♂ (RSKM).

USA: California: 1♂ (EMEC); Colorado: 2♀ (PCYU); Morrison (39.6677°N; 105.1968°W) (SE Red Rocks Amphitheatre), 16.vi.2017, T.M. Onuferko (1♂, PCYU); Idaho: 2♀, 1♂ (BBSL, PCYU); Utah: 1♀ (BBSL).

Non-barcoded material examined

Mexico: Baja California: San Vicente, 08.vii.1963, J. Powell (3♂, EMEC).

Canada: Alberta: 15♀, 10♂ (CNC); British Columbia: 16♀, 1♂ (CNC, ROM); Manitoba: 4♀, 7♂ (CNC, DEBU, ROM); Birds Hill Provincial Park (50.0100°N; 96.9100°W) (Division 12), 15.vii.2017, J. Gibbs and Nozoe (1♂, JBWM); Erickson, 03.viii.1983, D.H. Pengelly (1♀, JBWM); Fort Whyte (Winnipeg), 20.vi.1991, B.G. Elliot (1♂, JBWM), 13.vii.1991, B.G. Elliot (1♂, JBWM); Portage la Prairie, 29.vi.1976, T.D. Galloway (1♀, JBWM); Spruce Woods Provincial Park (49.6630°N; 99.2790°W) (Spirit Sands, Division 7), 07.vii.2017, J. Gibbs and Nozoe (1♂, JBWM); Winnipeg, 21.vi.1979, T.D. Galloway (1♂, JBWM), 06.vii.1991, B.G. Elliot (1♂, JBWM); Winnipeg Beach, 15.vii.1989, T.D. Galloway (1♀, JBWM), 15.vii.1989, T.D. Galloway (2♂, JBWM); Northwest Territories: 7♀ (CNC); Ontario: 9♀, 6♂ (CNC, PCYU, ROM); Caledon (Forks of the Credit Provincial Park), 25.vii.1968, P. MacKay (1♂, PCYU); Quebec: 3♂ (CNC); Saskatchewan: 9♀, 10♂ (CNC, PCYU); Borden Bridge, 01.viii.1976, T.D. Galloway (1♀, JBWM); Ernfold, 05.vii.1984, T.D. Galloway (1♀, JBWM); Sands Hills (7 km W Piapot), 26.vii.2003, D. Larson (1♂, PCYU); Yukon: 2♀ (CNC).

USA: California: 4♀, 4♂ (EMEC, UCR); 2 mi S Asilomar (Monterey County), 26.ix.1959, C.W. O’Brien (1♂, AMNH); Antioch (Contra Costa County), 20.ix.1958, J.R. Powers (1♀, AMNH), 28.viii.1976, N.J. Smith (1♀, UCBME); Bodega Head (Sonoma County), 14.v.1977, W.M. Oldham (1♂, UCBME); Carnelian Bay (Lake Tahoe), 29.vii.1962, R.M. Bohart (1♀, UCBME); Carson Pass (Alpine County), 13.vii.1966, R.M. Bohart (1♂, UCBME), 16.vii.1968, R.M. Bohart (1♂, UCBME), 16.vii.1968, W.W. Harberts (1♂, UCBME); Chipmunk Flat (Tuolumne County), 09.viii.1960, C.A. Toschi (1♀, AMNH); Dune Lakes (3 mi S Oceano, San Luis Obispo County), 01.vi.1972, J. Powell (1♂, EMEC), 03–04.x.1972, J. Powell (18♂, EMEC), 07.vi.1973, J. Powell (1♂, EMEC), 11.vii.1973, R. Coville (1♂, EMEC), 12.vii.1973, J. Powell (2♂, EMEC), 02.v.1974, J. Powell (1♂, EMEC); Holcomb Valley (San Bernardino County) (1♂, BBSL); Hot Creek (8 air mi E Mammoth Lakes, Mono County), 24.viii.1977, J. Powell (2♂, EMEC); Inglenook Fen (5 mi N Fort Bragg, Mendocino County), 27.v.1976, R. Coville (1♀, 3♂, EMEC); Inglenook Fen (Mendocino County), 22.vii.1972, E.I. Schlinger (1♂, EMEC); Lanphere-Christensen Dunes Preserve (4 mi W Arcata, Humboldt County), 26.vii.1975, M.E. Buegler and E.I. Schlinger (1♂, EMEC); Lobos Creek (San Francisco County), 10.v.1979, J. Powell (1♀, 3♂, EMEC), 15.vi.1960, G.I. Stage (1♀, AMNH); Mad River Beach (Humboldt County), 26.vi.1969, J. Powell (1♀, EMEC); McClures Beach (Marin County), 27.vi.1969, R.W. Thorp (1♀, UCBME); North Beach (Point Reyes National Seashore, Marin County), 10.v.1980, K. Standow (1♀, EMEC), 30.viii.1974, P.A. Opler (2♀, 1♂, EMEC); North Fork, Del Puerto Creek (Del Puerto Canyon, Stanislaus County), 25.v.1974, E. Schlinger (1♂, EMEC); Point Reyes National Seashore (Marin County), 03.iii.1968, R.W. Thorp (1♀, UCBME), 23.vii.1974, P.A. Opler (1♀, EMEC); San Bruno Mountain (San Mateo County), 23.v.1961, G.I. Stage (1♀, AMNH), 23.v.1961 (1♀, AMNH), 23.viii.1960, G.I. Stage (1♂, AMNH); San Francisco Bay Salt Marshes, viii.1907?, Thompson (3♂, EMEC); San Francisco Sand Dunes, 25.vi.1954, J.G. Rozen (1♂, EMEC), 25.vi.1954, P.D. Hurd (1♀, EMEC); Santa Cruz Island (Christi Beach, Santa Barbara County), 23.ix.1968, R.W. Thorp (1♀, UCBME); Sierra Valley (Sierra County), 06.vii.1972, R.M. Bohart (1♀, UCBME); Simonton Cove (San Miguel Island, Santa Barbara County), 11.vii.1970, A.A. Grigarick and R.C. Schuster (1♀, UCBME); Toms Place (Mono County), 01.ix.1965, A.J. Slater (1♀, EMEC); Yuba Pass (Sierra County), 11.viii.1978, R.M. Bohart (1♂, KUNHM); Colorado: Rock Creek Park (Colorado Springs), 19.viii.1937 (1♀, 1♂, AMNH); Florida: 1♀ (PCYU); Idaho: 1♀ (PCYU); Daniels Reservoir (Oneida County), 11.vii.1997, F.D. Parker (3♂, BBSL); Illinois: 1♂ (FMNH); Minnesota: Detroit, 26.viii.1924, O.A. Stevens (1♀, AMNH); Montana: 1♀ (KUNHM); 11 mi SE Ennis (Madison County), 18.viii.1966, D.R. Miller (1♀, UCBME); Nebraska: Cedar Point Biological Station (8 mi N Ogallala, Keith County), 11–18.vii.1988, J.G. Rozen and E. Quinter (1♀, AMNH); Fort Robinson (Dawes County), 11.viii.1971, J.G., B.L., and K.C. Rozen (3♀, AMNH), 12.viii.1971, J.G., B.L., and K.C. Rozen (2♀, AMNH), 09–11.viii.1972, J.G. Rozen, K.C. Rozen, and R. McGinley (2♀, 4♂, AMNH); Warbonnet Canyon (Sioux County), 24.vii.1968, R.R. Snelling (1♂, LACM); Nevada: Fallon, 01.vi.1930, E.L. Bell (1♀, AMNH), 06.vi.1930, E.L. Bell (1♀, AMNH), 10.vi.1930, E.L. Bell (1♀, AMNH); Mount Rose Summit (Washoe County), 09.vii.1964, R.M. Bohart (1♀, UCBME); New Mexico: Santa Fe, 09.vi.1931, F.E. Lutz (1♀, 1♂, AMNH); North Dakota: Gascoyne, 19.vi.1918, O.A. Stevens (1♀, AMNH); Jamestown, 16.viii.1913, O.A. Stevens (1♀, AMNH); Marmarth, 04.vii.1949, O.A. Stevens (3♂, AMNH); McKenzie, 05.viii.1913, O.A. Stevens (1♀, AMNH); Monango, 03.vii.1913, O.A. Stevens (1♀, AMNH); Pleasant Lake, 11.viii.1913, O.A. Stevens (1♀, AMNH); Washburn, 23.vii.1926, O.A. Stevens (3♀, 2♂, AMNH); Williston, 09.viii.1915, O.A. Stevens (1♂, AMNH); Oregon: 1♂ (KUNHM); South Dakota: 1♀ (BIML); Utah: Indian Canyon (Duchesne County), 18.vii.1965, G.F. Knowlton (1♀, UCBME); NE Ruby’s Inn (Garfield County), 17.viii.1995, V.J. Tepedino and F.D. Parker (2♀, BBSL).

Epeolus nebulosus sp. n.

Figs 71, 72, 99A

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. nebulosus apart from all other North American Epeolus except E. basili, E. novomexicanus, and E. pusillus: the axilla is large, with the tip extending well beyond the midlength of the mesoscutellum but at most to the band of pale tomentum along its posterior margin, dilated laterally, and ferruginous to some degree whereas the mesoscutellum is typically all black; the axilla’s free portion is clearly less than 2/5 as long as its entire medial length; the mesopleuron is closely (most i<1d) and evenly punctate, that of the female is obscured by white tomentum only in the upper half (with a large, sparsely hairy circle occupying much of the ventrolateral half) whereas that of the male (excluding the hypoepimeral area) is entirely obscured by white tomentum; T2–T4 have complete and evenly broad fasciae; the T2 fascia has lobe-like anterolateral extensions of tomentum; and the pseudopygidial area of the female is lunate and wider than long (the apex ≤2 × the medial length). Epeolus basili, E. nebulosus, E. novomexicanus, and E. pusillus are all extremely similar to one another. Epeolus nebulosus is most similar to E. novomexicanus, but in E. novomexicanus the mesoscutum usually has distinct paramedian bands and at least the integument beneath the T1 apical fascia is ferruginous, as are sometimes the rest of the tergum and other terga, whereas in E. nebulosus the mesoscutum is entirely obscured by pale tomentum and the metasomal terga (excluding the brown translucent apical margins) are entirely black. In E. basili the metasomal terga are also ferruginous to some degree, and the T2 and T3 (for female) or T2–T4 (for male) fasciae are narrowed medially and removed from the apical margin (in E. nebulosus the T2–T4 fasciae are on or very little removed from the apical margin), and the pseudopygidial area of the female is ≥2 × the medial length. Whereas in E. pusillus the flagellum, except sometimes F1, and metasomal sterna are consistently brown or black and clearly not the same reddish-orange color as the legs (tibiae to tarsi), in E. nebulosus the flagellum, at least ventrally, is the same reddish-orange color as the legs (tibiae to tarsi) as are usually the metasomal sterna. Epeolus nebulosus is also similar to E. scutellaris in that the axilla is large, with the lateral margin arcuate, and that the apical fasciae are complete. However, in E. scutellaris the pseudopygidial area of the female is much wider (the apex ~2.5–3 × the medial length) than in E. nebulosus, and the mesopleuron of both the female and male is obscured by white tomentum only in the upper half (with a large, sparsely hairy circle occupying much of the ventrolateral half).

Description

MALE: Length 7.2 mm; head length 2.0 mm; head width 2.7 mm; fore wing length 5.5 mm.

Integument coloration. Mostly black; notable exceptions as follows: at least partially ferruginous on mandible, antenna, pronotal lobe, tegula, axilla, legs, pygidial plate, and metasomal sterna. Mandible with apex darker than rest of mandible; preapical tooth slightly lighter than mandibular apex. Antenna brown and orange in part. Pronotal lobe and tegula pale ferruginous to amber. Wing membrane subhyaline, apically dusky. Legs more extensively reddish orange than brown or black. S1–S6 reddish orange.

Pubescence. Face with tomentum densest on clypeus and around antennal socket, slightly sparser on upper paraocular area and vertexal area. Dorsum of mesosoma and metasoma with bands of off-white to pale yellow short appressed setae. Mesoscutum largely obscured by pale tomentum. Mesopleuron (excluding hypoepimeral area) entirely obscured by white tomentum. Metanotum with tomentum uninterrupted, uniformly off white. T1 with narrow and short discal patch largely obscured by pale tomentum. T2–T6 each with complete fascia, T2 with fascia with wide basomedially convergent anterolateral extensions of tomentum. S4 and S5 with long coppery to silvery subapical hairs, which individually are often darker apically.

Surface sculpture. Punctures dense. Labrum with larger and sparser punctures (i=1–2d) than clypeus (i<1d) (difficult to see in holotype because clypeus entirely obscured by tomentum; described from paratypes with hair removed). Small impunctate shiny spot lateral to lateral ocellus. Mesoscutum, mesoscutellum, and axilla coarsely and densely rugose-punctate. Tegula densely punctate mesally (i≤1d), less so laterally (i=1–2d). Mesopleuron with ventrolateral half densely punctate (i<1d) to rugose; mesopleuron with punctures more or less equally dense throughout (not visible in holotype because mesopleuron entirely obscured by tomentum; described from paratypes). Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc.

Structure. Preapical tooth obtuse. Labrum with pair of small subapical denticles not preceded by carinae (difficult to see in holotype; described from paratypes). Frontal keel not strongly raised. Scape with greatest length 2.0 × greatest width. F2 noticeably longer than wide (L/W ratio = 1.2). Preoccipital ridge not joining hypostomal carina, from which it is separated by about 1.5 MOD at its terminal. Mesoscutellum weakly bigibbous. Axilla large, its lateral margin (L) half as long as mesoscutellar width (W) (L/W ratio = 0.5) and tip extending well beyond midlength of mesoscutellum but not as far back as its posterior margin; axilla with tip clearly visible, but unattached to mesoscutellum for less than 2/5 the medial length of axilla; axilla with lateral margin arcuate. Fore wing with three submarginal cells. Pygidial plate apically rounded, with large deep, well-separated punctures, with the interspaces shining.

FEMALE: Description as for male except for usual secondary sexual characters and as follows: F2 even longer than wide (L/W ratio = 1.5); mesopleuron densely hairy, except for two almost entirely bare patches (one beneath base of fore wing (hypoepimeral area), a larger circular patch occupying much of ventrolateral half of mesopleuron); T5 with large, continuous patch of pale tomentum bordering and contacting pseudopygidial area present only in female; T5 with pseudopygidial area lunate, its apex twice as wide as medial length, indicated by silvery setae on disc of apicomedial region elevated from rest of tergum; S4 and S5 with much shorter hairs (S5 with apical fimbria of coppery to silvery hairs not extending beyond apex of sternum by much more than 1/4 MOD); pygidial plate apically truncate, with small, denser punctures.

Figure 71. 

Epeolus nebulosus A female allotype, lateral habitus (scale bar 3 mm) B female allotype, dorsal habitus (scale bar 3 mm) C male holotype, lateral habitus (scale bar 3 mm), and D female allotype axillae and mesoscutellum, dorsal view (scale bar 0.5 mm; blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length).

Etymology

The name is in reference to the pale tomentum obscuring much of the integument of this species. From the Latin, “nebulosus” (hazy).

Distribution

California and probably western Nevada (Fig. 72).

Figure 72. 

Occurrence records of E. nebulosus known to the author (yellow circles).

Ecology

HOST RECORDS: The host species of E. nebulosus is/are presently unknown.

FLORAL RECORDS: Labels of examined voucher specimens indicate a floral association with Ericameria nauseosa.

Discussion

Epeolus nebulosus is a cryptic species within the “pusillus group” that closely resembles some specimens of E. novomexicanus, and the ranges of the two species overlap to some extent. The morphological differences (in integument coloration and patterns of pubescence) among the four members of the “pusillus group” are subtle. The status of E. nebulosus as a separate species is further supported by a separate BIN and large barcode sequence divergence (>3.2%) from its nearest neighbor, E. novomexicanus. Although most species of Epeolus were described from a female name-bearing type, a male specimen is designated as the holotype of E. nebulosus because a barcode-compliant sequence is associated with it and because much of the pubescence is discolored or rubbed off in the available female specimen, which is herein designated as the allotype. Since this species is described from very few specimens, efforts should be made to collect additional representatives of E. nebulosus for DNA barcoding to determine if the morphological differences between it and E. novomexicanus reported here are consistent.

Material studied

Type material. Primary: USA: California: Gilbert Pass on Hwy 168 (37.4305°N; 117.9388°W) (N Deep Springs Valley, Inyo County), 14.ix.2013, M.C. Orr (holotype ♂ [CCDB-28239 F01], BBSL).

Secondary: USA: California: 3.2 km S Pearblossom (Los Angeles County), 07.xi.1977, R.R. Snelling (allotype ♀, LACM); Gilbert Pass on Hwy 168 (37.4305°N; 117.9388°W) (N Deep Springs Valley, Inyo County), 14.ix.2013, M.C. Orr (paratypes 2♂, BBSL).

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ACZ0767. See Type material for specimens examined and sequenced (indicated by unique CCDB-plate and well number).

Epeolus novomexicanus Cockerell, 1912

Figs 73, 74, 97E, 99B

Epeolus novomexicanus Cockerell, 1912. Ann. Mag. Nat. Hist. (8) 10: 487 (♂).

Diagnosis

The following morphological features in combination (excluding any that are specific to the opposite sex of the one being diagnosed) can be used to tell E. novomexicanus apart from all other North American Epeolus except E. basili, E. nebulosus, and E. pusillus: the axilla is large, with the tip extending well beyond the midlength of the mesoscutellum but at most to the band of pale tomentum along its posterior margin, dilated laterally, and ferruginous to some degree whereas the mesoscutellum is typically all black; the axilla’s free portion is clearly less than 2/5 as long as its entire medial length; the mesopleuron is closely (most i<1d) and evenly punctate, that of the female is obscured by white tomentum only in the upper half (with a large, sparsely hairy circle occupying much of the ventrolateral half) whereas that of the male (excluding the hypoepimeral area) is entirely obscured by white tomentum; T2–T4 have complete and evenly broad fasciae; the T2 fascia has lobe-like anterolateral extensions of tomentum; and the pseudopygidial area of the female is lunate and wider than long (the apex ≤2 × the medial length). Epeolus basili, E. nebulosus, E. novomexicanus, and E. pusillus are all extremely similar to one another. Epeolus novomexicanus is most similar to E. nebulosus, but in E. nebulosus the mesoscutum is entirely obscured by pale tomentum and the metasomal terga (excluding the brown translucent apical margins) are entirely black whereas in E. novomexicanus the mesoscutum usually has distinct paramedian bands and at least the integument beneath the T1 apical fascia is ferruginous, as are sometimes the rest of the tergum and other terga. In E. basili the metasomal terga are also ferruginous to some degree, but the T2 and T3 (for female) or T2–T4 (for male) fasciae are narrowed medially and removed from the apical margin (in E. novomexicanus the T2–T4 fasciae are on or very little removed from the apical margin), and the pseudopygidial area of the female is ≥2 × the medial length. Whereas in E. pusillus the flagellum, except sometimes F1, and metasomal sterna are consistently brown or black and clearly not the same reddish-orange color as the legs (tibiae to tarsi), in E. novomexicanus the flagellum, at least ventrally, is the same reddish-orange color as the legs (tibiae to tarsi) as are usually the metasomal sterna. Epeolus novomexicanus is also similar to E. scutellaris in that t