Research Article |
Corresponding author: Zachary M. Portman ( zportman@gmail.com ) Academic editor: Thorleif Dörfel
© 2022 Zachary M. Portman, Mike Arduser, Ian G. Lane, Daniel P. Cariveau.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Portman ZM, Arduser M, Lane IG, Cariveau DP (2022) A review of the Augochloropsis (Hymenoptera, Halictidae) and keys to the shiny green Halictinae of the midwestern United States. ZooKeys 1130: 103-152. https://doi.org/10.3897/zookeys.1130.86413
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Augochloropsis and other shiny green Halictinae have had various taxonomic issues and are often misidentified. One prevailing taxonomic issue is that Augochloropsis metallica (Fabricius) has two subspecies, that have long been recognized as morphologically distinct (Augochloropsis metallica metallica and Augochloropsis metallica fulgida (Smith), but the subspecies are inconsistently applied in the literature. Here, we review the Augochloropsis of the Midwest and further address the Augochloropsis species in the broader United States to resolve the outstanding taxonomic issues with the midwestern species. We provide identification keys and diagnoses for the genera and species of the shiny green Halictinae of the midwestern United States, which includes the genera Agapostemon, Augochlora, Augochlorella, and Augochloropsis. This work results in taxonomic changes to Augochloropsis. Augochloropsis sumptuosa (Smith) is split into two species, with the name Augochloropsis sumptuosa retained for the eastern form, and Augochloropsis humeralis (Patton), stat. nov., reinstated for the western form. Augochloropsis metallica is split into five species, with two of those species occurring in the midwestern United States: Augochloropsis metallica and Augochloropsis viridula (Smith), stat. nov. Examination of the holotype of Augochloropsis fulgida (Smith) revealed that it does not agree with the prevailing concept of Augochloropsis metallica fulgida; it is reinstated as Augochloropsis fulgida, stat. nov., but is currently known only from the holotype female from Florida. Augochloropsis cuprea (Smith), long considered to be a synonym of Augochloropsis metallica, is also distinct, and we are reinstating Augochloropsis cuprea, stat. nov., though the range of this species is unclear. We further recognize Augochloropsis fulvofimbriata (Friese), stat. nov., from South and Central America, as distinct. These changes result in a total of three Augochloropsis species in the Midwest and seven named species in the United States. We are aware of additional species from the southern and southwestern United States that are undescribed, and we highlight additional taxonomic work that remains to be done.
Agapostemon, Augochlora, Augochlorella, identification, Paraugochloropsis, species complex
The genus Augochloropsis Cockerell contains approximately 140 species, recognizable by their metallic coloration and distinctly-shaped tegula (
The species of Augochloropsis in the United States and Canada have undergone many taxonomic changes. In the first revisionary work,
There has been inconsistent use of the names and species concepts of the Augochloropsis in the United States in recent bee diversity studies, primarily with the usage of the subspecies of Augochloropsis metallica, with some researchers recognizing them and others not. As a result, when papers refer to “Augochloropsis metallica” it is often impossible to know whether they were referring to “Augochloropsis metallica metallica” or “Augochloropsis metallica fulgida,” especially given that many publications fail to cite the identification resources or taxon concepts they used (see
More broadly, the green Halictinae in general are plagued by misidentifications, particularly of males. This is in part due to the lack of up-to-date or high-quality identification resources. Indeed, some widely used identification resources contain characters that are too variable to be useful or are outright incorrect (e.g., the keys on discoverlife.org;
Here, we review the Augochloropsis species of the Midwest, recognizing three species from the region: Augochloropsis humeralis, Augochloropsis metallica sensu stricto, and Augochloropsis viridula. While we originally aimed to simply clarify the subspecies of Augochloropsis metallica, it necessarily expanded into a larger project after examination of the type specimens revealed numerous issues that necessitated a geographic expansion and a more in-depth update of the taxonomy. As a result of the updated taxonomy, we are making the following changes: Augochloropsis humeralis is resurrected from synonymy with Augochloropsis sumptuosa, we define Augochloropsis fulgida in a different sense than it has traditionally been used, and Augochloropsis viridula is resurrected from synonymy and recognized as a valid species. We further recognize as valid species two former synonyms of Augochloropsis metallica: Augochloropsis cuprea and Augochloropsis fulvofimbriata Friese. We also point to more work that remains to be done, as we recognize seven species in the United States, but there appear to be at least four more undescribed or unrecognized species. Lastly, we provide an illustrated key to the Augochloropsis and the other shiny green Halictinae of the midwestern United States, which covers the genera Agapostemon Guérin-Méneville, Augochlora, Augochlorella Sandhouse, and Augochloropsis.
The keys used here are variously adapted and modified from existing sources, primarily from
The keys and diagnoses follow the species concepts from the most recent revisions of those groups:
The following museum and collection acronyms are used in the paper:
CNBL The collection of the Cariveau Native Bee Lab, St. Paul Minnesota, USA (Z. Portman).
CRC Catherine Reed Collection. Currently resides in the Cariveau Native Bee Lab and will be accessioned into the UMSP.
EERC Elaine Evans Research Collection (E. Evans). Housed at the Cariveau Native Bee Lab (CNBL) and will be accessioned into the UMSP.
IDNP Indiana Dunes National Park. Examined specimens deposited at the UMSP.
iNat Selected high-quality records from the community science portal iNaturalist.com were examined for Augochloropsis humeralis. All record information is included in the material examined section.
MASR Mike Arduser specimen record. Includes a combination of specimens in Mike Arduser’s personal collection, as well as specimens Mike Arduser has personally identified but no longer has in hand.
MNDNR The Minnesota Department of Natural Resources, St. Paul, MN, USA (J. Petersen and N. Gerjets). These are primarily deposited in the UMSP except for a small synoptic collection.
NHMD Natural History Museum of Denmark, Copenhagen, Denmark (L. Vilhelmsen).
OSUC C.A. Triplehorn Insect Collection, Ohio State University Columbus, Ohio, USA (L. Musetti).
OUMNH University Museum of Natural History, Oxford, United Kingdom (J. Hogan).
This study represents material from multiple sources, often examined over the course of many years. As a result, in the reports of material examined we are reporting a combination of specimen-level and county-level data. Historic specimens were manually georeferenced using Google Earth Pro software (v. 7.3.4.8248). For county level records, points were mapped to the county centroids. Specimen images were taken using an Olympus DP27 camera mounted on an Olympus SZX16 stereo microscope, with the images stacked using CombineZP software (
Identification of the shiny green Halictinae of the midwestern United States.
1 | Tegula enlarged and asymmetric, with the inner posterior margin hooked or angled (Fig. |
Augochloropsis Cockerell |
– | Tegula normal and ovoid (Fig. |
2 |
2 | Propodeum with posterior surface encircled by a raised rim or carina (Fig. |
Agapostemon Guérin-Méneville |
– | Propodeum with posterior surface not encircled by a carina (Fig. |
3 |
3 | Female S1 with central keel (Fig. |
Augochlora pura (Say) |
– | Female S1 without keel; both sexes with paraocular lobe not prominent, forming an obtuse or right angle (Fig. |
Augochlorella Sandhouse |
Both sexes of Agapostemon are diagnosed by the complete carina on the rear face of the propodeum (Fig.
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).
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.
Note: We include two principally western species, Agapostemon angelicus Cockerell and Agapostemon melliventris Cresson, that may potentially occur in the midwestern states.
Key to females
1 | Metasoma black or amber-colored (Fig. |
2 |
– | Metasoma metallic green like thorax (Fig. |
3 |
2 | Metasomal terga blackish (Fig. |
virescens (Fabricius) |
– | Metasomal terga amber-colored, at least in part (Fig. |
melliventris Cresson |
3 | Scutum and scutellum “doubly punctate”, i.e., with uniformly scattered large punctures among the more numerous small ones (Fig. |
texanus Cresson or angelicus Cockerell |
– | Scutum densely punctate, rugosopunctate or weakly reticulate (Fig. |
4 |
4 | Pronotum with dorsolateral angle pointed and produced at a distinct right angle; dorsolateral ridge sharply edged (Fig. |
sericeus (Forster) |
– | Pronotum with dorsolateral angle and dorso-lateral ridge blunted (Fig. |
splendens (Lepeletier) |
Agapostemon thorax characters (anterior of bee to left, “mc” refers to the base of the middle coxa) A Agapostemon sericeus close-up of rugosopunctate scutum B Agapostemon splendens close-up of densely punctate scutum C Agapostemon sericeus with ventral pleural tubercle flush with rest of plate D Agapostemon splendens with ventral pleural tubercle slightly upraised. Yellow arrows indicate the ventral pleural tubercle.
Key to males
1 | S4 entirely flat, without a low, transverse swelling along the apical or preapical margin (Fig. |
virescens (Fabricius) |
– | S4 (and S3 to a lesser extent) with a low, transverse swelling along the apical or preapical margin (Fig. |
2 |
2 | Hind femur skinny, not swollen at all (Fig. |
melliventris Cresson |
– | Hind femur moderately to grossly swollen (Fig. |
3 |
3 | Hind legs quite swollen, width of hind femur about half the length (Fig. |
splendens (Lepeletier) |
– | Hind legs only moderately swollen (Fig. |
4 |
4 | F1 slightly more than half length of F2 (Fig. |
sericeus (Forster) |
– | F1 at least three-fourths length of F2 (Fig. |
5 |
5 | Hind tibia with brown to black stripe present anteriorly (and posteriorly) (e.g., Fig. |
texanus Cresson |
– | Lacking brown to black stripe on anterior surface of hind tibia (Fig. |
angelicus Cockerell |
Females of Agapostemon angelicus can be recognized by the unique double-punctate scutum (as in Fig.
Agapostemon angelicus is primarily a western species, though
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.
Male Agapostemon melliventris can be recognized by having the metasoma primarily yellow (Fig.
Agapostemon melliventris is not known from the midwestern US, though
The female of Agapostemon sericeus can be recognized by the combination of the metallic green metasoma (as in Fig.
Male Agapostemon sericeus have S3 and S4 with a low transverse swelling and generally have distinct yellow marks on the apical sterna (Fig.
Agapostemon sericeus was previously known as Agapostemon radiatus (Say) (e.g.,
Females of Agapostemon femoratus Crawford, primarily a western species not recorded east of New Mexico, Colorado and Wyoming by
The female of Agapostemon splendens can be recognized by the combination of the metallic green metasoma (as in Fig.
Male Agapostemon splendens can be recognized from all other midwestern Agapostemon by their very enlarged hind femur, with the length twice the width (Fig.
Some previous works (e.g.,
The females of Agapostemon texanus have the metasoma metallic green (Fig.
Male Agapostemon texanus have S3 and S4 with a low transverse swelling and generally have distinct yellow marks on the apical sterna (Fig.
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.
Agapostemon texanus and Agapostemon angelicus largely do not overlap in range in the midwestern region, though
Females of Agapostemon virescens are the only midwestern species that has the metasoma dark (Fig.
Males of Agapostemon virescens can be recognized by the lack of a transverse swelling on S4 (Fig.
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
The genus Augochlorella can be recognized by the combination of a normal oval-shaped tegula (as in Fig.
Key to females
1 | Propodeal triangle with striae continuing to posterior margin, or very nearly so (Fig. |
aurata (Smith) |
– | Propodeal triangle with striae not reaching posterior margin, leaving a distinct smooth portion (Fig. |
persimilis (Viereck) |
Key to males
1 | Inner edge of hind basitarsus with hairs on apical two-thirds only slightly elongate, their length about equal to width of basitarsus (Fig. |
aurata (Smith) |
– | Inner edge of hind basitarsus with hairs on apical two-thirds distinctly elongate, their length distinctly longer than width of basitarsus (Fig. |
persimilis (Viereck) |
Augochlorella characters A Augochlorella aurata female face B Augochlorella persimilis female face C–E Augochlorella aurata female propodeal triangles demonstrating the range of variation C apical margin carinate D apical margin largely lacking a carina E apical margin completely lacking a carina and striae partially removed from margin F–H Augochlorella persimilis female propodeal triangles demonstrating the range of variation F propodeum with narrow smooth margin G propodeum with broad smooth margin H propodeum with narrow and irregular smooth margin I Augochlorella aurata male hind basitarsus J Augochlorella persimilis male hind basitarsus. Scale bars: 1 mm (A, B); 500 µm (C–H, I, J).
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.
Female Augochlorella aurata are also often confused with Augochlora pura, but Augochlorella aurata have the paraocular lobes less protuberant (Fig.
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.
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.
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
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
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.
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.
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 (
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.
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.
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.
Key to females
1 | Vertex (in frontal view) rising above ocelli by at least one ocellar diameter (Fig. |
humeralis (Patton) |
– | Vertex (in frontal view) not rising above ocelli (Fig. |
2 |
2 | T2 hair fringe on apical margin with thickened flattened hairs unlike the hairs elsewhere on T2, and arranged closely together and appearing like the teeth of a comb along the apical margin of T2 (Fig. |
metallica (Fabricius) |
– | T2 hair fringe on apical margin with hairs identical to the hairs elsewhere on T2 (Fig. |
viridula (Smith) |
Key to males
1 | Vertex (in frontal view) rising above ocelli by at least one ocellar diameter (Fig. |
humeralis (Patton) |
– | Vertex (in frontal view) flattened, not rising above ocelli (Fig. |
2 |
2 | T2 with apical fringe of distinctly thickened hairs (Fig. |
metallica (Fabricius) |
– | T2 with apical fringe composed of unthickened hairs (Fig. |
viridula (Smith) |
Augochloropsis female characters A Augochloropsis humeralis face B Augochloropsis metallica face C Augochloropsis viridula face D Augochloropsis humeralis well-developed, 90-degree pronotal flange indicated by red arrow E Augochloropsis metallica obtuse pronotal flange indicated by red arrow F Augochloropsis viridula obtuse pronotal flange indicated by red arrow G Augochloropsis humeralis metasoma H Augochloropsis metallica metasoma I Augochloropsis viridula metasoma. Scale bars: 1 mm (A–F); 2 mm (G–I).
Augochloropsis female T2 hair fringes A Augochloropsis metallica with thickened hairs B Augochloropsis viridula with unthickened, slender hairs C Augochloropsis sp. (likely undescribed species from Eastland, Texas) with intermediate hairs. Note the specimens in A and B have their metasomas stretched out, revealing the brown basal part of the tergum that is normally hidden under the preceding tergum; this was done to increase contrast of the hairs and make the differences clearer. Scale bars: 1 mm, all images at the same scale.
Augochloropsis male characters A Augochloropsis humeralis face B Augochloropsis metallica face C Augochloropsis viridula face D Augochloropsis humeralis well-developed, 90-degree pronotal flange indicated by red arrow E Augochloropsis metallica obtuse pronotal flange indicated by red arrow F Augochloropsis viridula obtuse pronotal flange indicated by red arrow G Augochloropsis humeralis metasoma H Augochloropsis metallica metasoma I Augochloropsis viridula metasoma. Scale bars: 1 mm (A–F); 2 mm (G–I).
Augochlora humeralis
Patton, 1879: 365 ♀♂. Lectotype: ♀ USA, North-western Kansas, 8 Sep 1877 leg. S.W. Williston, on goldenrod [
Agapostemon caeruleus
Ashmead, 1890: 7 ♂ (not ♀) (syn.
Augochlora sumptuosa bolliana
Cockerell, 1909: 31 ♀ (syn. Under Augochloropsis caerulea by
Halictus (Augochlora) pattoni
Vachal, 1903: 132 (proposed replacement name for humeralis Patton; syn. by
Augochlora (Augochloropsis) humeralis
(in
Augochloropsis caerulea
(in
Augochloropsis humeralis
(in
Augochloropsis (Paraugochloropsis) sumptuosa
(in
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
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.
What has previously been called Augochloropsis sumptuosa by
Two syntypes of Augochloropsis humeralis (1 male and 1 female) were located in the
Augochlora sumptuosa bolliana Cockerell is from Texas and was synonymized with Augochloropsis sumptuosa by
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.
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 ♀ [
Andrena metallica
Fabricius, 1793: 309 ♀. Holotype: ♀ “America” [NHMD 308680]. Images examined by ZP and MA (Fig.
Augochlora fervida
Smith, 1853: 81 ♂ (syn. [under cuprea] by
Augochlora (Augochloropsis) cleomis
Titus, 1901: 135 ♀♂ (syn. by
Halictus chorisis
Vachal, 1903: 136 ♀ (syn. by
Megilla metallica
(in
Augochlora fervida
(in
Augochloropsis cuprea
(in
Augochloropsis (Paraugochloropsis) metallica metallica
(in
Augochloropsis (Paraugochloropsis) metallica
(in
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
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
The holotype of Augochloropsis metallica (Fig.
We define Augochloropsis metallica in a much more restricted sense than previous authors, who lumped multiple taxa under Augochloropsis metallica (e.g.,
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
Another synonym with issues is Augochloropsis chorisis Vachal, which was originally synonymized under Augochloropsis cuprea by
Augochlora (Augochloropsis) cleomis was described from a male and female specimen from near Fort Collins, Colorado (
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.
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 ♀ [
Augochlora viridula
Smith, 1853: 81 ♂. Holotype: ♂ USA, New York, Trenton Falls [
Augochlora lucidula
Smith, 1853: 81 ♀ (syn.
Halictus (Augochlora) viridissimus Viereck, 1910: 688 (proposed replacement name for viridula Smith, preoccupied in Halictus).
Augochlora viridula
(in
Augochloropsis cuprea
(in
Augochloropsis viridula
(in
Augochloropsis metallica
(Fabricius) (in
Augochloropsis (Paraugochloropsis) metallica fulgida
(in
Augochloropsis (Paraugochloropsis) fulgida
(in
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.
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
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.
Augochloropsis viridula is a polylectic ground-nester. Nesting and sociality were documented by
Augochloropsis viridula occurs throughout the eastern United States (Fig.
This species has historically been referred to as Augochloropsis metallica fulgida sensu
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
That Augochloropsis viridula (Smith) and Augochloropsis lucidula (Smith) were different sexes of the same species was recognized by
In the generic revision of augochlorine bees by
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 ♀ [
Augochlora anonyma
Cockerell, 1922: 15 ♀. Holotype: ♀ USA, Florida, No Name Key [
Augochloropsis cuprea
(in
Augochloropsis (Paraugochloropsis) anonyma
(in
Both sexes of Augochloropsis anonyma can be recognized by the short propodeal triangle, which is impressed and narrower than the metanotum (Fig.
Augochloropsis anonyma is also similar to Augochloropsis fulgida because both share the character of dark hairs on the metasoma (see Fig.
Augochloropsis anonyma is known from the far southeastern US, and we have examined material from Florida and Georgia (Fig.
Augochlora cuprea
Smith, 1853: 79 ♀. Images examined by ZP and MA (Fig.
Augochloropsis cuprea
(in
The type of Augochloropsis cuprea (Fig.
Augochloropsis cuprea was considered a junior synonym of Augochloropsis metallica metallica by both
The range of Augochloropsis cuprea is unclear as
Augochlora fulgida
Smith, 1853: 79 ♀. Holotype: ♀ USA, Florida, St. John’s Bluff, East Florida [NHMUK014024970]. Images examined by ZP and MA (Fig.
Augochlora fulgida
(in
Augochloropsis cuprea
(in
We define Augochloropsis fulgida differently than previous workers because examination of the type specimen (Fig.
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.
Augochlora sumptuosa Smith, 1853: 82 ♀. Syntype(s?): ♀ North America. Type or types missing and presumed lost.
Augochlora lacustris
Cockerell, 1922: 14 ♀ (syn.
Augochlora floridica
Cockerell, 1922: 14 ♂ (syn.
Augochlora sumptuosa
(in
Augochlora humeralis
(in
Augochloropsis caerulea
(in
Augochloropsis (Paraugochloropsis) sumptuosa
(in
Augochloropsis sumptuosa is most similar to Augochloropsis humeralis (refer to the diagnosis of that species to separate them).
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
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
We are aware of at least four additional potential species of Augochloropsis in the United States. We are listing them here in order to alert readers to their presence, as many have been incorrectly lumped together under existing species, particularly Augochloropsis metallica. However, we do not treat them further. We lack sufficient material of these species, and it is unknown whether they are undescribed or not, as they may be described from Mexico or they may be one of the many poorly known species described by Cockerell. The potential species and their locations include:
Here, we have revised the Augochloropsis of the Midwest and made additional changes to the Augochloropsis of the broader United States. This work will allow for the confident identification of the species in the midwestern United States and allow the species’ ranges to be better understood. However, there are areas of the southern United States (particularly Florida and Texas) where any Augochloropsis identifications must be undertaken with great care due to the number of undescribed or unknown species. We estimate there are an additional four species of Augochloropsis in the United States that are unknown or undescribed, not counting Augochloropsis fulgida, which is only known from the type and has the male now unknown. In addition, more work needs to be done to check the status of some of the current synonyms of Augochloropsis sumptuosa and Augochloropsis humeralis from Texas and Florida. Even the genus name may change at some point, as
The taxonomic changes and identification resources provided here will allow for more accurate identification of Augochloropsis and the other shiny green Halictinae. However, similar to the situation in Augochloropsis, more taxonomic work is still needed in the other shiny green Halictinae. For example the Agapostemon of the United States were last revised 50 years ago (
Our work also demonstrates the difficulty, indeed the futility, of attempting to monitor many bee groups that are in taxonomic disarray (
Comparison of names and species concepts applied over various keys and revisions of Augochloropsis. Dashes indicate that the species was not treated by the author.
|
|
|
|
Current name |
---|---|---|---|---|
fervida (Smith) | cuprea (Smith) | cuprea (Smith) | metallica metallica (Fabricius) | metallica (Fabricius) |
– | – | cuprea (Smith) | ||
– | – | fulvofimbriata (Friese) | ||
viridula (Smith) | viridula (Smith) | metallica fulgida (Smith) | viridula (Smith) | |
– | – | fulgida (Smith) | ||
– | – | anonyma (Cockerell) | anonyma (Cockerell) | |
– | caerulea (Ashmead) | humeralis (Patton) | sumptuosa (Smith) | humeralis (Patton) |
– | – | sumptuosa (Smith) |
We gratefully acknowledge the work of many people for loaning specimens and especially for tracking down and photographing the various type specimens: we thank Joseph Monks (