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).

(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).

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).

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

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.

This species was recently redescribed (Onuferko 2017).

Great Plains to southwestern Ontario (Fig. 5).

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.

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

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

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).

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).

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.

This species was recently redescribed (Onuferko 2017).

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).

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.

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

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]).

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

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

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).

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.

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.

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.

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

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

FLORAL RECORDS: Unknown.

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%).

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).

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

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.

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.

Central and southern California (Fig. 11).

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.

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. asperatusholotype, 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.

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]).

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).

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).

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.

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.

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.

New Mexico and southern Colorado (Fig. 13).

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

FLORAL RECORDS: Unknown.

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.

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).

Unavailable.

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).

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.

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).

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.

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.

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).

Unavailable.

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).

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.

This species was recently redescribed (Onuferko 2017).

Eastern North America (Fig. 17).

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

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

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]).

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

USA: New York: 1♀ (AMNH).

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 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.

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.

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.

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).

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.

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.

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).

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

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).

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.

Maryland to North Carolina (Fig. 21).

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).

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.

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).

Unavailable.

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).

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.

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.

Arizona to west Texas (Fig. 23).

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.

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. asperatushas 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.

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

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).

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).

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).

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.

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

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

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).

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.

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).

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

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.

This species was recently redescribed (Onuferko 2017).

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

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.

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.

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

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

USA: Florida: 1♂ (FSCA).

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).

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).

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.

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.

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).

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).

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.

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).

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

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.

This species was recently redescribed (Onuferko 2017).

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

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.

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

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]).

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).

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).

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.

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.

South Atlantic states (Fig. 33).

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).

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.

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).

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).

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 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.

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.

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

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

HOST RECORDS: The female PCYU paratype collected by H.T. Ngo (see Material studied) is labelled with the same collection information as three Colletesspecimens (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).

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.

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).

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

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.

This species was recently redescribed (Onuferko 2017).

Western North America (Fig. 37).

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

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.

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).

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).

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).

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.

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.

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.

Florida and coastal Georgia (Fig. 40).

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).

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.

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).

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

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.

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.

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).

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

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).

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.

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).

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

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.

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.

South Atlantic states (Fig. 44).

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).

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).

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).

Unavailable.

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).

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.

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.