Both sexes of Agapostemon are diagnosed by the complete carina on the rear face of the propodeum (Fig. 2A, B). Other metallic green Halictinae genera, such as Augochloropsis, can have a pair of lateral carinae (e.g., Fig. 2C), but these are well-separated dorsally and never forming a complete carina as in Agapostemon. Females can be further recognized by having the hind tibial spurs with broad teeth. Males can be further recognized by having the metasoma black and yellow striped rather than metallic green and by having the basitarsus fused with the next tarsal segment (Fig. 2E).

Figure 2. 

Characters to separate Agapostemon from Augochlorini: Complete raised carina on the rear face of the propodeum in Agapostemon A female and B male. Incomplete carina in C Augochloropsis female and D Augochlorella male E Agapostemon male hind leg with first two tarsomeres fused (red arrow pointing to point of fusion).

Figure 3. 

Characters to separate Augochlora and Augochlorella A S1 of Augochlora pura female with a median keel indicated by red arrow B Augochlora pura male rear propodeum showing distinct punctures C Augochlora pura female face with protruding paraocular lobe indicated by red arrow D Augochlorella aurata female face with undeveloped paraocular lobe indicated by red arrow E Augochlora pura male face with protruding paraocular lobe indicated by red arrow F Augochlorella aurata male with undeveloped paraocular lobe indicated by red arrow. Scale bars: 1 mm.

Females of Agapostemon angelicus can be recognized by the unique double-punctate scutum (as in Fig. 5A), a character they share with Agapostemon texanus. Our efforts to find characters to separate the females of these two species for the most part have been fruitless, and like workers before us (Roberts 1972), we consider females of Agapostemon angelicus and Agapostemon texanus to be morphologically indistinguishable. Males of Agapostemon angelicus can be separated from males of Agapostemon texanus using the leg and genitalia characters given in the key.

Agapostemon angelicus is primarily a western species, though Roberts (1972) records it from central Iowa and extreme eastern Kansas. As far as we have been able to determine, there are no recent records of Agapostemon angelicus east of the 98^th Meridian. A 2018–2019 statewide survey of Kansas bees by Morphew (2017) did not find any Agapostemon angelicus males (verified by MSA) east of the central part of the state, with the easternmost records from Ellsworth and Rice counties. Recent survey efforts in eastern Nebraska have not found any Agapostemon angelicus east of the Grand Island area (Hall Co.). No confirmed recent records are known from Minnesota, Missouri, or Iowa, despite extensive collection efforts in those states. However, care should be taken to look for it and more work needs to be done to confirm the eastern extent of the range of Agapostemon angelicus. In areas where the two species may overlap, it is recommended that females identified morphologically as Agapostemon angelicus or Agapostemon texanus be treated as a single morphospecies. The two species can also be separated by DNA barcodes.¹

Female Agapostemon melliventris can be recognized by having the apex of the clypeus yellow as well as their non-metallic, light-colored metasoma. The terga are generally amber-colored but can be dark enough (e.g., Fig. 4B) to resemble Agapostemon virescens.

Male Agapostemon melliventris can be recognized by having the metasoma primarily yellow (Fig. 9A), with just thin dark bands, and they also have the hind femur much skinnier (Fig. 8A) than any of the other species treated here.

Agapostemon melliventris is not known from the midwestern US, though Roberts (1972) records if from eastern Nebraska and Kansas, so there is the potential for it to be found in Missouri and Iowa. We are not aware of any recent collections east of the 98^th Meridian.

The female of Agapostemon sericeus can be recognized by the combination of the metallic green metasoma (as in Fig. 4C) and the reticulate sculpturing of the scutum (Figs 5B, 6A). It is most similar to Agapostemon splendens, but Agapostemon splendens has the scutum more punctured (Fig. 6B) rather than reticulate, and Agapostemon sericeus can be further distinguished by its sharply angled dorsolateral ridge of the pronotum (Fig. 5B) and by having the ventral pleural tubercle flush with the plate (Fig. 6C).

Male Agapostemon sericeus have S3 and S4 with a low transverse swelling and generally have distinct yellow marks on the apical sterna (Fig. 7C). They are most similar to males of Agapostemon texanus but can be distinguished by the relative lengths of F1 and F2: in Agapostemon sericeus F1 is slightly more than half the length of F2, whereas in Agapostemon texanus F1 is about three-fourths the length of F2 (Fig. 10). They can also be separated by the genitalia (see Roberts 1972).

Agapostemon sericeus was previously known as Agapostemon radiatus (Say) (e.g., Mitchell 1960; Roberts 1972) but was synonymized by Day and Fitton (1977).

Females of Agapostemon femoratus Crawford, primarily a western species not recorded east of New Mexico, Colorado and Wyoming by Roberts (1972), are essentially identical to females of Agapostemon sericeus, though the males are quite distinct, possessing a grossly enlarged hind femur, its width and length equal or nearly so. The key in Roberts (1972) indicates the scutum of female Agapostemon sericeus is more distinctly punctate than Agapostemon femoratus, but we do not consider this a reliable separating character. Curiously, there are several Missouri records of Agapostemon femoratus from the 1960s identified by Roberts in separate online databases, (discoverlife.org, Ascher and Pickering 2022), but these were not included in his 1972 revision. We have not seen these specimens, but assume they represent mis-determined females of Agapostemon sericeus, not Agapostemon femoratus.

The female of Agapostemon splendens can be recognized by the combination of the metallic green metasoma (as in Fig. 4C) and the coarsely punctured sculpturing of the scutum (Figs 5C, 6B). It is similar to Agapostemon sericeus, but that species has the sculpturing of the scutum more reticulate (Figs 4B, 6A). Agapostemon splendens can be further distinguished by the obtuse dorsolateral ridge (Fig. 5C), the upraised ventral pleural tubercle (Fig. 6D), and it is generally larger than Agapostemon sericeus (though their sizes can intergrade).

Male Agapostemon splendens can be recognized from all other midwestern Agapostemon by their very enlarged hind femur, with the length twice the width (Fig. 8C). They also tend to be larger than related species and have darker wings, but this character is subtle.

Some previous works (e.g., Mitchell (1960) and the keys on discoverlife.org) have used the shape of the ridges of the propodeal triangle (which often form a depressed medial triangle) to separate female Agapostemon splendens from Agapostemon sericeus (which have parallel striae throughout the propodeal triangle), but we have found the character variable and it can be quite subtle, particularly in smaller Agapostemon splendens. Agapostemon splendens is largely restricted to areas of deep sands. We have examined material from throughout the range of Agapostemon splendens, and there are many individuals, especially in the southern US, that have the scutal sculpturing more reticulate, similar to Agapostemon sericeus. More work is needed to determine whether this represents normal variation or is potentially due to a cryptic species complex.

The females of Agapostemon texanus have the metasoma metallic green (Fig. 4C) and can be recognized by the “double-punctured” scutum, which has a combination of intermixed large and small punctures (Fig. 5A). Females cannot be reliably distinguished from Agapostemon angelicus, so they should be separated based on range or DNA barcodes (see remarks for Agapostemon angelicus above).

Male Agapostemon texanus have S3 and S4 with a low transverse swelling and generally have distinct yellow marks on the apical sterna (Fig. 7B, D). They are extremely similar to Agapostemon angelicus, but Agapostemon texanus have the hind tibia with black stripes on the front and back (Fig. 8D), whereas Agapostemon angelicus has the hind tibia yellow anteriorly (Fig. 11). In addition, the two species can be separated based on the genitalia characters given in the key (Fig. 12), and at least in the midwestern US, the range of the two species largely does not overlap.

Male Agapostemon texanus are also similar to (and frequently misidentified as) Agapostemon sericeus but can be distinguished based on the relative lengths of F1 and F2: Agapostemon texanus has F1 about three-fourths the length of F2 (Fig. 10A), whereas Agapostemon sericeus has F1 slightly more than half the length of F2 (Fig. 10B).

Agapostemon texanus and Agapostemon angelicus largely do not overlap in range in the midwestern region, though Roberts (1972) reports Agapostemon angelicus from Iowa and eastern Kansas (see remarks under Agapostemon angelicus, above).

Females of Agapostemon virescens are the only midwestern species that has the metasoma dark (Fig. 4A), rather than metallic green (but see comments on dark Agapostemon melliventris).

Males of Agapostemon virescens can be recognized by the lack of a transverse swelling on S4 (Fig. 7A). In addition, S5 and S6 are usually all dark (Fig. 7A), whereas those sterna usually (though not always) have at least some yellow in other Agapostemon species. Finally, Agapostemon virescens males have a relatively slender hind femur compared to most other midwestern Agapostemon species (see Fig. 8E).

Two western species with females with non-metallic metasomas have been recorded from nearby states though they have not been recorded from Iowa, Minnesota, or Missouri. Agapostemon melliventris has been found as far east as eastern Kansas and Nebraska, but they have the metasoma lighter and the apex of the clypeus yellow, compared to black in Agapostemon virescens females. In addition, Agapostemon coloradinus (Vachal) is a Great Plains species which occurs as far east as eastern Kansas, though Agapostemon coloradinus is usually noticeably larger than Agapostemon virescens with finer, closer striations on the hypostomal area on the underside of the head (see Roberts 1972). Males of Agapostemon coloradinus are similar to males of Agapostemon virescens, but Agapostemon coloradinus males have a dark stripe on the posterior surface of the hind femur and the inner gonostylar flap of Agapostemon coloradinus lacks a pronounced, medially-directed process basally, which is present in Agapostemon virescens.

The genus Augochlorella can be recognized by the combination of a normal oval-shaped tegula (as in Fig. 1B), the incomplete carina on the rear face of the propodeum (Fig. 2D), and the lack of a protruding paraocular lobe (Fig. 3D, F). Females lack the keel on S1 seen in Augochlora and they have simple hind tibial spurs. Males are quite similar to Augochlora, but Augochlorella males have the S4 apical margin weakly to strongly concave, versus straight in Augochlora. In addition, Augochlorella males lack distinct punctures on the rear of the propodeum (Fig. 2D) compared to Augochlora males which do have distinct punctures (Fig. 3B). Some Augochlorella are more of a greenish-bronze color.

Augochlorella aurata is very similar to Augochlorella persimilis. Female Augochlorella aurata can be recognized by having the striations of the propodeum continuing to the posterior margin (Fig. 13C–E), which often, but not always, is bordered by a carina (e.g., Fig. 13C). In contrast, females of Augochlorella persimilis always have a distinct smooth portion before the margin of the propodeum (Fig. 13F–H). In addition, female Augochlorella aurata are generally larger, have the head slightly longer and the apex of the clypeus is black only on the apical fourth (Fig. 13A). In contrast, Augochlorella persimilis females are generally quite small, have the head slightly broader, and the apex of the clypeus is black on the apical third (Fig. 13B).

Female Augochlorella aurata are also often confused with Augochlora pura, but Augochlorella aurata have the paraocular lobes less protuberant (Fig. 3D, F) than Augochlora pura, and Augochlorella aurata also lack a keel on S1.

Male Augochlorella aurata can be separated from Augochlorella persimilis by the hair on the apical two-thirds of the inner edge of the hind basitarsus, which is short in Augochlorella aurata, with the length of the hairs about equal to the width of the basitarsus (Fig. 13I), whereas Augochlorella persimilis has the hairs distinctly longer than the width of the basitarsus (Fig. 13J). In addition, the striae on the propodeal triangle of Augochlorella aurata reach the posterior margin (as in Fig. 13C–E) whereas male Augochlorella persimilis generally have a smooth portion before the margin.

Male Augochlorella aurata are often confused with Augochlora pura males, but Augochlorella aurata have the margin of S4 concave rather than straight, and they lack distinct punctures on the rear of the propodeum (Fig. 2D), compared to distinctly punctured in Augochlora pura (Fig. 3B).

Augochlorella aurata and Augochlorella persimilis are often confused in collections and some females can intergrade to the degree where they are impossible to differentiate. Males are also frequently confused because the hind basitarsus character is often misinterpreted since both species have the basal third of the basitarsus with distinctly shorter hairs, which can cause confusion in keys that focus on the length of the basal hairs rather than the apical hairs, such as Coelho (2004), or the keys on discoverlife.org that incorrectly state that Augochlorella aurata males have the “hair on rear basitarsus all about the same length”.

Given the high level of variation in Augochlorella aurata, it seems likely that it is a species complex. Supporting this hypothesis are the various forms that Ordway (1966) recognized, one of which was elevated to species rank by Coelho (2004), as well as the high barcode diversity found in the species (Sheffield et al. 2009).

Augochlorella persimilis is very similar to Augochlorella aurata. Females can be distinguished by the lack of rugae at the rear of the propodeal triangle, though this character can often be subtle (Fig. 13F–H). In addition, Augochlorella persimilis tend to be smaller than Augochlorella aurata, and they have a more extensive apical black mark on the clypeus, with the black part taking up approximately one-third of the length of the clypeus (Fig. 13B), compared to approximately one-fourth the length of the clypeus in Augochlorella aurata (Fig. 13A). Note that there are often females of Augochlorella persimilis and Augochlorella aurata that cannot be reliably separated.

Male Augochlorella persimilis can be separated from Augochlorella aurata by the length of the hairs on the inner side of the hind basitarsus: Augochlorella persimilis have the hairs very short for the basal third, then the hairs flare out to about twice the width of the basitarsus (Fig. 13J). In contrast, the hairs on Augochlorella aurata are short for the basal third, and only get slightly longer, about equal in length to the width of the basitarsus (Fig. 13I). Like females, the males of Augochlorella persimilis also have a lack of rugae at the rear of the propodeal triangle but it is less distinct (see Fig. 13F–H).

This species has a more southern distribution than Augochlorella aurata, though the species commonly overlap and co-occur. The northern extent of the range of Augochlorella persimilis reaches the southern part of Michigan, Minnesota, and Wisconsin (Wolf and Ascher 2008; Gibbs et al. 2017).

Augochlora pura is the only species of Augochlora that occurs in the midwestern United States.

Augochlora pura is most similar to Augochlorella aurata and Augochlorella persimilis. Both sexes of Augochlora pura can be recognized by the distinct and prominent facial lobes (Fig. 3C, E), which extend below the level of the base of the mandible and are stronger than those found in Augochlorella, but the difference is subtle and easy to confuse. Female Augochlora pura are unique in having a keel on S1 (Fig. 3A) and the hind tibial spur is simple. In addition, the mandibles of Augochlora pura females are more robust, with 2 distinct and nearly equally-sized apical teeth, whereas Augochlorella have 1 main tooth and a smaller subapical tooth.

Males of Augochlora pura can be further recognized from Augochlorella by their straight apical margin on S4 (compared to concave in Augochlorella) and they have distinct punctures on the rear of the propodeum (Fig. 3B), compared to impunctate or obscure punctures in Augochlorella males (Fig. 2D).

Augochlora and Augochlorella are frequently confused in collections, especially males. Midwestern specimens of Augochlora pura fall under subspecies Augochlora pura pura. More work is needed to determine whether Augochlora pura pura and Augochlora pura mosieri Cockerell are distinct taxa.

Both sexes of Augochloropsis are diagnosed by the unique shape of the tegula, which has the inner posterior margin hooked (Fig. 1A). Females have the inner hind tibial spur with multiple straight teeth compared to broad teeth in Agapostemon or untoothed spurs in Augochlorella and Augochlora. Augochloropsis males have a uniquely-shaped S4 (see Fig. 20C, H, M), with a median point and lateral arms, though the sternum is typically hidden. Though typically strongly metallic green, many individuals are metallic bluish or even purplish.

Both sexes of Augochloropsis humeralis can be distinguished from Augochloropsis metallica and Augochloropsis viridula by multiple characters. The pronotal flange of Augochloropsis humeralis has the lateral edges approaching 90 degrees (Figs 14D, 16D) whereas, the lateral edges of the pronotal flange in Augochloropsis metallica and Augochloropsis viridula are obtuse (Figs 14E, F, 16E, F, note they do still have a distinct pronotal flange as well). In addition, the vertex of Augochloropsis humeralis rises distinctly above the ocelli (Figs 14A, 16A) whereas it does not rise above the ocelli in metallica and viridula (Figs 14B, C, 16B, C). Finally, the strongly tessellate and “silky” texture of Augochloropsis humeralis (Figs 14G, 16G) is distinct in comparison to Augochloropsis metallica and Augochloropsis viridula (Figs 14H, I, 16H, I).

Augochloropsis humeralis is similar in most respects to Augochloropsis sumptuosa. The females can be separated by the more densely punctate metasomal terga: Augochloropsis humeralis has the punctures on T1 and T2 close together and separated by about one puncture width (at least over most of the terga), whereas Augochloropsis sumptuosa has the punctures always well-separated (about 3–5 puncture widths apart). In addition, females of Augochloropsis sumptuosa have a weak but distinct semicircular carina around the propodeal triangle (Fig. 17D), which Augochloropsis humeralis lacks (Fig. 17A). More work is needed on how to separate the males of Augochloropsis sumptuosa, but the male of Augochloropsis humeralis appears to have the median emargination of S4 more acute (Fig. 20C), compared to more rounded truncate in Augochloropsis sumptuosa (see Mitchell 1960: fig 111 (mislabeled as S5)), though this character is variable and it’s not clear how reliable it is.

What has previously been called Augochloropsis sumptuosa by Mitchell (1960) is not a single species but rather a species complex. Therefore, we have reinstated the name Augochloropsis humeralis Patton for the species occurring in the midwestern United States and retained the name Augochloropsis sumptuosa Smith for the species occurring in the southeastern United States. The exact extent of the range of Augochloropsis sumptuosa is unclear, and it is not clear to what extent the ranges of Augochloropsis humeralis and Augochloropsis sumptuosa may overlap. However, we have so far found no evidence that the two species overlap in range, with Augochloropsis sumptuosa appearing to be limited to the east coast of the United States and Augochloropsis humeralis appearing to be limited to the prairie region (Fig. 18). Historic records of Augochloropsis sumptuosa from Ohio were found to be misidentified Augochloropsis metallica. Since the identity of the midwestern species is clear, we have decided to proceed with a formal split; further delineation of the range of Augochloropsis sumptuosa must be accomplished in future research.

Two syntypes of Augochloropsis humeralis (1 male and 1 female) were located in the ANSP collection, where Sandhouse (1937) reported examining them. Though the specimens are undated and not clearly labeled as type specimens, the labels indicate that they are from the type locality. In addition, the specimens bear labels stating “Augochlora humeralis n.s.” and “Augochlora humeralis n.sp.”. The combination of the little-used name, the “n. sp.”, the type locality, and the fact that Sandhouse (1937) considered these types, makes us confident that these are indeed Patton’s syntypes. As a result, we have designated the female as the lectotype, making the male a paralectotype. Additional paralectotypes may potentially be present at the Smithsonian, as Titus (1901) states “Mr. Ashmead very kindly examined specimens in the U.S.N.M. of A. humeralis Patt., marked ‘N. W. Kans., Williston’”. However, our inquiries to the Smithsonian have received no answer.

Sandhouse (1937) considered the name Augochloropsis caerulea (Ashmead) to have priority because the name humeralis is a secondary homonym in the genus Halictus. However, humeralis is not a secondary homonym in the genus Augochloropsis and the substitute name is no longer in use, so following IZCN Article 59.3, the name humeralis is available and has priority.

Augochlora sumptuosa bolliana Cockerell is from Texas and was synonymized with Augochloropsis sumptuosa by Sandhouse (1937). Based on the online images of one of the syntypes, we are tentatively assigning it as a synonym of Augochloropsis humeralis, but a more critical evaluation of the specimen, with additional Texas material, should be performed.

Augochloropsis humeralis occurs throughout the prairie region, ranging from North Dakota and Minnesota south to New Mexico and Texas, extending to Colorado in the west and Indiana in the east (Fig. 18). Specimens from Indiana Dunes National Park represent the easternmost records.

Figure 17. 

Augochloropsis female propodea A Augochloropsis humeralis B Augochloropsis metallica C Augochloropsis viridula D Augochloropsis sumptuosa E Augochloropsis anonyma F Augochloropsis sp. (likely undescribed species from Eastland, Texas). Scale bars: 1 mm, all images at the same scale.

Augochloropsis humeralis is polylectic and nests are associated with deep sand (MA, pers. obs.). The sociality and the specifics of the nesting biology are unknown.

Colorado: Adams Co.: Denver (39.8207, -104.8613): 1 ♂ [iNat], 29 Aug 2019, @francesco167 leg.; Douglas Co.: (39.3467, -104.7511): 1 ♀ [iNat], Jul 2020, @calebcam leg.; Logan Co.: (40.7752, -103.2721): 1 ♂ [iNat], 22 Aug 2014, R. Webster leg. Illinois: Hancock Co.: Warsaw (40.3427, -91.4493): 1 ♂ [iNat], 14 Aug 2016, A. Moorehouse leg., Monarda punctata; Madison Co.: [MASR]; Mason Co.: (40.3921, -89.9104): 1 ♀ [iNat], 18 Jun 2019, A. Moorehouse leg., Asclepias sp. Indiana: Lake Co.: Indiana Dunes NP, Marquette Trail (41.6111, -87.2365): 1 ♀ [IDNP], 19 Jun 2019, McGill leg., blue pan; Indiana Dunes NP, Miller woods (41.6057, -87.2644): 1 ♂ [IDNP], 12 Sep 2018, McGill leg., white pan; 1 ♀ [IDNP], 4 Jun 2019, McGill leg., yellow pan; 1 ♀ [IDNP], 23 Jul 2019, McGill leg., yellow pan; Indiana Dunes NP, Miller woods (41.6071, -87.2644): 1 ♀ [IDNP], 23 Jul 2019, McGill leg., yellow pan; Newton Co.: Kankakee Sands (41.0848, -87.402): 1 ♀ [iNat], 24 May 2018, D. Lucas leg.; Porter Co.: Indiana Dunes National Lakeshore, Mnoke Prairie (41.6185, -87.1012): 1 ♀ [IDNP], 29 Jun 2017, J. Villalpando leg., bee bowl. Minnesota: Faribault Co.: (43.7, -93.96): 1 ♀ [UMSP], 18 Sep 1911; Fillmore Co.: Pin Oak SNA (43.79261, -92.21915): 1 ♀ [MNDNR], 24 Jul 2017, bowl; Hennepin Co.: (44.9, -93.4): 1 ♂ [UMSP], date unknown; Norman Co.: Agassiz Dunes SNA (47.51154, -96.28976): 1 ♂ [MNDNR], 24 Aug 2015, bowl; Sherburne Co.: Sherburne National Wildlife Refuge (45.46477, -93.67435): 2 ♀ [EERC], 15 Aug 2016, E. Evans leg., bowl; Wabasha Co.: Weaver Dunes (44.27746, -91.93892): 1 ♀ [UMSP], 28 May 2015, M.J. Hatfield leg., Ceanothus herbaceus; Weaver Dunes TNC/SNA (44.25096, -91.93795): 21 ♀ [MNDNR], 6 May 2017, bowl; 15 ♀ [MNDNR], 26 Jun 2017, bowl; 8 ♂ [MNDNR], 24 Jul 2017, bowl; 3 ♂ [MNDNR], 19 Aug 2017, bowl; 1 ♀ [MNDNR], 21 Sep 2017, bowl; Washington Co.: Belwin Conservancy (44.9241, -92.7931): 1 ♀ [EERC], 4 Sep 2015, J. Gardner leg., net, Solidago nemoralis; Belwin Conservancy (44.92569, -92.80001): 1 ♀ [CRC], 12 Jun 1995, C.C. Reed leg., net, Penstemon grandifloris; 1 ♀ [UMSP], 12 Jun 1995, C.C. Reed leg., net, P. grandiflorus; 9 ♀ [CRC], 15 Jun 1995, C.C. Reed leg., net, P. grandifloris; 6 ♀ [CRC], 16 Jun 1995, C.C. Reed leg., net, P. grandifloris; 1 ♀ 3 ♂ [CRC, UMSP], 15 Aug 1995, C.C. Reed leg., net, Dalea purpurea; 3 ♀ [CRC], 13 Jun 1997, C.C. Reed leg., net, P. grandifloris; Gray Cloud Dunes (44.79, -92.957): 1 ♀ 4 ♂ [UMSP], 9 Jul 1988; Grey Cloud Dunes (44.79, -92.957): 1 ♂ [CNBL], 23 Jul 2018, J. Petersen leg., net; Grey Cloud Dunes (44.7912, -92.9601): 1 ♂ [iNat], 14 Sep 2018, A. Birkey leg.; Grey Cloud Dunes SNA (44.79004, -92.95536): 1 ♂ [MNDNR], 9 Oct 2018, net, S. nemoralis; Grey Cloud Dunes SNA (44.790046, -92.955076): 1 ♂ [MNDNR], 31 Jul 2018, net, D. villosa; Winona Co.: Whitewater WMA (44.15033, -92.00066): 1 ♀ [MNDNR], 6 May 2017, bowl; 1 ♀ [MNDNR], 26 Jun 2017, bowl. Missouri: Clark Co.: [MASR]; Scott Co.: [MASR]. Nebraska: Hooker Co.: [MASR]; Rock Co.: (42.5, -99.8): 1 ♂ [iNat], Sep 2018, @allysond leg.; Thomas Co.: Neb Ntl For, near Halsey, 1 ♂ [WRME], 9 Aug 1991, Arduser leg.; Neb. Ntl For. Nr Whitetail campground: 1 ♀ [WRME], 10 Aug 1991, Arduser leg., Helianthus petiolarus. New Mexico: Chaves Co.: [MASR]. North Dakota: Ransom Co.: (46.474534, -97.342645): 1 ♂ [iNat], 9 Aug 2021, E. Wood leg.. Oklahoma: Ellis Co.: [MASR]. South Dakota: Clay Co.: Missouri National Recreation River (42.76215, -96.9743): 1 ♂ [iNat], 7 Jul 2021, @stenthesnake leg. Texas: Taylor Co.: (32.32, -99.92): 1 ♀ [OSUC], 18 Jun 1952, J.N. Knull, D.J. Knull leg.

Figure 18. 

Map of specimens or observations examined for this study from the species Augochloropsis humeralis, Augochloropsis anonyma, and Augochloropsis fulgida.

Both sexes of Augochloropsis metallica are most similar to Augochloropsis viridula, but Augochloropsis metallica can be separated from Augochloropsis viridula by the thicker hair fringe on the apical edge of T1 and T2; Augochloropsis metallica has the hairs noticeably thicker than the other hairs of the metasoma (Figs 14H, 15A, 16H), whereas the fringe hairs of Augochloropsis viridula are not noticeably thicker than the other metasomal hairs (Figs 14I, 15B, 16I). In addition, the terga of Augochloropsis metallica are more closely punctured, separated by about one puncture width on T2 (Fig. 14H, 16H), whereas the terga of Augochloropsis viridula are more sparsely punctured and separated by at least 2–4 puncture widths on T2 (Figs 14I, 16I).

Both sexes of Augochloropsis metallica can be separated from Augochloropsis humeralis by the shape of the pronotal flange and angle, which is smaller and has an obtuse lateral angle in Augochloropsis metallica (Figs 14E, 16E), compared to the larger flange and 90-degree lateral angle in Augochloropsis humeralis (Figs 14D, 16D). In addition, Augochloropsis metallica has the metasomal terga shining, with at most weak tessellation (Figs 14H, 16H), whereas Augochloropsis humeralis has the metasomal terga strongly and densely tessellate, resulting in dull, silky coloration (Figs 14G, 16G).

The holotype of Augochloropsis metallica (Fig. 19) is missing its head, but the punctures and hair bands on the metasoma, combined with the locality of “America” (Moure (1960) states “probably middle eastern U.S.A.”) are sufficient to confirm its identity.

We define Augochloropsis metallica in a much more restricted sense than previous authors, who lumped multiple taxa under Augochloropsis metallica (e.g., Sandhouse 1937; Mitchell 1960). We are here splitting Augochloropsis metallica (as defined by Mitchell 1960) into five taxa: A. metallica, A. cuprea, A. fulgida, A. fulvofimbriata, and A. viridula. In the original revision of the Augochloropsis of the United States, Sandhouse (1937) lumped at least six distinct taxa under the name Augochloropsis cuprea (Table 1). One reason for this over-lumping appears to be that Sandhouse (1937) did not actually examine any of the type specimens, and synonymized many species based on the description alone or through correspondence. Examination of Sandhouse-determined material in the UMSP shows that she consistently lumped specimens of Augochloropsis metallica, Augochloropsis viridula, and a third unknown species together. Mitchell (1960) clearly recognized that Augochloropsis metallica and Augochloropsis viridula [as Augochloropsis metallica fulgida] were distinct, and it is unclear why he only split them into subspecies.

The traditional view that Augochloropsis metallica extends down through Mexico and Central America is almost certainly incorrect and merely an artifact of the erroneously broad definition of the species adopted by previous workers. Though we have examined relatively little material from south of the United States, the material we have examined has not matched any of the US Augochloropsis treated here. The synonymy of Augochloropsis fulvofimbriata Friese, described from Costa Rica, is almost certainly incorrect. The source of the synonymy of Augochloropsis fulvofimbriata was originally made by Sandhouse (1937), who synonymized the male of Augochloropsis fulvofimbriata under Augochloropsis ignita and the female of Augochloropsis fulvofimbriata under Augochloropsis cuprea. However, Michener (1954), in his revision of the bees of Panama, treated Augochloropsis fulvofimbriata as valid, and though he did not mention the synonymy of A. fulvofimbriata under Augochloropsis cuprea, he did state about A. fulvofimbriata: “Sandhouse incorrectly placed this species in the synonymy if [sic] ignita.” Based on that, Michener (1954) clearly considered Augochloropsis fulvofimbriata a valid species (and he certainly would have been familiar with Augochloropsis metallica, which was then called A. cuprea). Moure (1960) did not list Augochloropsis fulvofimbriata as a synonym of Augochloropsis metallica metallica. However, following that, the works of Mitchell (1960), Hurd (1979), Moure and Hurd (1987), and Moure et al. (2007) all treat Augochloropsis fulvofimbriata as a synonym of Augochloropsis metallica metallica, but none of them indicate it is a new synonym, which suggests they were just carrying over the synonymy by Sandhouse (1937). Here, though we have not examined any material of Augochloropsis fulvofimbriata, we follow the classification of Michener (1954) who was the last worker to treat the species, and we formally treat Augochloropsis fulvofimbriata Friese (stat. nov.), as valid. This is supported by the findings of Celis and Cure (2017), who recognized Augochloropsis fulvofimbriata and even classified it in a separate subgenus than Augochloropsis metallica (Augochloropsis s.s. rather than Paraugochloropsis).

Another synonym with issues is Augochloropsis chorisis Vachal, which was originally synonymized under Augochloropsis cuprea by Sandhouse (1937) and listed as a synonym of Augochloropsis metallica metallica by Michell (1960). The type series of Augochloropsis chorisis contains specimens ranging from Georgia and Texas to Brazil (Rasmussen 2012), making it undoubtedly a composite series (Cockerell 1949). The specimen from Georgia was designated as a lectotype by Moure and Hurd (1987), who considered it a synonym of Augochloropsis metallica metallica. We have not been able to examine the lectotype which cannot be located at the National Museum of Natural History in Paris, France (A. Touret-Alby, pers. comm.) and we are nominally accepting the synonymy.

Figure 19. 

Augochloropsis metallica holotype female A lateral view B metasoma. Images provided by Lars Vilhelmsen, Sree Selvantharan, and Anders Illum of the Natural History Museum of Denmark, and used with permission. Scale bars: 5 mm (A); 1 mm (B).

Augochlora (Augochloropsis) cleomis was described from a male and female specimen from near Fort Collins, Colorado (Titus 1901). The types were not examined and it is not entirely clear from the description whether it is a synonym of Augochloropsis metallica or Augochloropsis humeralis. It was originally synonymized with Augochloropsis cuprea by Sandhouse (1937) and was later synonymized with Augochloropsis metallica metallica by Moure (1960). Sandhouse (1937) did not examine any specimens and it is unclear whether Moure (1960) did. Regardless, it is likely a synonym of Augochloropsis metallica, so we are nominally accepting the synonym.

Augochloropsis metallica is polylectic and nests in the ground. However, the specifics of the nesting biology and sociality are unknown. Augochloropsis metallica is often associated with sandy areas, and it has been found in natural habitats (e.g., native prairies, wetlands), as well as disturbed sites and urban areas.

Augochloropsis metallica occurs in the eastern states and across the Great Plains (Fig. 21A). Recent surveys (2009 to present) by MSA and co-workers in Oklahoma, Kansas and Nebraska have found Augochloropsis metallica throughout these states and further to the west (whereas Augochloropsis viridula is absent from those western areas).

Arkansas: Arkansas Co.: [MASR]; Faulkner Co.: [MASR]; Franklin Co.: [MASR]; Jackson Co.: [MASR]; Monroe Co.: [MASR]; White Co.: [MASR]; Woodruff Co.: [MASR]. Illinois: Calhoun Co.: [MASR]; Carroll Co.: [MASR]; Jasper Co.: [MASR]; Madison Co.: [MASR]; Marion Co.: [MASR]; Randolph Co.: [MASR]; Williamson Co.: [MASR]. Iowa: Jasper Co.: [MASR]. Kansas: Barton Co.: [MASR]; Bourbon Co.: [MASR]; Butler Co.: [MASR]; Chase Co.: [MASR]; Coffey Co.: [MASR]; Dickinson Co.: [MASR]; Douglas Co.: (38.88, -95.29): 1 ♀ [UMSP], 11 Jun 1919, W.F. Hoffman leg.; 1 ♀ [UMSP], 2 Jul 1919, W.F. Hoffman leg.; Geary Co.: [MASR]; Gove Co.: [MASR]; Greenwood Co.: [MASR]; Hodgeman Co.: [MASR]; Lane Co.: [MASR]; Lyon Co.: [MASR]; Morris Co.: [MASR]; Osage Co.: [MASR]; Pawnee Co.: [MASR]; Pottawatomie Co.: [MASR]; Reno Co.: [MASR]; Rice Co.: [MASR]; Riley Co.: [MASR]; Sheridan Co.: [MASR]; Thomas Co.: [MASR]; Trego Co.: [MASR]. Minnesota: Anoka Co.: Bunker Hills Reg. Pk. (45.2176, -93.2898): 1 ♀ [EERC], 8 Jun 2015, J. Gardner leg., net, Tradescantia occidentalis; Bunker Hills Regional Park (45.2176, -93.2899): 1 ♀ [EERC], 13 Jul 2016, E. Evans leg., bowl; Bunker Pr. Dunes (45.21, -93.27): 1 ♀ [UMSP], 20 Jun 1947; Cedar Creek Ecosystem Science Reserve (45.4323, -93.1894): 2 ♀ [EERC], 22 May 2015, J. Gardner leg., bowl trap; Cedar Creek Nat. Hist. (45.402673, -93.202601): 1 ♂ [CRC], 1 Aug 1991, C.C. Reed leg., net, Dalea purpurea; 1 ♂ [UMSP], 20 Aug 1991, C.C. Reed leg.; Cedar Creek Natural History Area (45.402673, -93.202601): 1 ♀ [UMSP], 23 Jul 1986; 1 ♂ [UMSP], 30 Jul 1990; 1 ♀ [UMSP], 21 Sep 1992; 1 ♀ [UMSP], 15 Aug 1995; Cedar Creek Ecosystem Science Reserve (45.4037, -93.1834): 1 ♂ [EERC], 12 Aug 2015, J. Gardner leg., net, D. villosa; Helen Allison Savanna SNA (45.38454, -93.16319): 1 ♀ [MNDNR], 6 May 2017, bowl; Rum River Cent. Reg. Pk. (45.2907, -93.3811): 1 ♀ [EERC], 12 Jun 2015, E. Evans leg., net, Amorpha fruticosa; Anoka/Isanti Co.: Cedar Creek Natural History Area (45.402673, -93.202601): 1 ♀ [UMSP], 15 Aug 2000; Hennepin Co.: Crow Hassan Park Reserve (45.2, -93.63): 2 ♀ [UMSP], 13 Jul 1995, C.C. Reed leg., net, Aster sericeus; Isanti Co.: Cedar Creek Natural History Area (45.402673, -93.202601): 1 ♀ [UMSP], 29 Aug 1981; 1 ♀ [UMSP], 13 Jul 1991; 1 ♀ [UMSP], 14 Jul 1992; 1 ♀ [UMSP], 30 Sep 1992; 1 ♀ [UMSP], 11 Aug 1993; Irving & John Anderson County Park (45.4602, -93.0594): 1 ♂ [EERC], 20 Jul 2015, E. Evans leg., net, Asclepias tuberosa; Lincoln Co.: Hole in the Mountain (44.25680554, -96.29248338): 1 ♀ [UMSP], 15 Jun 2016, N. Pennarolla, J. Leone leg., bowl; 1 ♀ [UMSP], 27 Jun 2017, N. Pennarolla, J. Leone leg., bowl; Hole-in-the-Mountain TNC (44.2412, -96.29963): 2 ♀ [MNDNR], 6 Jun 2016, bowl; 1 ♀ [MNDNR], 27 Jun 2016, bowl; Murray Co.: (44.0709, -95.5718): 1 ♀ [CNBL], 29 Jun 2019, Bee Bowls; Pipestone Co.: Prairie Coteau SNA (44.1241, -96.15275): 1 ♀ [MNDNR], 6 Jun 2016, bowl; Stearns Co.: St. Cloud (45.44, -94.16): 1 ♀ [UMSP], 22 Jun 1967; Yellow Medicine Co.: Mound Spring Prairie SNA (44.74521, -96.42999): 6 ♀ [MNDNR], 6 Jun 2016, bowl. Missouri: Barry Co.: [MASR]; Barton Co.: [MASR]; Benton Co.: [MASR]; Boone Co.: Columbia (38.943, -92.333): 1 ♀ [OSUC], 19 Oct 1955, W.A. Dimmitt leg.; Camden Co.: [MASR]; Douglas Co.: [MASR]; Franklin Co.: [MASR]; Harrison Co.: [MASR]; Howard Co.: Fayette (39.141, -92.686): 9 ♂ [UMSP], 25 Sep 1966, D.B. Crockett leg.; Jackson Co.: [MASR]; Jasper Co.: [MASR]; Jefferson Co.: [MASR]; Laclede Co.: [MASR]; Lafayette Co.: [MASR]; Linn Co.: [MASR]; Macon Co.: [MASR]; Mercer Co.: [MASR]; Miller Co.: [MASR]; Monroe Co.: [MASR]; Newton Co.: [MASR]; Pettis Co.: [MASR]; Ray Co.: [MASR]; Reynolds Co.: [MASR]; Saline Co.: [MASR]; Scott Co.: [MASR]; St. Clair Co.: [MASR]; St. Louis Co.: [MASR]; Ste. Genevieve Co.: [MASR]; Stoddard Co.: [MASR]; Sullivan Co.: [MASR]; Taney Co.: [MASR]. Nebraska: Co.: Halsey (41.904, -100.27): 1 ♀ [UMSP], 3 Sep 1924, R.W. Dawson leg.; Lancaster Co.: [MASR]; Richardson Co.: [MASR]. North Carolina: Wake Co.: Raleigh (35.799, -78.617): 1 ♀ [UMSP], 26 May 1940; 1 ♂ [UMSP], 17 Nov 1940. Ohio: Gallia Co.: (38.82, -82.3): 1 ♀ [OSUC], 23 Aug 1942, C.H. Kennedy leg.; Jackson Co.: (39.01, -82.61): 1 ♂ [OSUC], 9 Aug 1942, J.E. Gillaspy leg.; 1 ♀ [OSUC], 9 Aug 1942, R.W. Strandtmann leg.; Lawrence Co.: (38.6, -82.52): 1 ♂ [OSUC], 8 Aug 1942, R.W. Strandtmann leg.; 4 ♀ 1 ♂ [OSUC], 9 Aug 1942, R.W. Strandtmann leg.; 6 ♀ 1 ♂ [OSUC], 23 Aug 1942, C.H. Kennedy leg.; Muskingum Co.: New Concord (39.995, -81.741): 1 ♀ [OSUC], 22 May 1975, C. Dasch leg. Oklahoma: Ellis Co.: [MASR]. South Dakota: Co.: Black Hills (43.96, -103.77): 1 ♀ [UMSP], 15–30 Jun 1931, F. Miller leg. Texas: Dallas Co.: (32.73, -96.8): 1 ♀ [UMSP], 14 May 1937, H.C. Knutson leg., Marshallia caespitosa; Smith Co.: (32.39, -95.26): 1 ♀ [UMSP], May 1947, Barr leg. Virginia: Arlington Co.: (38.87, -77.09): 2 ♂ [UMSP], 20 Jul 1929, C.E. Michel leg.; Fauquier Co.: Warrentown (38.721, -77.799): 1 ♂ [UMSP], 28 Jul 1929, C.E. Michel leg.

Both sexes of Augochloropsis viridula can be recognized primarily by the lack of a thickened hair fringe on T1 and T2; the hairs that are present along the margin are slender and the same size and width as the rest of the hairs on the terga (Fig. 15B). This is in contrast to the females of Augochloropsis metallica which have the fringe hairs noticeably thickened (Fig. 15A). In addition, the terga of Augochloropsis viridula are more shining and sparsely punctured, and this is most apparent on T1 and T2, with punctures on T2 separated by at least 2–4 puncture widths on T2 (Figs 14I, 16I). In contrast, Augochloropsis metallica has the punctures on T1 and T2 closer together (punctures typically separated by 1–2 puncture widths on T2) and the interspaces between the punctures have slight tessellation, though they are still somewhat shining (Figs 14H, 16H).

Figure 20. 

Augochloropsis male terminalia A Augochloropsis humeralis dorsal genitalia B Augochloropsis humeralis ventral genitalia C Augochloropsis humeralis S4 D Augochloropsis humeralis S8 E Augochloropsis humeralis S7 F Augochloropsis metallica dorsal genitalia G Augochloropsis metallica ventral genitalia H Augochloropsis metallica S4 I Augochloropsis metallica S8 J Augochloropsis metallica S7 K Augochloropsis viridula dorsal genitalia (note the gonobase was torn off) L Augochloropsis viridula ventral genitalia M Augochloropsis viridula S4 N Augochloropsis viridula S8 O Augochloropsis viridula S7. Scale bars: 1 mm.

Augochloropsis viridula can be separated from Augochloropsis humeralis by its less developed pronotal flange, which is weak and forms an obtuse angle in Augochloropsis viridula (Figs 14F, 16F) compared to extensive and forming a right angle in Augochloropsis humeralis (Figs 14D, 16D), and Augochloropsis viridula has the terga smooth and shining (Figs 14I, 16I), compared to heavily tessellate and dull in Augochloropsis humeralis (Figs 14G, 16G).

Augochloropsis viridula can be separated from Augochloropsis anonyma and Augochloropsis fulgida by the relative lack of black hairs on the metasoma, having at most a few scattered black hairs on the apical terga (Fig. 14I), compared to copious black hairs over most of the terga in Augochloropsis anonyma and Augochloropsis fulgida (Fig. 23C, D).

Augochloropsis viridula is a polylectic ground-nester. Nesting and sociality were documented by Gibbs (2017) [as Augochloropsis metallica fulgida]. Gibbs (2017) found a nest with two females, one of which had undeveloped ovaries and could have been a worker, suggesting that this bee may be primitively eusocial, however more work is needed to understand the degree of sociality of this species.

Augochloropsis viridula occurs throughout the eastern United States (Fig. 21B). It does not spread as far west as Augochloropsis metallica (see Fig. 21A), and recent surveys (2009 to present) by MSA and co-workers in Oklahoma, Kansas and Nebraska have found viridula only on the extreme eastern margins of these states, while metallica has been found throughout these states and further to the west. Mitchell records it [as metallica fulgida] extending south to Florida, though we have not evaluated material from the purported southernmost portion of the range.

This species has historically been referred to as Augochloropsis metallica fulgida sensu Mitchell (1960). However, after examination of the holotype of Augochloropsis fulgida (Fig. 23), we found that it does not match the species concept used by Mitchell (1960) for Augochloropsis metallica fulgida. As a result, we resurrect the name Augochloropsis viridula (Smith), which was previously used by Robertson (1902) and Dreisbach (1945). Augochloropsis fulgida is reinstated as a separate species (see remarks for that species).

The species-level (rather than subspecies-level) recognition of Augochloropsis metallica and Augochloropsis viridula (previously classified as Augochloropsis metallica metallica and Augochloropsis metallica fulgida, respectively, by Mitchell (1960)) is supported by multiple characters, including the hair fringe on T2 (see Fig. 15A, B), difference in the degree of punctures and tessellation, the male terminalia, and differences in the extent of range. In particular, the male terminalia are distinct, with Augochloropsis metallica having the gonostyli more expanded (Fig. 20F, G), the gonocoxites more parallel-sided (Fig. 20F, G), the lateral arms of S4 more straight (Fig. 20H), and the lateral apodemes of S7 wider (Fig. 20J), in comparison, Augochloropsis viridula have the gonostyli narrower (Fig. 20K, L), the gonocoxites diverging apically (Fig. 20K, L), the lateral arms of S4 more curved (Fig. 20M), and the lateral apodemes of S7 narrower (Fig. 20O). The shape of S8 (Fig. 20I, N) appears to be too variable to be useful as a splitting character.

That Augochloropsis viridula (Smith) and Augochloropsis lucidula (Smith) were different sexes of the same species was recognized by Patton (1879), Robertson (1895), and Cockerell (1905). However, both names were synonymized under Augochloropsis cuprea (along with Augochloropsis anonyma) by Sandhouse (1937). The name Augochloropsis viridula was then correctly applied by Dreisbach (1945). Mitchell (1960) clearly did not consider viridula and lucidula conspecific as he considered Augochloropsis viridula a junior synonym of Augochloropsis metallica metallica and Augochloropsis lucidula a junior synonym of Augochloropsis metallica fulgida. Moure (1960) considered both viridula and lucidula junior synonyms of Augochloropsis metallica. Here, after examination of the primary types, we agree with the interpretation of Patton (1879) and Robertson (1895) in considering Augochloropsis viridula and Augochloropsis lucidula as both conspecific and a true species.

In the generic revision of augochlorine bees by Eickwort (1969), the subspecies of Augochloropsis metallica were not recognized. However, the illustrations of the genitalia and other characters are clearly of Augochloropsis viridula (rather than Augochloropsis metallica) based on the apically diverging lateral margins of the gonostyli.

USA: Alabama: Hale Co.: [MASR]. Arkansas: Lawrence Co.: [MASR]; Monroe Co.: [MASR]; White Co.: [MASR]; Woodruff Co.: [MASR]. Georgia: Catoosa Co.: [MASR]. Illinois: Carroll Co.: [MASR]; Jasper Co.: [MASR]; Madison Co.: [MASR]; Marion Co.: [MASR]; Ogle Co.: (41.8751, -89.3474): 1 ♀ [NACH], 1 Jul 2017, B. Bruninga-Socolar leg., net, Parthenium integrifolium; (41.896, -89.3461): 1 ♀ [NACH], 13 Jun 2017, B. Bruninga-Socolar leg., net, Trifolium pratense; Randolph Co.: [MASR]; Williamson Co.: [MASR]. Indiana: Lake Co.: Indiana Dunes NP, Miller woods (41.6057, -87.2644): 1 ♀ [IDNP], 23 Aug 2019, McGill leg., blue pan. Iowa: Clayton Co.: [MASR]; Jasper Co.: [MASR]; Pottawattamie Co.: [MASR]; Story Co.: Ames (42.016, -93.624): 2 ♀ [UMSP], 16 Jun 1930, B.A. Haws leg., Swept from sweet clover. Kansas: Johnson Co.: [MASR]; Linn Co.: [MASR]. Maine: Knox Co.: (44.04, -69.04): 1 ♀ [OSUC], 15 Jul 1956, D.J. Borror leg. Michigan: Cheboygan Co.: (45.48, -84.49): 1 ♀ [OSUC], date unknown, C.H. Kennedy leg.; Gladwin Co.: [MASR]. Minnesota: Anoka Co.: Bunker Hills Regional Park (45.2143, -93.2797): 1 ♀ [EERC], 24 Jun 2016, J. Gardner leg., net, Crepis tectorum; Cedar Creek Nat. Hist. (45.402673, -93.202601): 1 ♂ [UMSP], 1 Aug 1991, C.C. Reed leg.; Cedar Creek Natural History Area (45.402673, -93.202601): 1 ♀ [UMSP], 10 May 1993; Rum River Cent. Reg. Pk. (45.28686656, -93.37669731): 1 ♀ [EERC], 12 Jun 2015, E. Evans leg., net, Rosa arkansana; Rum River Cent. Reg. Pk. (45.2883, -93.38): 2 ♀ [EERC], 12 Jun 2015, E. Evans leg., net, Zizia aurea; Rum River Cent. Reg. Pk. (45.2907, -93.3811): 7 ♀ [EERC], 12 Jun 2015, E. Evans leg., net, Amorpha fruticosa; Anoka/Isanti Co.: Cedar Creek Natural History Area (45.402673, -93.202601): 1 ♀ [UMSP], 17 Sep 2004; Blue Earth Co.: Gilfillan Lake WMA (44.21091, -93.8494): 1 ♀ [MNDNR], 3 Oct 2016, net, Symphyotrichum lanceolatum; Maple River WMA (43.979867, -94.042629): 1 ♂ [MNDNR], 14 Aug 2015, net, Solidago altissima; Carver Co.: Schneewind WMA (44.80941, -93.82892): 1 ♀ [MNDNR], 8 Aug 2018, net, Melilotus alba; Schneewind WMA (44.80952, -93.82793): 1 ♀ [MNDNR], 16 Jul 2018, bowl; Chisago Co.: Wild River SP (45.5215, -92.7309): 1 ♀ [MNDNR], 22 Jun 2020, N. Gerjets leg., pantrap; Douglas Co.: StaffansonTNC (45.81606, -95.74604): 1 ♀ [CNBL], 5 Jun 2018, G. Pardee leg., net, Z. aptera; 1 ♀ [CNBL], 5 Jun 2018, I. Lane leg., net, Z. aptera; Fillmore Co.: (43.7, -92.2): 1 ♀ [UMSP], 24 May 1937, G. Kohls leg.; Goodhue Co.: Frontenac (44.53, -92.351): 1 ♀ [UMSP], 29 May 1930, C.E. Michel leg.; Spring Creek Prairie SNA (44.55522, -92.59502): 1 ♂ [MNDNR], 11 Aug 2017, net, Asclepias verticillata; Goodhue/Wabasha Co.: E Frontenac, Lake Pepin (44.53, -92.351): 1 ♀ [UMSP], 29 May 1941, M.W. Wing leg.; Frontenac, Lake Pepin (44.53, -92.351): 1 ♀ [UMSP], 29 May 1941, M.W. Wing leg., net; Hennepin Co.: (44.9, -93.4): 1 ♀ [UMSP], 27 May 1922, A.A. Nichol leg.; Crow-Hassan Park Reserve (45.2018, -93.6311): 1 ♀ [MNDNR], 25 Aug 2015, bowl; Minnesota Valley National Wildlife Refuge (44.79892, -93.38589): 1 ♀ [MNDNR], 17 Jul 2017, net, Solanum dulcamara; St Bonifacius: 6 Mile Marsh (44.9113, -93.71958): 1 ♀ [CNBL], 28 Jul 2018, Z. Portman leg., net, M. alba; St Bonifacius: 6 Mile Marsh (44.9121, -93.7217): 1 ♀ [CNBL], 5 Jun 2020, Z. Portman leg., net, Z. aurea; Houston Co.: (43.67, -91.5): 3 ♀ [UMSP], 21 May 1938, H.E. Milliron leg.; (43.68, -91.47): 2 ♀ [UMSP], 23 May 1936, C.E. Michel leg.; 4 ♀ [UMSP], 23 May 1936, D. Murray leg.; 1 ♀ [UMSP], 23 May 1936, O. Elster leg.; 3 ♀ [UMSP], 23 May 1936, R. Cottrell leg.; 1 ♀ [UMSP], 22 May 1937, H.S. Telford leg.; 2 ♀ [UMSP], 24 May 1937, C.E. Michel leg.; 1 ♀ [UMSP], 20 May 1938, P. Nicholson leg.; 3 ♀ [UMSP], 21 May 1938, C.E. Michel leg.; 2 ♀ [UMSP], 21 May 1938, H.E. Milliron leg.; 1 ♀ [UMSP], 21 May 1938, R. Anderson leg.; 1 ♀ [UMSP], 22 May 1938, R. Anderson leg.; 1 ♀ [UMSP], 21 Jun 1938, C.E. Michel leg.; 1 ♀ [UMSP], 26 May 1940, I. Tarshie leg.; Beaver Crk. Valley St. Park (43.642, -91.581): 2 ♀ [UMSP], 4 Jul 1973, Malaise trap; Eitzen (43.51, -91.46): 1 ♀ [UMSP], 23 May 1936; Mississippi Bluff, 1–2 m N State Line (43.524, -91.28): 1 ♀ [UMSP], 30 May 1941, J.H. Hughes leg.; 1 ♀ [UMSP], 27 May 1950; 1 ♀ [UMSP], May 1957; Mississippi Bluffs 1 mi N. New Albin, Ia. (43.514, -91.279): 1 ♀ [UMSP], 29 May 1960; Mound Prairie SNA (43.76248, -91.42277): 1 ♀ [MNDNR], 26 Jun 2017, bowl; S.E. tip of county (43.52, -91.29): 1 ♀ [UMSP], 24 May 1935, H. Dodge leg.; Winnebago Cr. Vy., 2–4 m NE Eitzen (43.541, -91.415): 1 ♀ [UMSP], 27 Jun 1956; Isanti Co.: Cedar Creek Natural History Area (45.402673, -93.202601): 1 ♀ [UMSP], 1 Aug 1985; 2 ♀ [UMSP], 21 May 1987, Rubus sp; 1 ♂ [UMSP], 30 Sep 1992; 1 ♀ [UMSP], 1 Sep 1993; 2 ♀ [UMSP], 27 Jul 1994; 1 ♂ [UMSP], 17 Sep 1994; 1 ♀ [UMSP], 19 Aug 2000; 1 ♀ [UMSP], 19 Aug 2001; Jackson Co.: Des Moines River SNA (43.79222, -95.09111): 6 ♀ [MNDNR], 6 Jun 2016, bowl; Kanabec Co.: Rice Creek WMA (45.7389, -93.2044): 1 ♀ [MNDNR], 30 Jun 2020, D. Drons leg., net, Rhus glabra; Kandiyohi Co.: Brenner (45.4006, -95.2462): 1 ♀ [CNBL], 7 Jun 2018, G. Pardee leg., net, Z. aptera; Brenner Lake WPA (45.39926, -95.24568): 2 ♀ [MNDNR], 6 Jul 2016, bowl; Nelson (45.35289989, -95.11923718): 1 ♀ [CNBL], 26 Jun 2017, R. Tucker leg., net, Cirsium arvense; 1 ♀ [CNBL], 4 Jun 2018, I. Lane leg., net, Z. aurea; 3 ♀ [CNBL], 4 Jun 2018, S. Marconie leg., net, Z. aurea; 1 ♀ [CNBL], 4 Jun 2018, T. Eicholz leg., net, Z. aurea; Rudningen (45.32725, -95.17902): 1 ♀ [CNBL], 26 Jun 2017, C. Herron-Sweet leg., net, Achillea millefolium; Le Sueur Co.: Dove Lake WMA (44.22547, -93.7065): 1 ♀ [MNDNR], 1 Sep 2017, net, So. Sp; Kasota Prairie SNA (44.26502, -94.00384): 1 ♀ [MNDNR], 6 May 2017, bowl; Lyon Co.: Glynn Prairie SNA (44.2637757, -95.69623097): 1 ♀ [UMSP], 20 Jul 2017, N. Pennarolla, J. Leone leg., bowl; Mille Lacs Co.: Kunkel WMA (45.5741, -93.6623): 1 ♀ [MNDNR], 24 Jun 2020, D. Drons leg., pantrap; Princeton (45.571, -93.578): 1 ♀ [UMSP], 3 Oct 1994, A. Johnson leg.; Murray Co.: (44.0709, -95.5718): 1 ♀ [CNBL], 29 Jun 2019, Bee Bowls; Olmsted Co.: Oronoco Prairie SNA (44.14002349, -92.48913144): 1 ♀ [MNDNR], 13 Sep 2013, bowl; Pine Co.: Chengwatana State Forest (45.819, -92.7864): 1 ♂ [MNDNR], 16 Jul 2020, N. Gerjets leg., net, Veranicastrum virginicum; St. Croix SP (45.9543, -92.5799): 1 ♀ [MNDNR], 25 Aug 2020, N. Gerjets leg., net, So. Sp.; Pope Co.: Glacial Lakes State Park (45.541, -95.531): 1 ♂ [UMSP], 25 Jul 1973, Malaise trap; Ramsey Co.: Bald Eagle Otter Lk. Reg. Pk. (45.09588474, -93.0494575): 1 ♀ [EERC], 5 Sep 2015, E. Evans leg., bowl trap; Battle Creek Reg. Pk. (44.9346, -93.0141): 1 ♀ [EERC], 10 Jun 2015, J. Gardner leg., net, Cornus sericea; Battle Creek Reg. Pk. (44.93505, -93.015467): 2 ♀ [EERC], 27 May 2015, J. Gardner leg., net, Geranium maculatum; Battle Creek Regional Park (44.9345, -93.013): 1 ♀ [EERC], 17 May 2016, E. Evans leg., bowl trap; Battle Creek Regional Park (44.94, -93.001): 2 ♀ [EERC], 8 Jun 2016, J. Gardner leg., net, Ru. Allegheniensis; Roseville, 3035 Fairview Avenue N (45.03262, -93.17757): 1 ♀ [UMSP], 5–7 Sep 2014, R.W. Holzenthal leg.; St Anthony Park (44.98, -93.2): 1 ♀ [UMSP], Jun year unknown; Redwood Co.: Cedar Mountain SNA (44.50489, -94.89886): 12 ♀ [MNDNR], 6 Jul 2016, bowl; Renville Co.: Morton Outcrops SNA (44.5501, -94.9902): 1 ♀ [MNDNR], 6 Jul 2016, bowl; 1 ♀ [MNDNR], 18 Jul 2016, bowl; Sherburne Co.: Sherburne National Wildlife Refuge (45.4973, -93.6851): 1 ♀ [EERC], 10 Jun 2016, E. Evans leg., net, R. arkansana; Uncas Dunes SNA (45.42750645, -93.69554017): 1 ♀ [MNDNR], 11 Jun 2013, net; Stearns Co.: Avon Hills Forest SNA (45.63589, -94.50259): 1 ♂ [MNDNR], 13 Sep 2018, net, So. altissima; St. Cloud (45.44, -94.16): 2 ♀ [UMSP], 25 May 1968; 1 ♂ [UMSP], 30 Jul 1968; Stevens Co.: Freeman WMA (45.46042, -95.97334): 1 ♀ [MNDNR], 21 Jun 2015, net, R. sp; Verlyn Marth Memorial Prairie SNA (45.7451, -96.00017): 3 ♀ [MNDNR], 6 Jul 2016, bowl; Swift Co.: Rice WPA (45.34486541, -95.32010344): 1 ♀ [UMSP], 26 Jun 2016, N. Pennarolla, J. Leone leg., bowl; Wabasha Co.: Reads Landing (44.402, -92.08): 1 ♀ [UMSP], 22 Jun 1934, C.E. Michel leg.; Washington Co.: (45.04, -92.89): 1 ♀ [UMSP], 9 May 1959; Afton State Park (44.846, -92.789): 1 ♀ [CRC], 11 Sep 1992, C.C. Reed leg., net; Arcola Bluffs SAC (45.1209, -92.7509): 1 ♀ [CNBL], 31 May 2018, K. Friedrich leg., vac, G. maculatum; 1 ♀ [CNBL], 14 Jun 2018, K. Friedrich leg., vac, Erigeron philadelphicus; Big Marine Park Res. (44.2014, -92.8796): 6 ♀ [EERC], 7 Jun 2016, J. Gardner leg., net, R. woodsii; Lost Valley Prairie SNA (44.80086892, -92.81775955): 1 ♀ [MNDNR], 13 Sep 2013, bowl; Lost Valley SNA (44.802885, -92.823067): 1 ♀ [UMSP], 19 Sep 1990, C.C. Reed leg.; 1 ♀ [CRC], 19 Sep 1990, C.C. Reed leg., net; 1 ♀ [UMSP], 28 Jul 1992, C.C. Reed leg.; St. Croix Savanna SNA (45.00322082, -92.78344361): 1 ♀ [MNDNR], 16 Sep 2013, bowl; St. Croix Savanna SNA (45.00540834, -92.78347343): 1 ♀ [MNDNR], 13 Sep 2013, bowl; St. Croix Savanna SNA (45.006475, -92.785823): 1 ♀ [UMSP], 5 Aug 1994, C.C. Reed leg., Monarda fistulosa; Winona Co.: Great River Bluffs SP (43.93895, -91.4113): 1 ♀ [MNDNR], 19 Aug 2017, bowl; Wright Co.: Lake Maria SP (45.31787, -93.93487): 1 ♀ [MNDNR], 6 May 2017, bowl. Mississippi: Bolivar Co.: Cleveland (33.741, -90.742): 1 ♀ [UMSP], 21 Apr 1937, R.W. Dawson leg. Missouri: Atchison Co.: [MASR]; Barry Co.: [MASR]; Barton Co.: [MASR]; Benton Co.: [MASR]; Bollinger Co.: [MASR]; Callaway Co.: [MASR]; Camden Co.: [MASR]; Crawford Co.: [MASR]; Dallas Co.: [MASR]; Dent Co.: [MASR]; Douglas Co.: [MASR]; Franklin Co.: [MASR]; Greene Co.: [MASR]; Grundy Co.: [MASR]; Harrison Co.: [MASR]; Jackson Co.: [MASR]; Jasper Co.: [MASR]; Jefferson Co.: [MASR]; Johnson Co.: [MASR]; Laclede Co.: [MASR]; Lafayette Co.: [MASR]; Lewis Co.: [MASR]; Lincoln Co.: [MASR]; Macon Co.: [MASR]; Madison Co.: [MASR]; Mercer Co.: [MASR]; Monroe Co.: [MASR]; Montgomery Co.: [MASR]; Pemiscot Co.: [MASR]; Pettis Co.: [MASR]; Putnam Co.: [MASR]; Randolph Co.: [MASR]; Ray Co.: [MASR]; Reynolds Co.: [MASR]; Saline Co.: [MASR]; Shannon Co.: [MASR]; St. Francis Co.: [MASR]; St. Louis Co.: [MASR]; Ste. Genevieve Co.: [MASR]; Stoddard Co.: [MASR]; Taney Co.: [MASR]; Warren Co.: [MASR]. New York: Tompkins Co.: Ithaca (42.442, -76.501): 1 ♀ [OSUC], 27 Aug 1950, J. Cillie leg. North Carolina: Sampson Co.: Ivanhoe (34.58, -78.25): 1 ♀ [UMSP], 3 May 1945, T.B. Mitchell leg. Ohio: Champaign Co.: (40.13, -83.77): 1 ♂ [OSUC], 24 Jul 1954; 1 ♀ [OSUC], 8 Jun 1994, N.F. Johnson leg., Malaise trap; Delaware Co.: (40.27, -83.01): 1 ♀ [OSUC], 2 Aug 1942, R.W. Strandtmann leg.; Fairfield Co.: (39.75, -82.63): 1 ♀ [OSUC], 16 Jun 1994, A. Sharkov leg.; Franklin Co.: (39.97, -83.01): 1 ♂ [OSUC], 21 Aug 1942; 1 ♀ [OSUC], 18 Jun 1952; Greene Co.: [MASR]; (39.69, -83.89): 1 ♀ [OSUC], 6 Jun 1956, J.N. Knull leg.; 1 ♀ [OSUC], 20 Jun 1957, J.N. Knull, D.J. Knull leg.; Hocking Co.: (39.49, -82.48): 1 ♀ [OSUC], 10 May 1935, R.C. Osburn leg.; 1 ♀ [OSUC], 14 Jun 1943, R.C. Osburn leg.; 1 ♀ [OSUC], 23 May year unknown, J.N. Knull, D.J. Knull leg.; 1 ♀ [OSUC], 14 Jun year unknown, R.C. Osburn leg.; Jackson Co.: (39.01, -82.61): 1 ♀ [OSUC], 9 Aug 1942, R.W. Strandtmann leg.; Lawrence Co.: (38.6, -82.52): 1 ♂ [OSUC], 8 Aug 1942, R.W. Strandtmann leg.; Logan Co.: (39.54, -82.41): 1 ♀ [UMSP], 16 Jul 1930, J. Patton leg.; Lucas Co.: [MASR]; (41.68, -83.47): 1 ♀ [OSUC], 19 May 2003, M. Arduser leg., Lupinus perennis; Ottawa Co.: Catawba Island (41.579, -82.836): 1 ♀ [OSUC], 27 Jun 1902, J.G. S. leg.; Put-in-Bay (41.649, -82.816): 1 ♀ [OSUC], 20–30 Jun 1924; 1 ♀ [OSUC], 14 Jul 1935, R.C. Osburn leg.; 1 ♀ [OSUC], 22 Aug 1941, R.C. Osburn leg.; 1 ♀ [OSUC], date unknown, C.H. Kennedy leg.; Paulding Co.: Charloe (41.131, -84.434): 1 ♀ [OSUC], 12 May 1951, H.F. Price leg.; Scioto Co.: (38.82, -82.99): 1 ♀ [OSUC], 6 Aug 1942, R.W. Strandtmann leg.; 1 ♀ [OSUC], 9 Jun 1943, J.N. Knull, D.J. Knull leg.; Summit Co.: Ira (41.182, -81.585): 1 ♀ [OSUC], date unknown, J.S. Hine leg.; Vinton Co.: (39.25, -82.49): 1 ♀ [OSUC], 20 Jun 1901; Williams Co.: Bryan (41.472, -84.553): 2 ♀ [OSUC], date unknown. Tennessee: Davidson Co.: Nashville: [MASR]. Wisconsin: Burnett Co.: (43.5, -88.71): 1 ♀ [UMSP], M. Sabourin leg.; Crawford Co.: Barnum (43.218, -90.839): 1 ♀ [UMSP], 2 Aug 1922, A.M. Holmquist leg.; Dane Co.: Madison (43.094, -89.321): 1 ♀ [OSUC], 25 Jun 1916; La Crosse Co.: [MASR]; Oconto Co.: Lakewood (45.3, -88.523): 1 ♀ [UMSP], 15 Jul 1948, H.E. Milliron leg.; Polk Co.: Tewksbury SACN (45.3031, -92.7312): 1 ♀ [CNBL], 25 May 2017, K. Friedrich leg., vac, Barbarea vulgaris; 2 ♀ [CNBL], 8 Jun 2017, K. Friedrich leg., vac, Ru. Sp.; 1 ♀ [CNBL], 4 Jun 2018, K. Friedrich leg., vac, Ru. Sp. Canada: Ontario: Middlesex Co.: London: [MASR].

Both sexes of Augochloropsis anonyma can be recognized by the short propodeal triangle, which is impressed and narrower than the metanotum (Fig. 17E). Augochloropsis anonyma are most likely to be confused with Augochloropsis viridula, as they overlap in range, and both have shining integument and poorly developed apical hair fringes on the metasoma. Augochloropsis anonyma females can be recognized by the presence of dark pubescence on T2–T4, whereas Augochloropsis viridula lacks dark pubescence. In addition, Augochloropsis anonyma has the propodeum dorsal surface impressed, shining, and relatively narrow with its medial length slightly less than the medial length of the metanotum (Fig. 17E). In comparison, Augochloropsis viridula has the propodeum dorsal surface flat throughout, not impressed, and relatively broad, its medial length dorsally as long (or longer) than the medial length of the metanotum (Fig. 17C).

Figure 21. 

Map of specimens examined for this study A Augochloropsis metallica B Augochloropsis viridula. Due to the limited geographic focus of our study, the easternmost extents of the ranges are relatively incomplete.

Augochloropsis anonyma is also similar to Augochloropsis fulgida because both share the character of dark hairs on the metasoma (see Fig. 23C, D for fulgida). However, the narrow propodeal triangle of Augochloropsis anonyma (Fig. 17E) separates these two species, as Augochloropsis fulgida has the propodeal triangle broader (visible in Fig. 23C).

Augochloropsis anonyma is known from the far southeastern US, and we have examined material from Florida and Georgia (Fig. 18). Mitchell (1960) reports it occurring as far north as North Carolina.

The type of Augochloropsis cuprea (Fig. 22) was located in the Oxford Museum, and based on the label, it was examined by J.S. Moure in 1957 and C.D. Michener in 1965. The specimen is not clearly labeled as the type, but both Moure and Michener agreed that it was likely the holotype. Sandhouse (1937) appears to have only examined the type by proxy through sawfly taxonomist R. B. Benson and Mitchell (1960) did not examine it.

Augochloropsis cuprea was considered a junior synonym of Augochloropsis metallica metallica by both Moure (1960) and Mitchell (1960). However, examination of the type of Augochloropsis cuprea reveals that it is distinct from Augochloropsis metallica based on the short T2 fringe (Fig. 22D), the relatively short T3 impressed area (Fig. 22D), the presence of some black hairs on the metasoma (Fig. 22A, D), the more shining propodeum (Fig. 22C), and the short posterior carina of the propodeum (Fig. 22C, D). As a result, we recognize the two forms as heterospecific and thus Augochloropsis cuprea as a valid species.

Figure 22. 

Augochloropsis cuprea holotype female A lateral view B head C rear dorsal view of propodeum D metasoma. Scale bars: 5 mm (A); 1 mm (B); 1 mm (D). Images provided by Dr. James Hogan (OUMNH).

The range of Augochloropsis cuprea is unclear as Smith (1853) reports the type locality as “North America”. We have in our possession a single female from Oklahoma (from Four Canyon Preserve headquarters, Ellis County) that may be a match for Augochloropsis cuprea, but this must be considered tentative, especially since the Oklahoma specimen lacks the number of black hairs on the metasoma seen in the type. There is also the possibility that Augochloropsis cuprea is a Mexican species, especially since multiple Mexican species share the character of the broadly shiny propodeal triangle. The male remains unknown and additional investigation, with more material, is sorely needed.

We define Augochloropsis fulgida differently than previous workers because examination of the type specimen (Fig. 23) revealed that it does not match the species concept used for Augochloropsis metallica fulgida by Mitchell (1960). Even though Mitchell (1960) examined the type, it does not key out correctly in his key or match his description. As it stands, the type of Augochloropsis fulgida does not match any Augochloropsis species we are familiar with. The type female, from St. John’s Bluff Florida (Fig. 18), is most similar to Augochloropsis anonyma in that it has erect dark hairs on the metasoma (Fig. 23C, D), but it differs in having a larger and more tessellate propodeal triangle (Fig. 23C) compared to the narrow and shining propodeal triangle in Augochloropsis anonyma (Fig. 17E), and the hair fringe on T2 of Augochloropsis fulgida appears to be slightly more prominent than in Augochloropsis anonyma.

More work is needed to clarify this species as it is currently only known from the type and the male is unknown. However, we have not performed a dedicated search for more material that could match Augochloropsis fulgida. It is also a possibility that the type of Augochloropsis fulgida is mislabeled and not from Florida or even the United States. However, a more likely explanation is that any Augochloropsis fulgida material has been misidentified as Augochloropsis anonyma due to the presence of black pubescence on the metasoma.

Augochloropsis sumptuosa is most similar to Augochloropsis humeralis (refer to the diagnosis of that species to separate them).

Figure 23. 

Augochloropsis fulgida holotype female A lateral view B head C rear dorsal view D metasoma. Scale bars: 5 mm (A); 1 mm (B). Images provided by Dr. Joseph Monks (NHMUK).

We use Augochloropsis sumptuosa in a more restricted sense than previous authors because we have split it into two species: Augochloropsis sumptuosa and Augochloropsis humeralis. Now, Augochloropsis sumptuosa refers to the species occurring in the southeastern United States though the exact range is unknown at this time, and it remains to be seen to what degree, if any, the range of the two species overlaps.

The type or types of Augochloropsis sumptuosa have been lost. The type could not be located by Sandhouse (1937) or Mitchell (1960). The last report of possible types is from Cockerell (1897), who stated that “Col. C.T. Bingham” examined syntypes in the Natural History Museum (London, UK). However, there are currently no specimens in the Natural History Museum that could possibly be syntypes (J. Monks, pers. comm., Jun 2021). Despite the missing types, it is our opinion that there is not a need for a neotype because the identity of Augochloropsis sumptuosa can be determined from the original description. Specifically, the original description states “the base of the metathorax enclosed by an arched ridge, the enclosed space granulated, the sides of the truncation margined by sharp carinae.” This description matches the southeastern species (which has the propodeal triangle surrounded by a weak semicircular carina; Fig. 17D) but not the species found in the Midwest. On this basis, we are retaining the name Augochloropsis sumptuosa Smith for the southeastern species. The oldest available name for the midwestern species is Augochloropsis humeralis (Patton), the types of which were collected in western Kansas.

More work remains to be done on the taxonomy of Augochloropsis sumptuosa because we have not critically evaluated the status of two synonyms: Augochlora lacustris Cockerell and Augochlora floridica Cockerell. They were originally synonymized with Augochloropsis sumptuosa by Sandhouse (1937), and Mitchell (1960) agreed with that designation, stating “Examination of the types has failed to reveal any significant difference that would justify the recognition of either lacustris or floridica.” We have examined images of the types, which are clear enough for us to tentatively agree. However, given that we have split Augochloropsis sumptuosa into two species and there is potentially a third similar species in Florida, these types should be critically reexamined as part of a reevaluation of the Florida fauna.