Oligochlora Engel

ENGEL, MICHAEL S., 2000, Classification Of The Bee Tribe Augochlorini (Hymenoptera: Halictidae), Bulletin of the American Museum of Natural History 2000 (250), pp. 1-89 : 47-51

Subgenus Oligochlora Engel s.s.

Oligochlora Engel, 1996b: 336 . Type species: Oligochlora eickworti Engel, 1996b , original designation.

DIAGNOSIS: This subgenus differs from Soliapis by the presence of an acarinarium and the obsolescent anteior border of the basitibial plate. The only other augochlorine with an acarinarium is the South American genus Thectochlora . Oligochlora is distinguished from Thectochlora by an unproduced pronotal lateral angle, absence of a lamella on the pronotal dorsal ridge, a broadly rounded mesoscutum, absence of a dorsal hook on the mesotrochanter, and abence of dense tomentum on the basal area of the propodeum.

DESCRIPTION: As for the genus with the following additions: Female. Basitibial plate bordered posteriorly, margin obsolete ante- riorly. Anterior surface T1 weakly modified into an acarinarium. Male. Unknown.

REVISIONS: At present this group contains three species. The subgenus was newly diagnosed by Engel (1997a) based on the discovery of a third species.

PALEOBIOLOGY: Two of the three species of Oligochlora are associated with astigmatid mites (Engel, 1996b; Fain et al., 1999). Mitebee associations are common, although poorly understood (Eickwort, 1979b, 1994; Fain et al., 1999). Oligochlora , like the genus Thectochlora (see below), possesses an acarinarium on the first metasomal tergum suggesting a mutualistic association between these ancient bees and mites.

Soliapis , new subgenus Figure 82

TYPE SPECIES: Oligochlora (Soliapis) rozeni , new species (appendix 1).

DIAGNOSIS: See Diagnosis for Oligochlora s.s. (above).

DESCRIPTION: As for the genus with the following additions: Female. Basitibial plate strongly bordered on all sides. Anterior surface of T1 unmodified, not developed into an acarinarium. Male. Unknown.

ETYMOLOGY: The subgeneric name is a combination of the Latin words sola (meaning ‘‘alone’’) and apis (meaning ‘‘bee’’) and is a reference to the lack of an acarinarium and thereby the mutualism with mites in these species.

REVISIONS: Two species are included in this subgenus. The first is described in appendix 1 as the type species of the group while a second is treated by Engel and Rightmyer (in press).

Genus Paroxystoglossa Moure

Paroxystoglossa Moure, 1940: 59 . Type species Oxystoglossa jocasta Schrottky, 1911 , original designation.

Paraoxystoglossa Roubik, 1989: 392. Lapsus calami.

DIAGNOSIS: Refer to the Diagnosis for Megaloptilla (above).

DESCRIPTION: Female. Mandible with moderate subapical tooth. Labral distal process narrowly triangular; basal elevation orbicular; teeth weak. Prementum not greatly

elongate. Galeal apex rounded; galeal comb absent; galeal base extending to stipital base. Hypostomal ridge carinate; anterior angle rounded. Length of malar space less than basal mandibular width. Epistomal sulcus obtuse. Ocelli not greatly enlarged; ocellar furrow absent. Vertex not expanded or ridged behind ocelli. Preoccipital ridge angled, distinctly not carinate. Pronotal lateral angle variable; dorsal ridge carinate; lateral ridge rounded to angled. Mesoscutal anterior bor- der narrowed; mesoscutal lip angled. Tegula oval. Anterior basitarsal brush present. Basitibial plate bordered on all sides. Inner hind tibial spur serrate. Apex of marginal cell truncate. Distal hamuli with irregular spacing pattern. Basal area of propodeum weakly striate to rugulose; propodeal pit narrow. Male. Mandible simple. Labrum with distal process; basal area notched. Antenna long, extending back to propodeum; F2 longer than F1. Inner hind tibial spur serrate. Metasoma oval to slightly elongate. Apical margins of S4–S6 emarginate. Apical margins of S7 and S8 with median projection. Proctiger unmodified. Gonobasal bridge narrow; dorsal lobes strong. Parapenial lobe and basal process of gonostylus absent. Ventral surface of penis valve with prong.

REVISIONS: The genus was revised by Moure (1960), who recognized nine species.

BIOLOGY: The nesting biology of Paroxystoglossa has been studied by Michener and Lange (1958a, 1958b) and Michener and Seabra (1959).

DISTRIBUTION: Species of the genus range in Argentina, southern Brazil, and Paraguay.

Genus Pereirapis Moure Figures 27 , 49 , 52 , 62

Pereirapis Moure, 1943a: 461 . Type species: Pereirapis rhizophila Moure, 1943a [= Halictus semiauratus Spinola, 1853 ], monobasic and original designation.

Pereirapsis Alves dos Santos, 1997: 6. Lapsus calami.

DIAGNOSIS: Pereirapis generally resembles species of the more common genera Augochlorella and Augochlora . It can be separated from the latter genus by the absence of a deeply acute epistomal sulcus that projects into the base of the clypeus and by the acute marginal cell apex. Pereirapis differs from Augochlorella in the presence of a large basal lobe on the inner metatibial spur. Pereirapis are small, brilliant metallic bees that are frequently green, with some populations slightly bluish and others more cupreous.

DESCRIPTION: Female. Mandible with moderate to strong subapical tooth. Labral distal process narrowly triangular; basal elevation orbicular; teeth strong. Prementum not greatly elongate. Galeal apex rounded; galeal comb absent; galeal base extending to base of stipes. Hypostomal ridge carinate; anterior angle rounded. Length of malar space less than basal mandibular width. Epistomal sulcus orthogonal. Ocelli not greatly enlarged; ocellar furrow absent. Vertex not expanded or ridge behind ocelli. Preoccipital ridge carinate. Pronotal lateral angle not produced, orthogonal to obtuse; dorsal ridge carinate; lateral ridge angled. Mesoscutal anterior border rounded; mesoscutal lip rounded. Tegula oval. Anterior basitarsal brush absent. Basitibial plate bordered on all sides. Inner hind tibial spur serrate, basally with an expanded tooth (fig. 49). Apex of marginal cell acute. Distal hamuli with irregular spacing pattern. Basal area of propodeum striate; propodeal pit narrow. Male. Mandible simple. Labrum with distal process; basal area not notched. Antenna extending back to scutellum; F2 approximately equal in length to F1. Inner hind tibial spur serrate. Metasoma oval. Apical margin of S4 concave. Apical margin of S5 unmodified. Apical margin of S6 emarginate. Apical margin of S7 unmodified. Apical margin of S8 with median process; spiculum narrow. Proctiger unmodified. Gonobasal bridge narrow; dorsal lobes strong. Basal process of gonostylus present; parapenial lobe absent. Ventral surface of penis valve unmodified.

REVISIONS: There is only the one included species (see appendix 1).

BIOLOGY: The nesting biology, phenology, and primitively eusocial behavior of Pereirapis was investigated by Oliveira Campos (1980). McGinley (1989) cited the description of immature stages for Pereirapis edentata (Michener) , but this is a species of Augochlorella .

DISTRIBUTION: Pereirapis ranges from Mexico eastward into the West Indies and south to southern Brazil.

Genus Pseudaugochlora Michener Figures 10 , 22–23 , 63 , 65 , 71

Caenaugochlora (Pseudaugochlora) Michener, 1954b: 77 . Type species: Halictus nigromarginatus Spinola, 1841 [= Megilla graminea Fabricius, 1804 ], original designation.

DIAGNOSIS: Pseudaugochlora is superficially most similar to Caenaugochlora . The genus can be separated from Caenaugochlora by the presence of a strong ridge on the vertex, pointed galeal apex, and hooked apex of F 11 in males.

DESCRIPTION: Female. Mandible with strong subapical tooth. Labral distal process narrowly triangular; basal elevation orbicular; teeth weak. Prementum not greatly elongate. Galeal apex acute; galeal comb absent; galeal base extending to stipital base. Hypostomal ridge carinate; anterior angle rounded. Length of malar space less than basal mandibular width. Epistomal sulcus orthogonal. Ocelli not greatly enlarged; ocellar furrow absent. Vertex with ridge behind ocelli. Preoccipital ridge rounded. Pronotal lateral angle not produced, obtuse; dorsal ridge carinate; lateral ridge rounded. Mesoscutal anterior border rounded; mesoscutal lip rounded. Tegula oval. Anterior basitarsal brush present. Basitibial plate bordered on all sides. Inner hind tibial spur pectinate. Apex of marginal cell truncate. Distal hamuli with irregular spacing pattern. Basal area of propodeum rugose; propodeal pit narrow. Male. Mandible simple. Labrum with distal process; basal area notched. Antenna extending back to scutellum or metanotum; F2 approximately equal in length to F1; F11 hooked at apex. Inner hind tibial spur serrate. Metasoma oval. Apical margins of S4 and S5 emarginate; S4 with medio-apical setal patch; S5 with lateroapical setal patches. Apical margin of S6 emarginate. Apical margin of S7 produced laterally. Apical margin of S8 with median process; spiculum broad. Proctiger with anal filaments. Gonobasal bridge narrow or membranous; dorsal lobes strong. Parapenial lobe present; basal process of gonostylus absent. Ventral surface of penis valve unmodified.

REVISIONS: The genus has never been revised. Moure and Hurd (1987) list seven named species while an eighth species is proposed in appendix 1.

BIOLOGY: The nesting biology of P. graminea and P. sordicutis (Vachal) has been investigated by Michener and Kerfoot [1967 as species of Pseudaugochloropsis , the latter under the name P. nigerrima (Friese) ].

DISTRIBUTION: Pseudaugochlora extends from the southwestern United States to Argentina and east into the West Indies.

COMMENTS: This group has generally been known under the name Pseudaugochloropsis by recent authors (e.g., Eickwort, 1967 Michener, 1974; Eickwort and Sakagami 1979; Moure and Hurd, 1987; Roubik, 1989 Michener et al., 1994; Radchenko and Pesenko, 1994b; Griswold et al., 1995; Danforth and Eickwort, 1997). However, the type species of Pseudaugochloropsis is a species of Augochloropsis (Paraugochloropsis) , and thus the valid name for the genus is Pseudaugochlora (see also discussions in Michener, 1954b, 1994). As if this confusion were not enough, the type species of this genus has been consistently cited as Halictus nigromarginatus Spinola, 1851 (Eickwort, 1969a Michener, 1994, 1997), which is obviously a homonym of the actual type species of Pseudaugochlora (namely, H. nigromarginatus Spinola, 1841 ). Spinola’s 1851 nigromarginatus is in actuality a species of Caenohalictus (= Caenohalictus oblitus Moure and Hurd 1987 ).

Genus Temnosoma Smith

Figures 7–9 , 20 , 30 , 40 , 56–57 , 67 , 69 , 76

Temnosoma Smith, 1853: 38 . Type species: Temnosoma metallicum Smith, 1853 , monobasic.

Tamnosoma Taschenberg, 1883: 93. Lapsus calami.

Micraugochlora Schrottky, 1909a: 138 . Type species: Micraugochlora sphaerocephala Schrottky, 1909a , monobasic.

Temnosoma (Temnosomula) Ogloblin, 1953: 2 Type species: Temnosoma (Temnosomula) platensis Ogloblin, 1953 [= Micraugochlora sphaerocephala Schrottky, 1909a ], monobasic and original designation. Proposed as new again in Ogloblin, 1954.

Themnosoma Sakagami, 1979: 83. Lapsus calami

Microaugochlora Schlindwein, 1998: 52. Lapsus calami.

DIAGNOSIS: This is one of the most distinctive genera of Augochlorini . The coarse sculpturing of the body gives these cleptoparasitic bees the general appearance of chrysidid wasps. Among augochlorine genera, Temnosoma ’s appearance comes closest to species of Glyptochlora . Temnosoma can be separated from Glyptochlora by an oval tegula, absence of a scopa, the depressed apical margins of the first two metasomal terga, absence of a medio-apical cleft in the fifth metasomal tergum of females, and the presence of a broad division in the seventh metasomal tergum of males, among other characters.

DESCRIPTION: Female. Body coarsely punctured. Mandible simple. Labral distal process quadrate; basal elevation weakly bilobed; keel absent; teeth absent. Prementum not greatly elongate. Galeal apex rounded; galeal comb absent; galeal base extending to base of stipes. Hypostomal ridge carinate; anterior angle rounded. Length of malar space less than basal mandibular width. Epistomal sulcus obtuse. Ocelli not greatly enlarged; ocellar furrow absent. Vertex expand- ed behind ocelli. Preoccipital ridge sharply angled and weakly carinate. Pronotal lateral angle not produced, obtuse; dorsal ridge carinate; lateral ridge rounded. Mesoscutal anterior border rounded; mesoscutal lip weak and rounded. Tegula oval; coarsely punctured. Anterior basitarsal brush absent. Mesofemoral brush absent. Scopa absent. Penicillus absent. Basitibial plate absent. Inner hind tibial spur serrate. Apex of marginal cell acute. Distal hamuli with irregular spacing pattern. Basal area of propodeum strongly striated. Apical margins of T1 and T2 strongly depressed and rimmed; apical margin of T5 without cleft. Male. Mandible simple. Labrum with quadrate distal process. Antenna extending back to mesoscutum; F2 longer than F1. Inner hind tibial spur serrate. Metasoma oval. Apical margin of T7 with broad division. Apical margins of S4 and S5 unmodified. Apical margin of S6 emarginate. Apical margin of S7 medially unsclerotized. Apical margin of S8 with median projection; spiculum narrow. Proctiger unmodified. Gonobasal bridge broad; dorsal lobes weak. Gonostylus with parapenial lobe present; basal process absent; ventral process greatly expanded with long setae; dorsal process reduced, small ridge with setae. Ventral surface of penis valve unmodified, with large dorsal process.

REVISIONS: Friese (1925) revised the species of Temnosoma known at the time recognizing six species. Moure and Hurd (1987) list seven species. There has been no modern treatment of the genus.

BIOLOGY: The biology of Temnosoma remains unstudied, but the species are presumably parasitic, perhaps on species of the genus Augochloropsis or Augochlora (Michener, 1978a) .

DISTRIBUTION: Temnosoma occupies a wide distribution with species found from as far south as northern Argentina, throughout South and Central America, north to Arizona (Timberlake, 1958), as well as occurring in Cuba and Jamaica (Eickwort, 1988). One potential host, Augochloropsis , does not occur in the West Indies, but the other, Augochlora , does.

Genus Thectochlora Moure Figures 1 , 42 , 47 , 53–55 , 72

Thectochlora Moure, 1940: 51 . Type species: Halictus alaris Vachal, 1904 , monobasic and original designation.

DIAGNOSIS: Thectochlora resembles to some degree the genera of the Augochlora - group (e.g., Augochlorella , Pereirapis ), but differs most notably in the presence of a well-developed acarinarium on the first metasomal tergum, presence of a strong dorsal hook on the mesotrochanter, a lamellate dorsal pronotal ridge, and the presence of dense tomentum on the basal area of the propodeum. The genus also differs in the combination of an obtuse epistomal sulcus and a pectinate inner metatibial spur.

DESCRIPTION: Female. Mandible with weak subapical tooth. Labral distal process narrowly triangular; basal elevation orbicular; teeth strong. Prementum not greatly elongate. Galeal apex rounded; galeal comb absent; galeal base extending to base of stipes. Hypostomal ridge carinate; anterior angle rounded. Length of malar space less than basal mandibular width. Epistomal sulcus obtuse. Ocelli not greatly enlarged; ocellar furrow absent. Vertex not expanded or ridged behind ocelli. Preoccipital ridge rounded. Pronotal lateral angle produced, obtuse; dorsal ridge lamellate; lateral ridge rounded. Mesoscutal anterior border narrowed; mesoscutal lip lamellate. Tegula oval. Anterior basitarsal brush absent. Mesotrochanter with strong dorsal hook. Basitibial plate rimmed on posterior edge, anterior border obsolete. Inner hind tibial spur pectinate. Apex of marginal cell truncate. Distal hamuli with irregular spacing pattern. Basal area of propodeum striate, extremely short, covered in dense tomentum. Anterior surface T1 modified into an acarinarium (a depressed shiny area bordered basally and partly on lateral margins by long plumose hairs). Male. Mandible simple. Labrum with weak distal process; basal area not notched. Antennae extending just beyond posterior margin of propodeum; F2 much longer than F1. Inner hind tibial spur serrate. Metasoma oval. S4 with lateral processes on apical margin possessing strong setae at apices; central patch of dense setae on disc. Apical margin of S5 emarginate. Apical margin of S6 emarginate. S7 unmodified. Apical margin of S8 with medial process; spiculum narrow. Proctiger unmodified. Gonobasal bridge narrow; dorsal lobes strong. Gonostylus with parapenial lobe present; basal process absent. Penis valve with prong on ventral surface.

REVISIONS: There is currently only one species recognized in the genus.

BIOLOGY: Nothing is known of Thectochlora biology, aside from its association with a specific genus of acarid mites — Thectochloracarus (Fain et al., 1999) . The modified anterior surface of T1 for the transport of mites suggests that this is perhaps a mutualistic relationship.

DISTRIBUTION: Thectochlora occurs in northern Argentina, southern Brazil, Paraguay, and Guyana.

Figures

Fig. 82. Photomicrograph of holotype female of Oligochlora (Soliapis) rozeni, new subgenus and new species, in Dominican amber (photograph by author).

Figs. 26–29. Head modifications. 26. Preoccipital area of Corynura (Corynura) chilensis (Spinola). 27. Preoccipital area of Pereirapis semiaurata (Spinola), carina present. 28. Compound eye of C. (Callistochlora) chloris (Spinola), elongate setae. 29. Compound eye of Augochlora (Augochlora) pura (Say), minute setae.

Figs. 46–51. Spurs and antenna cleaners of female Augochlorini. 46. Inner hind tibial spur of Chlerogella sp. 47. Inner hind tibial spur of Thectochlora alaris (Vachal). 48. Inner hind tibial spur of Augochlora (Augochlora) pura (Say). 49. Inner hind tibial spur of Pereirapis semiaurata (Spinola). 50 Antenna cleaner of Chlerogella sp. 51. Antenna cleaner of Megommation (Cleptommation) minutum (Friese).

Figs. 52–55. Tergal modifications of female Augochlorini. 52. Anterior surface of T1 of Pereirapis semiaurata (Spinola). 53. Anterior surface of T1 of Thectochlora alaris (Vachal) with acarinarium, mites removed. 54. Dorsal aspect of T1 of T. alaris with acarinarium, mites removed. 55. Same view as in figure 54 with mites present.

Figs. 60–63. Features of male Augochlorini. 60. Labrum of Caenaugochlora (Caenaugochlora) costaricensis (Friese). 61. From left to right: apex of scape, pedicel, F1, F2, and basal margin of F3 for Corynura (Callistochlora) prothysteres (Vachal). 62. From bottom to top: scape, pedicel, F1, F2, F3, and basal margin of F4 for Pereirapis semiaurata (Spinola). 63. Distal flagellomeres of Pseudaugochlora graminea (Fabricius), F11 hooked at apex.

Figs. 7–12. Head modifications of female Augochlorini. 7. Temnosoma metallicum Smith, dorsal aspect. 8. T. metallicum, profile. 9. T. metallicum, vertex. 10. Pseudaugochlora graminea (Fabricius), vertex. 11. Megalopta (Megalopta) genalis Meade-Waldo, vertex. 12. Clypeal apex of Megommation (Megaloptina) ogilviei (Cockerell).

Figs. 21–25. Labiomaxillary complex structures of Augochlorini. 21. Maxilla in profile Corynura (Corynura) chilensis (Spinola). 22. Maxilla of Pseudaugochlora graminea (Fabricius). 23. Galeal apex of P. graminea. 24. Galeal apex of C. chilensis. 25. Galeal comb of C. chilensis.

Figs. 64–67. Metasomal modifications of male Augochlorini. 64. Relatively unmodified sterna (S5– S6) of Neocorynura papallactensis Engel. 65. S3–S6 of Pseudaugochlora graminea (Fabricius). 66. S3– S6 of Megalopta (Megalopta) genalis Meade-Waldo. 67. T7 of Temnosoma metallicum Smith.

Figs. 68–71. Hidden and fused male sterna S7–S8; spicula are on the left margin and apical processes are shown on the right. 68. Augochloropsis (Paraugochloropsis) sumptuosa (Smith). 69. Temnosoma metallicum Smith. 70. Megalopta (Megalopta) genalis Meade-Waldo. 71. Pseudaugochlora graminea (Fabricius).

Figs. 17–20. Labra of female Augochlorini. 17. Augochlora (Augochlora) pura (Say). 18. Rhinocorynura briseis (Smith). 19. Megalopta (Megalopta) genalis Meade-Waldo. 20. Temnosoma metallicum Smith.

Figs. 30–33. Mesosomae of female Augochlorini. 30. Temnosoma metallicum Smith, profile. 31 Rhinocorynura briseis (Smith), profile. 32. Mesoscutum of Megommation (Cleptommation) minutum (Friese), dorsal aspect. 33. Mesoscutum of R. briseis, dorsal aspect, pronotal dorsal lamellae evident on upper corners of mesosoma.

Figs. 38–41. Features of female Augochlorini. 38. Scutellum, metanotum, and basal area of propodeum of Megommation (Cleptommation) minutum (Friese). 39. Scutellum, metanotum, and basal area of propodeum of Chlerogella sp. 40. Hind leg of Temnosoma metallicum Smith, scopa absent from femur and tibia. 41. Hind leg of Augochlora (Augochlora) pura (Say), scopa on femur and tibia.

Figs. 56–59. Metasomal modifications of female Augochlorini. 56. Metasoma of Temnosoma metallicum Smith, dorsal aspect. 57. Metasoma of T. metallicum, profile. 58. S1 of Augochlora (Augochlora) pura (Say), median apical ridge. 59. Apex of T5 of Megalopta (Megalopta) genalis Meade- Waldo, medioapical cleft of pseudopygidial area.

Figs. 72–77. Male genitalic structures. 72. Genital capsule of Thectochlora alaris (Vachal), ventral aspect. 73. Genital capsule of Megalopta (Megalopta) genalis Meade-Waldo, ventral aspect. 74. Genital capsule of Rhinocorynura briseis (Smith), ventral aspect. 75. Gonocoxite-gonostylus junction in R. briseis depicting basal gonostylar process on ventral surface, process with strong setae at apex. 76. Penis valve of Temnosoma metallicum Smith, profile, with large dorsal processes. 77. Basal gonostylar process of Corynura (Callistochlora) prothysteres (Vachal) lacking setae, partially hidden by setose ventral

Figs. 1–6. Heads of Augochlorini, frontal aspect. 1. Thectochlora alaris (Vachal), female. 2. Rhinocorynura briseis (Smith), female. 3. Megommation (Cleptommation) minutum (Friese), female. 4 Chlerogella sp., female. 5. M. (Megaloptina) ogilviei (Cockerell), female. 6. Megalopta (Megalopta genalis Meade-Waldo, male.

Figs. 42–45. Leg modifications of female Augochlorini. 42. Inner surface of mesotrochanter of Thectochlora alaris (Vachal) showing hook on dorsal surface. 43. Basitibial plate of Augochlora (Augochlora) pura (Say). 44. Basitibial plate of T. alaris. 45. Outer surface of metatibial base of Megommation (Cleptommation) minutum (Friese), showing absence of basitibial plate.