Classification of the Bee Tribe Augochlorini (Hymenoptera: Halictidae)

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provided by American Museum of Natural History Scientific Publications CLASSIFICATION OF THE BEE TRIBE AUGOCHLORINI (HYMENOPTERA: HALICTIDAE)

MICHAEL S. ENGEL Research Scientist Division of Invertebrate Zoology, American Museum of Natural History

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Number 250, 90 pages, 82 ﬁgures, 4 tables Issued April 7, 2000 Price: $8.10 a copy

Copyright ᭧ American Museum of Natural History 2000 ISSN 0003-0090 FRONTISPIECE. Dorsal habitus of Neocorynurella seeleyi Engel and Klein (from Engel and Klein, 1997).

‘‘Go to the bee, and learn how diligent she is, and what a noble work she produces, whose labors kings and private men use for their use, she is desired and honored by all, and though weak in strength she values wisdom and prevails.’’ —Proverbs, Chapter 6, Septuagint1 version of the Bible

1 The Septuagint, sometimes known as Interpretatio secundum septuaginta seniores (Transl.: ‘‘Translation Accord- ing to 70 Elders’’)or␩ ␧␬␬␭␩␴␫␣␴␶␫␬␩ ␧␬␦␱␴␫␵ (Transl.: ‘‘The Ecclesiastic Edition’’), is the oldest Greek version, perhaps also the oldest extant version, of the Old Testament; believed to have been translated in Alexandria at the request of Ptolemy II Philadelphus (285Ð247 B.C.) by 70Ð72 Jewish scholars. The number 70 for the scholars would have been based on the tradition that 70 elders accompanied Moses when he received the commandments, while it is believed by some that the number was increased to 72, with six scholars representing each of the 12 Hebrew tribes. Later versions, such as those based on the Vulgate Latin translation, remove this portion of this text and retain only the reference to the ‘‘ant.’’ CONTENTS Abstract ...... 4 Introduction ...... 5 Taxonomic Review ...... 5 The Present Study ...... 7 Acknowledgments ...... 7 Materials and Methods ...... 7 General Morphology ...... 10 Females ...... 10 Males ...... 23 Systematics ...... 26 Tribe Augochlorini Beebe ...... 26 Subtribe Augochlorina Beebe ...... 29 Subtribe Corynurina, new subtribe ...... 51 Key to Genera and Subgenera of Augochlorini ...... 55 Cladistics ...... 60 Description of Characters ...... 60 Data Analysis ...... 63 Discussion ...... 65 Phylogenetic Relationships ...... 65 Ethological Evolution ...... 70 Fossil History ...... 71 References ...... 71 Appendix 1 ...... 79 Appendix 2 ...... 87 Appendix 3 ...... 89 Appendix 4 ...... 90

3 4 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

ABSTRACT The halictid bee tribe Augochlorini is revised at the level of genus and subgenus. Forty- one genera and subgenera are recognized with two being extinct. New subgenera of Augo- chlora, Electraugochlora, and of Oligochlora, Soliapis, are described for new fossils from Tertiary amber of the Dominican Republic. Complete taxonomic histories are given along with information on revisions at the species level, biological studies, and general distribution for each family- and genus-group taxon. The following subgenera are newly synonymized: Ae- thechlora new junior subjective synonym of Oxystoglossella, Mycterochlora new junior subjective synonym of Augochlora, Vachalius new junior sybjective synonym of Neocorynurella, and Neocorynuroides new junior subjective synonym of Neocorynura. The group Glyptochlora is resurrected from synonymy as a valid subgenus of Augochloropsis. The males of Chlero- gella and Micrommation are described for the first time. A key to the genera and subgenera of the tribe is presented along with a key to the species of the rare Augochloropsis (Glypto- chlora). Suprageneric relationships in the tribe are explored cladistically. Characters of adult external morphology (72 characters) and ethology (12 characters) are coded for all recognized augochlorine genera as well as outgroups from the Halictini, Nomioidini, and Nomiinae. Results of the cladistic analysis are remarkably resolved but not very robust. The cladogram is summarized and infratribal groups defined. Most notably, the tribe is divided into two monophyletic subtribes: the Corynurina (new subtribe), containing the southern South American genera Corynura, Halictillus, Rhectomia, and Rhinocorynura; and the nominate subtribe, Augochlo- rina. Implications of the cladistic analysis on diversification in the Augochlorini and evolu- tionary patterns within the tribe are discussed. New distribution records are given for three species which extend the known ranges of their respective genera. Augochlora essequibensis is a new junior subjective synonym of A. nigrocyanea while A. cladopyga, A. seminigra, Augochlorella bidentata, Halictus caucasicus, H. cerasis, H. chrysaspis, H. myrrhites, H. simotes, and Pereirapis rhizophila are all new junior subjective synonyms of P. semiaurata. Megalopta intermedia, Augochlora nitidior, Augochlo- rella eusticta, Augochlorodes clementis, and Augochloropsis scabriceps are all recognized as nomina nuda. Vachalius cosmetor, from Colombia and Venezuela, is transferred to Neocory- nurella while Corynura biciliata, from Costa Rica, is recognized as a species of Halictini and is transferred to the genus Lasioglossum (Evylaeus) (new combinations). Four fossil and recent species new to science are described as Augochlora (Electraugochlora) leptoloba (fossil), Augochlora (Oxystoglossella) rightmyerae (extant), Oligochlora (Soliapis) rozeni (fossil), and Pseudaugochlora pulchra (extant). 2000 ENGEL: BEE TRIBE AUGOCHLORINI 5

INTRODUCTION Bees are one of the most ecologically im- in length, although some ‘‘giants’’ do occur portant groups of arthropods and are by far in the nocturnal genus Megalopta, etc. (ca. the most significant group of insect pollina- 18 mm) and minute species are known in tors. There are approximately 20,000 species Halictillus, etc. (ca. 3 mm). worldwide, with the greatest diversity found Many augochlorine species are gregarious, in xeric regions. The living bees are segre- nesting in societies ranging from communal gated into six families; Colletidae2, Halicti- to primitively eusocial, semisocial societies dae, Andrenidae, Melittidae, Megachilidae, apparently being the most common type and and Apidae; and together form a monophy- perhaps part of the augochlorine groundplan. letic group within the aculeate superfamily Nest architecture within the tribe is varied, Apoidea. Among the major lineages of bees, with nests ranging from simple tubular bur- the family Halictidae is one of the more bas- rows with lateral cells to more elaborate al, apparently being sister to all other bees chambers with supported clusters of brood except the most primitive family, Colletidae. cells constructed either in soil or rotting The family is relatively large, consisting of wood. This diversity has attracted the atten- approximately 3500 described species. Three tion of comparative ethologists and several main lineages are recognized within the fam- hypotheses have been presented to explain ily; the plesiomorphic and certainly paraphy- the patterns of change seen among augo- letic subfamily Rophitinae (ϭ Dufoureinae chlorine nests (e.g., Sakagami and Michener, of some authors), the Nomiinae, and the Ha- 1962; Eickwort and Sakagami, 1979). Like lictinae. The halictines, by far, dwarf the oth- most halictines, augochlorines are polylectic, er two subfamilies in size, and make up ap- visiting an array of flowers for resources; proximately 80% of the specific diversity for however, some modified foraging behaviors the family. The Halictinae is itself presently have arisen in the tribe. Most notably, the divided into three tribes: Nomioidini, Halic- transition to nocturnal flower visitation in tini, and Augochlorini. The present study fo- genera such as Megalopta is a peculiarity cuses on the last-named tribe and provides a among bees. True nocturnal behavior, not revised supraspecific classification. simply crepuscular, has apparently arisen at The Augochlorini is remarkable among least eight times among all bees; it is note- halictids for having its greatest diversity con- worthy that three of these derivations have centrated in the warm tropics of the New been within the Augochlorini (see Discus- World, while lineages of the others are di- sion). Likewise, based on their anatomy, spe- versified either in xeric, temperate, or cool cific groups appear to have specialized on tropical habitats. The tribe consists of ap- flowers with deep corollas (e.g., Chlerogas). proximately 525 valid species and is most noted for the brilliant metallic coloration, TAXONOMIC REVIEW which is frequently green (hence their name: Greek auge ϩ chloros meaning ‘‘shining The first known augochlorine species was green’’). Coloration throughout the tribe is described by Fabricius (1793) as Andrena quite variable, with a small number of spe- metallica (currently Augochloropsis metalli- cies being dull metallic or black (similar to ca). Fabricius went on to name the second most species of Halictini), others brown to augochlorine in 1804, but over a quarter of amber, while the majority are metallic red, a century would pass before another species gold, green, blue, violet, or any combination would be recognized (Halictus rubellus Hal- of these. Species are distributed from south- iday, 1836). From 1836 to the present, every ern Canada to northern Argentina and Chile decade has seen activity on the species-level and east into the West Indies. Most taxa are taxonomy of augochlorines, although after of a moderate size, being around 6Ð10 mm 1970 augochlorine taxonomy ground to a near halt with only two subgenera and two 2 I consider the Australian family Stenotritidae to be species named over the next quarter of a cen- a subfamily, at best, of Colletidae. tury (one of them, however, was merely a 6 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250 replacement name for a preoccupied species cific classification of the Apoidea in which of Caenaugochlora). Not until 1995 did he placed the then recognized augochlorine studies on the tribe resume. genera (namely Augochlora, Corynura, Me- At the generic level, most authors through- galopta, and Temnosoma) into two different out the nineteenth century followed Latreille groups: Augochlora, Corynura, and Mega- (1804) and placed augochlorines in the genus lopta (under the lapsus Megaloptera) were Halictus. It was Spinola (1851) who first pro- placed in the Halictinae (then a subfamily of posed a currently recognized augochlorine ge- Andrenidae) along with genera such as Aga- nus, Corynura. During the remainder of that postemon, Nomioides, and Systropha, where- decade, four other genus-group names were as the parasitic genus Temnosoma was proposed for augochlorine species (more than grouped with other parasitic short-tongued were proposed during the remainder of that bees (e.g., Sphecodes) in the subfamily Sphe- century). In the early twentieth century the codinae. Although Beebe (1925) later pro- generic classification of the Augochlorini ex- posed the family-group name Augochloridae, ploded with the number of genera more than the group as he conceived it was equivalent doubling (to 20) by 1910, mostly owing to to the Halictinae of Ashmead (1899). The the efforts of Schrottky. Although a few first suprageneric recognition of a group cor- groups were proposed over the intervening responding to the tribe of today was by years, it was not until the 1940s and 1960s Schrottky (1909b) who proposed the tribal that work on the tribe resumed with any con- name Oxystoglossini for those genera allied centration. At this time Moure proposed 18 to the genus Oxystoglossa (a junior synonym genera in connection with his studies designed of Augochlora); for the first time separating to monograph the New World halictine fauna. the augochlorines from Halictus and its al- In 1969, Eickwort (1969a) provided the first lied genera. At approximately the same time, synthesis of augochlorine genera; relegating Vachal was involved in developing his clas- several names to synonymy. Eickwort recog- sification of New World halictines (all under nized 24 genera and an additional seven sub- the genus Halictus!) in the form of a lengthy genera. At the time of his study, the genera identification key. Vachal (1911) proposed Chlerogella and Rhectomia were known only the division Halicti hexagoni which approx- on the basis of females, and the genus Chler- imately corresponded to Schrottky’s Oxy- ogas only on the basis of males. In the same stoglossini. Moure (1943a), perhaps unaware year, Moure (1969) described one additional of Beebe’s use of the name Augochloridae as genus, Micrommation, and later proposed two well as Schrottky’s tribal name, defined the new subgenera in Augochlora and Megom- tribe Augochlorini as new and to accommo- mation (Moure and Hurd, 1987), bringing the date the Halicti falcati subdivision of Vach- total composition of the Augochlorini to 25 al’s Halicti hexagoni. At the same time Mou- genera and nine additional subgenera. Since re proposed the tribe Augochloropsini and then several genera have been added to the the subfamily Megaloptinae for other subdi- tribe by the present author and coauthors as visions of Vachal’s group; the three supra- well as an additional genus, Vachalius,by generic groups together being equivalent to Moure (1999). the Augochlorini of today. Eickwort (1969b) Kirby (1802) divided the Linnean genus presented the first modern diagnosis of the Apis (Linnaeus, 1758), which was equivalent tribe, distinguishing it from the Halictini on to all six bee families of today, into short- the basis of both adult morphology and nest tongued and long-tongued bees [the Andre- architecture. Eickwort was apparently un- netae and Apiariae of Latreille (1802)]; a aware of both Beebe’s (1925) article and of higher classification which was perpetuated the older tribal name Oxystoglossini, using by Smith (1853, 1879). Under this classifi- Augochlorini as the valid name and attribut- catory structure the few augochlorines ing it to Moure. The usage of Augochlorini known at the time were placed in the Andre- has become universal and Oxystoglossini has netae along with the genus Halictus and, as not been used since Schrottky’s proposal of indicated earlier, frequently within that ge- the name in 1909. An application to suppress nus. Ashmead (1899) proposed a supraspe- Oxystoglossini has been made with the 2000 ENGEL: BEE TRIBE AUGOCHLORINI 7

I.C.Z.N. and is currently pending (Engel, ACKNOWLEDGMENTS 1999a). This work is adapted from a dissertation As of 31 December 1999, 697 species- submitted in partial fulfillment of the require- group and 64 genus-group (table 1) names ments for the degree of Doctor of Philosophy have been proposed in the Augochlorini, in- at Cornell University (Engel, 1998). I am including nomina nova, nomina nuda, and debted to my doctoral committee: James K. names newly proposed herein. Liebherr, Richard G. Harrison, Thomas D. Seeley, and Charles D. Michener. Jerome G. Rozen, Jr., David A. Grimaldi, and Molly G. THE PRESENT STUDY Rightmyer have, with great enthusiasm, provided me with constant encouragement and The present study provides a revised clas- support while at the AMNH. The numerous sification of the Augochlorini above the level curators (listed below) as well as a number of species, an attempt to elucidate phyloge- of individuals too numerous to mention gen- netic relationships among the supraspecific erously provided assistance and support; to taxa, and to provide commentary on the evo- each of them I am sincerely thankful. I am lution of the tribe. The classification devel- grateful for the influence of two men whose oped here recognizes 30 genera and an ad- Earthly labors have since come to a close: ditional 11 subgenera. Four genus-group taxa my late advisor, George C. Eickwort, and are newly placed into synonymy while a sub- Byron A. Alexander. During his life George genus of Augochloropsis is resurrected from was mentor to many and I am blessed that synonymy. Table 1 associates each genus- our paths crossed, even if only for a short group name with its current status. Mega- time. George and Byron were noteworthy re- loptilla, previously considered a subgenus of searchers, masterful teachers, inspiring men- Megommation, is elevated to generic rank as tors, and close friends. Lastly, I thank my it is not closely related to the later genus. The parents, Rev. A. G. Engel and Donna G. En- males of Chlerogella and Micrommation are gel, for their constant support and encouragement. described for the first time. By means of syn- Funding was provided by the following onymies, a taxonomic history is given for awards and agencies: Predoctoral Fellowship each family- and genus-group taxon. In ad- of the National Science Foundation, Ernst dition, for each genus information on the bi- Mayr Award of Harvard University’s Muse- ology and distribution of the included taxa is um of Comparative Zoology, CanaColl summarized and references provided for any Foundation of the Canadian National Collec- revisions or keys to species. A new key is tion, Short-term Fellowship of the Smithson- provided for the identification of all valid ian Tropical Research Institute, Kalbfleisch genera and subgenera. A cladistic analysis of Fund, Collection Study Grant of the AMNH, affinities among augochlorine genera is ex- Exploration Fund of the Explorer’s Club, the plored based on 84 characters of adult exter- Graduate School and Department of Ento- nal morphology and ethology. All suprage- mology of Cornell University. I am also neric groups are defined based on the clad- grateful to Robert G. Goelet, Chairman ogram and commentary provided on evolu- Emeritus of the AMNH Board of Trustees, tion within the tribe. for support of my studies at the AMNH.

MATERIALS AND METHODS

In the section below on Systematics, the betically under each subtribe; their order taxonomic history of each group is presented should not be taken as a reflection of rela- followed by a brief diagnosis separating the tionship. Phylogenetic relationships are elab- taxon from close relatives or superficially orated on in the Discussion. Following the similar genera. Genera are presented alpha- diagnosis for each genus a short description 8 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250 2000 ENGEL: BEE TRIBE AUGOCHLORINI 9 providing important morphological traits is Jr., and E. Quinter; ANSP, Philadelphia appended. Lastly, appended to each entry are Academy of Natural Sciences, Philadelphia, brief summaries on (1) the current classifi- Pennsylvania, D. Azuma; BMNH, The Nat- cation of each genus-group taxon, providing ural History Museum, British Museum, Lon- references to revisions, if they exist; (2) in- don, United Kingdom, G. Else, S. Lewis; formation and references to the biology of CMNH, Carnegie Museum of Natural His- each group; and (3) distribution information. tory, Pittsburgh, Pennsylvania, R. L. David- If, however, nothing is known concerning the son; CNC, Canadian National Insect Collec- biology for a given group, this section has tion, Ottawa, Canada, G. Gibson and L. Du- been omitted. A list of species examined dur- mouchel; CUIC, Cornell University Insect ing the course of this study to construct the Collection, Ithaca, New York, J. K. Liebherr cladistic matrix is presented in appendix 3. and E. R. Hoebeke; DZUP, Museu de En- In the descriptions the following abbrevia- tomologia Padre Jesu«s Santiago Moure, De- tions are used: F, flagellomere; S, metasomal partamento de Zoologia, Universidade Fed- sternum; T, metasomal tergum. The length of eral do Parana«, Curitiba, Brazil, J. S. Moure; the prementum and size of ocelli are charac- EMUS, U.S.D.A. Bee Biology and System- ters used to separate genera. An elongate pre- atics Laboratory, Utah State University, Lo- mentum is taken to mean that its length is gan, Utah, T. L. Griswold; FMNH, Field greater than seven times its width, and con- Museum of Natural History, Chicago, Illi- versely a prementum considered not elongate nois, P. P. Parrillo; FSCA, Florida State Col- has the length less than seven times its width. lection of Arthropods, Gainesville, Florida, J. Similarly, ‘‘ocelli greatly enlarged’’ is used Wiley and M. C. Thomas; GARM, Gabriel for those taxa in which the distance from the A. R. Melo Private Collection, Ribeiraõ upper margin of the compound eye to the lat- Preto, Brazil (formerly of SEMC); IIRB, eral ocellus is less than the diameter of the Museo de Invertebrados, Instituto de Inves- median ocellus. Further details on morpholog- tigacio«n de Recursos Biolo«gicos ‘‘Alexander ical terminology are provided below (under von Humboldt,’’ Santa Fe« de Bogota«, Co- General Morphology). lombia, F. Fernandez; INPA, Instituto Na- Specimens were prepared for scanning cional de Pequisas da Amazoˆnia, Manaus, electron microscopy as follows: (1) pinned Brazil, A. Y. Harada; IZAV, Universidad specimens were disarticulated and mounted Central de Venezuela, Maracay, Venezuela, directly onto carbon-covered stubs; (2) spec- J. A. Clavijo and M. A. Ganiani; LACM, imens originally stored in 70Ð90% ethanol The Natural History Museum of Los Angeles were disarticulated and then transferred step- County, Los Angeles, California, R. R. Snell- wise to 100% ethanol in which they were kept ing; MACT, Morone Amber Collection, Tu- for at least 24 hours, before transfer to hexa- rin, Italy, E. Morone (D. A. Grimaldi, methyldisalizane (HMDS) for one hour, and AMNH); MCZ, Museum of Comparative again to a fresh bath of HMDS. After the sec- Zoology, Cambridge, Massachusetts, P. Per- ond wash of HMDS, the specimens were kins and S. Cover; MEMU, Mississippi State placed under a fume hood and allowed to re- Insect Collection, Mississippi State, Missis- main there until all of the HMDS had evap- sippi, R. L. Brown; MIUP, Museo de Inver- orated (procedure further described by Brown, tebrados ‘‘G. B. Fairchild,’’ Universidad de 1993; Rumph and Turner, 1998; and other au- Panama«, Panama« City, Panama«, D. Quintero; thors). Each specimen was then mounted on MNHC, Museo Nacional de Historia Natu- an electron microscopy stub for study. ral, Havana, Cuba, J. A. Genaro; MNHN, The following institutions and curators Museum National d’Histoire Naturelle, Paris, provided material that was examined during France, J. Casevitz-Weulersse; MRSN, Spi- the course of this study. Abbreviations for nola Collection, Museo Regionale Scienze institutions are taken from Arnett et al. Naturali, Turin, Italy, P. L. Scaramozzino; (1993), except those collections not listed by MSUC, Michigan State University, Depart- them for which acronyms were newly cre- ment of Entomology Collection, East Lan- ated: AMNH, American Museum of Natural sing, Michigan, F. Stehr; PMAE, Provincial History, New York, New York, J. G. Rozen, Museum of Alberta, Edmonton, Canada, A. 10 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

Finnamore; SEMC, Snow Entomological eral de Viçosa, Viçosa, Brazil; UNCB, Univ- Collection, Natural History Museum, Uni- ersidad Nacional de Colombia, Santa Fe« de versity of Kansas, Lawrence, Kansas, C. D. Bogota«, Colombia, G. Nates-Parra; USNM, Michener and R. W. Brooks; STRI, Smith- United States National Museum, Smithsoni- sonian Tropical Research Institute, Balboa, an Institution, Washington, D.C., R. J. Panama«, D. W. Roubik; UADE, University McGinley and M. Mello; YUDB, Depart- of Arkansas, Fayetteville, Arkansas, S. A. ment of Biology, York University, North Cameron and J. B. Whitfield; UCD, Bohart York, Canada, L. Packer; ZMHB, Museum Museum of Entomology, Davis, California, fu¬r Naturkunde, Humboldt-Universita¬t, Ber- L. S. Kimsey and L. A. Baptiste; UFVB, lin, Germany, F. Koch and A. Kleine-Mo¬ll- Museu Entomologico de Universidade Fed- hoff.

GENERAL MORPHOLOGY The general morphology of the augochlo- area (fig. 17), but the base of the distal pro- rine Pseudaugochlora graminea (Fabricius) cess can be expanded (e.g., Megalopta and was examined in detail by Eickwort (1969a: Xenochlora) such that it is as broad as the as a species of Pseudaugochloropsis), and basal area (broadly triangular) (fig. 18). In such a study need not be repeated here. In- the genera Augochloropsis and Temnosoma stead, only those characters employed in this the distal process is broad on both ends, be- classification are briefly examined. The most coming almost quadrate (fig. 20). The distal common state of each character is presented process bears on its dorsal surface a longi- along with a discussion of variations found tudinal carina or ridge (the distal keel; figs. in the tribe. Morphological terminology gen- 17Ð19). In two groups the distal keel be- erally follows that of Michener (1944, 1965) comes expanded at its base such that it forms and Eickwort (1969a). a small plateau (fig. 19). The labral teeth along the lateral margins of the distal process FEMALES can be completely absent (fig. 18) or devel- The mandible is often equipped with a sub- oped into strong projections such that the lat- apical tooth that is little differentiated from eral margins appear nearly pectinate (fig. 17). the apex and is positioned slightly basad the The apex of the clypeus in Megommation s.s. apex on the upper margin of the mandible and Megaloptina is deeply concave (figs. 5, (fig. 16). This tooth can become strongly pro- 12) rather than being approximately straight duced from the apex with a deep concavity between two weakly developed lateral tuber- differentiating it and sometimes the tooth cles (figs. 1Ð4, 6). In the genus Rhinocory- reaches apicad as far as the mandibular apex. nura the clypeus frequently bears armature In these cases, the mandible is frequently has such as tubercles. smaller supplementary tubercles or teeth on The prementum is typically much less than the inner surface, basal to the subapical tooth seven times as long as wide (fig. 14). How- (fig. 16). In a few genera (e.g., Temnosoma, ever, in some genera (e.g., Megaloptidia and Cleptommation), the mandible lacks subapical relatives) it is greatly narrowed and elongate, or supplementary teeth (fig. 15). being seven or more times longer than wide The labrum consists of a transverse basal (fig. 13). The base of the galea frequently area that may bear various blunt tubercles extends to the stipital base or nearly so (fig. ranging in shape from orbicular to strongly 22), but in a few genera (treated below as the bilobed. Extending apicad and set below this subtribe Corynurina) it extends only half of basal region is a triangular distal process, po- this distance (fig. 21). In most augochlorines sitioned such that the pointed apex of the tri- the maxillary palpi are much shorter than the angle forms the most distal point of the la- prementum, or at most as long as the pre- brum (fig. 17). The base of this process, mentum. However, in Ariphanarthra they are where it joins with the basal area, is often greatly elongate and flattened so that the pal- slightly narrower than the width of the basal pi reach to the metasoma in repose. The ga- 2000 ENGEL: BEE TRIBE AUGOCHLORINI 11

Figs. 1Ð6. Heads of Augochlorini, frontal aspect. 1. Thectochlora alaris (Vachal), female. 2. Rhin- ocorynura 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. 12 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

Figs. 7Ð12. 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). 2000 ENGEL: BEE TRIBE AUGOCHLORINI 13

Figs. 13Ð16. Female mouthparts. 13. Labiomaxillary complex and hypostomal fossa of Megom- mation (Megaloptina) ogilviei (Cockerell), ventral aspect. 14. Labiomaxillary complex and hypostomal fossa of Corynura (Corynura) chilensis (Spinola), ventral aspect. 15. Clypeus and mandibles of M. (Cleptommation) minutum (Friese). 16. Mandible of Megalopta (Megalopta) genalis Meade-Waldo. lea bears a broad, rounded lobe at its apex border of the head it gently turns outwards, (fig. 24) and the inner surface typically lacks running behind the articulation of the man- modifications (fig. 23); however, in a few dible (fig. 14), or in some augochlorines the genera (e.g., Pseudaugochlora and relatives) point at which this carina turns outward it is the galea is sharply pointed at its apex (fig. developed into a strong point or angle instead 23). In the corynurine genera the inner sur- of a broad curve. The hypostomal ridge is face bears a strong galeal comb (figs. 24, 25). lamellate in Megommation s.s. and projects A strong ridge separates the hypostomal behind the posterior border of the head. fossa from the postgenal area. This ridge is The area of integument between the lower usually developed into the hypostomal carina border of the compound eye and the mandib- which runs from the posterior border of the ular base is the malar space. In most groups head anteriorly (figs. 13, 14). At the anterior this space is extremely short with the com- 14 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

Figs. 17Ð20. Labra of female Augochlorini. 17. Augochlora (Augochlora) pura (Say). 18. Rhino- corynura briseis (Smith). 19. Megalopta (Megalopta) genalis Meade-Waldo. 20. Temnosoma metallicum Smith. pound eye seemingly in contact with the towards the compound eyes is variously con- mandibular base (i.e., the length of the malar structed. In a few genera (e.g., Augochlora) space is much shorter than the basal mandib- this angle is strongly acute and projecting ular width: ﬁgs. 1Ð3, 5, 6, 8). In certain au- into the clypeal base as a small lobe. In gochlorine groups this space becomes greatly Chlerogelloides this is the most well devel- elongate with a separation between the man- oped with the lobe being extremely narrow dibular base and the compound eye at least and nearly reaching to the apical margin of three quarters of the basal mandibular width, the clypeus. In scattered genera the angle is but frequently many times longer (ﬁg. 4). approximately orthogonal (i.e., almost right The epistomal sulcus is subdivided into a angular), whereas in still others (e.g., Cory- number of individual sulci. The angle created nura) it is broadly obtuse (in some cases al- between the dorsal clypeo-genal sulcus and most not forming an angle, the two sulci be- the lateral clypeo-genal sulcus which opens ing almost linear with one other). 2000 ENGEL: BEE TRIBE AUGOCHLORINI 15

Figs. 21Ð25. 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. 16 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

Figs. 26Ð29. Head modiﬁcations. 26. Preoccipital area of Corynura (Corynura) chilensis (Spinola). 27. Preoccipital area of Pereirapis semiaurata (Spinola), carina present. 28. Compound eye of C.(Cal- listochlora) chloris (Spinola), elongate setae. 29. Compound eye of Augochlora (Augochlora) pura (Say), minute setae.

The antennal flagellum of bees is com- compound eyes support setae which are posed of 10 units (flagellomeres) in females about as long as the diameter of an omma- and 11 in males. In the genus Chlerogas the tidium (fig. 29). In a few groups (e.g., Cal- number of these segments is reduced by one listochlora or Caenaugochlora s.s.), these se- in each sex. tae are greatly lengthened, many times as The ocelli of most augochlorines are typ- long as an ommatidial diameter, and are fre- ical for bees (figs. 1Ð5, 7Ð10), being sepa- quently visible on pinned specimens without rated from the compound eyes by their own the aid of a microscope (fig. 28). diameter or more. In a few taxa the ocelli are The vertex is gently rounded or slightly greatly enlarged, coming into close proxim- flattened behind the ocelli but in some genera ity with the compound eyes (figs. 6, 11). The it can be fairly long (figs. 2, 7Ð9). In the 2000 ENGEL: BEE TRIBE AUGOCHLORINI 17

Figs. 30Ð33. 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. genera Megalopta s.l. and Xenochlora a deep distinguish close relatives. In these cases the interocellar furrow runs between and just be- mesosoma is strongly and coarsely punctured hind the lateral ocelli (fig. 11). In Pseudau- (fig. 30). The pronotum extends latero-pos- gochlora, a strong transverse ridge is present teriorly to form the pronotal lobe and in so on the vertex behind the ocelli (fig. 10). The doing forms a dorsal surface that creates the preoccipital area is gently rounded in most lateral angle as it bends posteriorly to form augochlorines (fig. 26); however, in a few the lobe. This angle can be strongly produced groups it bears a strong carina or a weak la- so that it projects anteriorly and forms a large mella (fig. 27). dorsal surface. The ridge between this dorsal The sculpturing of the mesosoma is quite surface and the lateral surface of the prono- variable among augochlorines. A few genera tum is the pronotal dorsal ridge (fig. 31), have distinctive sculpturing patterns that help while the ridge running from the point of the 18 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

Figs. 34Ð37. Characteristics of female Augochlorini. 34. Tegula of Augochloropsis (Paraugochlo- ropsis) sumptuosa (Smith), depicting notch on inner posterior margin. 35. Tegula of Rhinocorynura briseis (Smith). 36. Distal hamuli of hind wing of Megalopta (Megalopta) genalis Meade-Waldo. 37. Distal hamuli of Chlerogella sp. angle ventrally along the lateral surface is the forming a slight bottleneck appearance (fig. pronotal lateral ridge (fig. 31). These ridges 33) that can also be seen in profile (cf. figs. can be rounded, carinate, or in more unusual 30, 31). The tegula is ovoid and usually cases, lamellate. The dorsal surface of the slightly narrower on the anterior end than is pronotum is greatly expanded and convex, the broadly rounded posterior portion (fig. forming an inflated surface in Chlerogella 35). In Augochloropsis s.l., however, the in- and Chlerogelloides. The anterior border of ner posterior border of the tegula is indented the mesoscutum is frequently gently convex or deeply notched (fig. 34), so that the pos- and broadly rounded when viewed from terior part of the tegula is narrowed and bent above (fig. 32). In some genera (e.g., Neo- inward. corynura, Rhinocorynura), however, the me- The probasitarsus is frequently equipped dio-anterior margin is produced anteriorly on its outer edge with a distinctive row of 2000 ENGEL: BEE TRIBE AUGOCHLORINI 19

Figs. 38Ð41. 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. stiff setae known as the anterior basitarsal in order to draw future students to this fea- brush. In bees the protibial spur is modified ture as it appears to be a good character for into an antenna cleaner consisting of a back- species and species-groups. The mesotro- bone spine referred to as the malus and a flat, chanter is gently rounded on its upper bor- inner extension called the velum. The free der in all augochlorines except in Thecto- portion of malus (not attaching to the velum) chlora a strong hooked tubercle is present is variously modified and its margin can be on this surface (fig. 42). The metafemur and serrated, minutely ciliate (fig. 51), or densely metatibia bear the scopa, which is formed of pectinate with the teeth progressively becom- dense, plumose setae used in the transport ing shorter towards the apex (fig. 50). Al- of pollen (fig. 41). Three groups lack the though this structure is quite variable among scopa and have scattered simple setae over and within the genera, it is mentioned here these leg surfaces (fig. 40). On the outer 20 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

Figs. 42Ð45. Leg modifications of female Augochlorini. 42. Inner surface of mesotrochanter of Thectochlora alaris (Vachal) showing hook on dorsal surface. 43. Basitibial plate of Augochlora (Au- gochlora) pura (Say). 44. Basitibial plate of T. alaris. 45. Outer surface of metatibial base of Megom- mation (Cleptommation) minutum (Friese), showing absence of basitibial plate. margin of the metatibial base is frequently tatibial spur range from serrated (figs. 48, found a small, raised area, termed the basi- 49) to pectinate, and bear a variable number tibial plate. The plate is glabrous, although of teeth usually between 3 and 10 (figs. 46, a few groups have some short setae located 47); in Ischnomelissa and Ctenaugochlora, on this plate. In most genera the plate is bor- there are more than 10 teeth densely packed dered on all sides except the base by a dis- together. tinct rim or edge (fig. 43). In scattered gen- The marginal cell of the forewing is bor- era, the plate slopes anteriorly and blends dered anteriorly by vein R and posteriorly by into the integument of the tibia without a Rs. The apex of this cell is typically acute, distinct border along this margin (fig. 44), with Rs gently curving anteriorly before while in others the plate is completely ab- meeting R. In some genera, Rs turns more sent (fig. 45). The inner margins of the me- sharply anteriorly and bears a feeble distal 2000 ENGEL: BEE TRIBE AUGOCHLORINI 21

Figs. 46Ð51. Spurs and antenna cleaners of female Augochlorini. 46. Inner hind tibial spur of Chler- ogella 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). 22 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

Figs. 52Ð55. 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. spur which extends towards the wing apex mesosoma). The sculpturing of the basal area (feebly truncate and appendiculate). The dis- is quite variable; in some groups it is either tal hamuli along the anterior margin of the uniformly striate or imbricate. The basal area hind wing are typically arranged irregularly is typically about as long as the scutellum with a distinct spacing pattern produced by (fig. 38), although in some groups it is large separations between subgroups of the strongly declivitous and only as long as the hamuli (fig. 37). In Megalopta and Nocto- metanotum. In three augochlorine genera the raptor, the hamuli are regularly spaced and basal area is elongate and as long as, or fre- numerous (fig. 36). quently longer than, the scutellum and me- The basal area of the propodeum is the tanotum combined (fig. 39). The posterior dorsal-facing surface of the first abdominal surface of the propodeum bears a narrow me- segment (fused into the thorax to form the dian depression termed the propodeal pit 2000 ENGEL: BEE TRIBE AUGOCHLORINI 23

Figs. 56Ð59. Metasomal modiﬁcations of female Augochlorini. 56. Metasoma of Temnosoma metallicum Smith, dorsal aspect. 57. Metasoma of T. metallicum, proﬁle. 58. S1 of Augochlora (Augo- chlora) pura (Say), median apical ridge. 59. Apex of T5 of Megalopta (Megalopta) genalis Meade- Waldo, medioapical cleft of pseudopygidial area.

(seen on the far right side of fig. 38), the (figs. 56, 57). The medio-apical margin of T5 typical state is a simple, narrow pit; however, bears a strong cleft (fig. 59) that is lost in the in some corynurine genera this pit is set into parasitic genus Temnosoma. Some species of a distinct V-shaped depression. the genus Augochlora bear a strong median The anterior surface of the first metasomal spine or ridge on S1 (fig. 58). In all other tergum is typically flattened with scattered se- instances this surface is gently rounded. tae over smooth, imbricate, or lightly punctured integument (fig. 52). In two groups, this MALES surface is strongly concave and bordered by The basal area of the male labrum has a dense setae forming an acarinarium (figs. 53Ð similar construction to that of the female, al- 55). In one genus the apical borders of T1-2 though it is less developed, typically being are strongly depressed and rimmed posteriorly much shorter than that of the female. The 24 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

Figs. 60Ð63. 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. basal area sometimes bears a small notch. tends to the posterior border of the scutellum, The distal process is typically absent, al- while long antennae extend to the propo- though a small distal process is sometimes deum or beyond. The relative lengths of F1 present as a broad triangular extension of the and F2 are often diagnostic. In most instanc- apical margin (fig. 60). es these flagellomeres are of approximately The male antennae are variously construct- equal length (fig. 62), although some genera ed. The length varies dramatically with some have the second many times longer than the genera having short antennae whereby the first (fig. 61), while others have the first flagellum extends only posterad to the pos- many times longer than the second. The api- terior border of the mesoscutum. In moder- cal flagellomere (F11) in Pseudaugochlora is ately developed antennae the flagellum ex- hooked at its apex (fig. 63). 2000 ENGEL: BEE TRIBE AUGOCHLORINI 25

Figs. 64Ð67. Metasomal modiﬁcations of male Augochlorini. 64. Relatively unmodiﬁed sterna (S5Ð S6) of Neocorynura papallactensis Engel. 65. S3ÐS6 of Pseudaugochlora graminea (Fabricius). 66. S3Ð S6 of Megalopta (Megalopta) genalis Meade-Waldo. 67. T7 of Temnosoma metallicum Smith.

Typically, the legs of males are like those of plumose setae surround the propodeal spi- of females, excluding the usual sex differ- racles in Megaloptina and Cleptommation. ences (e.g., lack of pollen-collecting apparati The male metasoma is typically elongated, in males). However, in one species of Chler- the terga being as long as wide or longer, but ogelloides the mesofemur and mesotibia are in various genera, the metasoma is construct- greatly expanded and equipped with short, ed like that of the female, with each tergum blunt processes on their inner surfaces. The wider than long. In other groups the meta- inner hind tibial spur of males bears a finely soma is petiolate, T1 being much longer than serrated margin except in the genus Chlero- wide and greatly narrowed along its basal gas where it is pectinate with long teeth. half. Mesosomal pubescence varies dramatical- The apical margin of T7 is typically un- ly among the genera, but distinctive patches modified, uninterrupted by processes or 26 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250 emarginations. In the genus Temnosoma, ventral surface that meet medially to produce however, the apical margin of this tergum is the gonobasal bridge. This ventral bridge deeply concave and bilobed (fig. 67). The ranges from membranous to sclerotized. Dor- proctiger includes the membranous remnants sally, the gonobase is depressed along its me- of the internalized T9. The apical margin of dio-apical margin, producing two lobes. In the proctiger is sometimes equipped with dis- many genera these lobes are strongly devel- tinct anal filaments and/or an apical fringe of oped with a deep cleft or concavity separat- microtrichiae. The sterna are variously mod- ing the two; however, some augochlorines ified and some of these features are constant have merely a shallow depression, so that the among species of a given genus. The apical lobes are scarcely developed. The ventral margins of the sterna are typically entire, surface of the penis valve is equipped with a lacking processes or emarginations (fig. 64). broad prong (fig. 72). In a few genera this is In some genera, however, median processes replaced by a stiff keel, while in others it is are found on the apical margins of S3ÐS5, lost altogether. In the genus Temnosoma, the while in others the apical margins of S4ÐS6 dorsal surface of the penis valve bears a large genera bear depressions or clefts (fig. 66). apically directed process (fig. 76). The base Megalopta and Noctoraptor also possess of the gonostylus is infrequently equipped deep lateral notches in the apical margin of with a small thumblike projection known as S4 (fig. 66). The fourth sternum can also be the basal process (figs. 74, 75). This process adorned with a dense triangular setal patch is normally covered by short setae (figs. 74, medially (fig. 65). In Caenaugochlora s.s. 75); however, in Callistochlora the setae are this setal patch is positioned on a series of absent (fig. 77). A parapenial lobe often arises from just below the gonostylar processes median tubercles. Other setal patterns exist on the inner surface of the gonostylus (fig. and, while typically constant within a spe- 73). This does not appear to be homologous cies, do not appear to be shared across with the retrorse lobe found among the Ha- groups of higher rank. The seventh and lictini. The retrorse lobe arises from the ven- eighth sterna are fused and hidden within the tral surface of the gonocoxite and somewhat body just ventral to the genital capsule. The below where the parapenial lobe found apical margins of these sterna can be un- among the augochlorines originates. The modified, or equipped with processes (e.g., ventral gonostylar process is a well-devel- figs. 68Ð71). Along the anterior margin of S8 oped lobe of various shapes. In Augochlo- is a median process referred to as the spic- rella and Ceratalictus it is deeply divided ulum. This structure is present among all au- along its apex. The dorsal gonostylar process gochlorine genera and is frequently longer is also variously shaped, but in a few groups than it is wide (figs. 69Ð71), although in it is reduced to a distinctive setose ridge. The some groups it is quite broad (fig. 68). degree to which the dorsal process is scler- The gonobase possesses two arms on its otized is also diagnostic for some genera.

SYSTEMATICS TRIBE AUGOCHLORINI BEEBE World halictines, the augochlorines are most DIAGNOSIS: Augochlorines are most readily similar to (and likely most closely related to) recognized from other Halictinae by their genera of the Caenohalictus and Agaposte- brilliant metallic coloration; however, this is mon generic complexes; these groups also not a universal character, being neither frequently exhibit metallic-green body col- unique to nor fixed across Augochlorini. The oration. medio-apical cleft of the female fifth meta- DESCRIPTION: Female. Compound eyes somal tergum and the absence of a pygidial frequently with emargination just above level plate on the apical margin of the male sev- of antennae (figs. 1Ð7). Distal wing venation enth metasomal tergum separates Augochlo- strong; basal vein strongly arcuate. Apical rini from other Halictinae. Among New margin of T5 cleft (except in Temnosoma) 2000 ENGEL: BEE TRIBE AUGOCHLORINI 27

Figs. 68Ð71. Hidden and fused male sterna S7ÐS8; spicula are on the left margin and apical processes are shown on the right. 68. Augochloropsis (Paraugochloropsis) sumptuosa (Smith). 69. Tem- nosoma metallicum Smith. 70. Megalopta (Megalopta) genalis Meade-Waldo. 71. Pseudaugochlora graminea (Fabricius).

(ﬁg. 59). Male. Apical margin of T7 without genera (table 2) and until further work on pygidial plate, margin not recurved. Apical augochlorine immatures is undertaken, the margin of S6 cleft (except in Halictillus and validity of this character in supporting the some Augochlora). Spiculum present on an- monophyly of the tribe remains uncertain. terior border of S8 (ﬁgs. 68Ð71). Retrorse Authorship of the tribe has universally lobe of gonocoxite absent. been given to Moure (1943a). In actuality, COMMENTS: A potential larval character Beebe (1925), in a semipopular treatment of for the Augochlorini is the development of the fauna of Guyana, mentions the name the mandibular cusp into an acute, elongate ‘‘Augochloridae’’ to include tropical species projection (McGinley, 1981). Larvae, how- of the genera Augochlora and Halictus. Al- ever, are known for only a few, scattered though Beebe (op. cit.) proposed the group 28 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

Figs. 72Ð77. 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 process. 2000 ENGEL: BEE TRIBE AUGOCHLORINI 29

cavalierly, he is to be considered as the cor- ument frequently brilliant metallic. Male. rect author of the family-group name based Distal process of labrum frequently present. on Augochlora (Engel, 1999b). Apical margins of S4ÐS5 frequently modified. SUBTRIBE AUGOCHLORINA BEEBE Oxystoglossini Schrottky, 1909b: 482. Type ge- Genus Andinaugochlora Eickwort nus: Oxystoglossa Smith, 1853. Suppression in Andinaugochlora Eickwort, 1969a: 407. Type favor of Augochlorini Beebe, 1925, pending species: Andinaugochlora micheneri Eickwort, with I.C.Z.N. (Engel, 1999a, b). 1969a, monobasic and original designation. Augochloridae Beebe, 1925: 102. Type genus: Augochlora Smith, 1853. Corrected authorship DIAGNOSIS: Species of Andinaugochlora (not Moure, 1943a: see Engel, 1999b). superficially resemble larger species of the Augochloropsini Moure, 1943a: 462. Type genus: more widely distributed genus Neocorynura Augochloropsis Cockerell, 1897b. and even more similarly the related genus Megaloptinae Moure, 1943a: 479. Type genus: Neocorynurella. From the former group, An- Megalopta Smith, 1853. dinaugochlora can be separated by the DIAGNOSIS: This is the largest group of Au- broadly rounded mesoscutal anterior border gochlorini and is quite heterogeneous. The and the obsolescent anterior border to the elongate medial process of the premental metabasitibial plate, while from the latter ge- apex that extends anteriorly beyond the api- nus it differs in these same characters and ces of the lateral processes is unique to the also by the nearly orthogonal epistomal sul- Augochlorina and readily separates them cus and carinate preoccipital ridge. from the more primitive subtribe, Corynuri- DESCRIPTION: Female. Mandible with na. strong subapical tooth. Labral distal process DESCRIPTION: As for the tribe with the fol- narrowly triangular; basal elevation orbicu- lowing additions: Female. Angle formed by lar; teeth absent. Prementum not greatly epistomal sulcus variable. Premental apex elongate. Galeal apex rounded; galeal comb with median process elongate, extending be- absent; galeal base extending nearly to base yond apices of lateral processes. Strong ga- of stipes. Hypostomal ridge carinate; anterior leal comb absent (figs. 22, 23); galeal apex angle rounded. Length of malar space less variable, frequently rounded; galeal base ex- than basal mandibular width. Epistomal sul- tends posteriorly to near stipital base (fig. cus nearly orthogonal. Ocelli not greatly en- 22). Marginal cell with apex variable. Integ- larged; ocellar furrow absent. Vertex neither 30 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250 expanded nor ridged behind ocelli. Preoccip- pose extend posteriorly to the metasoma, a ital ridge carinate. Pronotal lateral angle not feature unique among the Augochlorini. produced and obtuse; dorsal ridge carinate; DESCRIPTION: Female. Mandible with lateral ridge rounded. Mesoscutal anterior moderately strong subapical tooth. Labral border rounded; mesoscutal lip rounded. distal process broadly triangular; basal ele- Tegula oval. Anterior basitarsal brush pre- vation orbicular; teeth absent. Prementum sent. Basitibial plate bordered posteriorly, greatly elongate; maxillary palpus elongate, margin obsolete anteriorly. Inner hind tibial extending posterad to metasoma. Galeal apex spur pectinate. Apex of marginal cell trun- pointed, comb absent. Hypostomal ridge car- cate. Distal hamuli with irregular spacing inate; anterior angle rounded. Length of ma- pattern. Basal area of propodeum granular; lar space less than basal mandibular width. propodeal pit narrow. Male. Mandible sim- Epistomal sulcus orthogonal. Ocelli not ple. Labrum with weak distal process; basal greatly enlarged; ocellar furrow absent. Ver- area not notched. Antenna long, extending tex short, barely as long as diameter of me- beyond propodeum; scape short; F2 longer dian ocellus. Preoccipital ridge rounded. than F1. Metasoma elongate. Apical margins Pronotal lateral angle not produced, obtuse; of S4 and S5 straight, unmodified. Apical dorsal ridge carinate; lateral ridge rounded. margin of S6 emarginate. Apical margins of Mesoscutal anterior border rounded; meso- S7 and S8 with medial projections; spiculum scutal lip rounded. Tegula oval. Anterior bas- broad. Proctiger unmodified. Gonobasal itarsal brush present. Basitibial plate with bridge narrow, dorsal lobes strong. Basal well-developed borders. Inner hind tibial process of gonostylus and parapenial lobe spur serrate. Apex of marginal cell acute. absent. Ventral surface of penis valve with Distal hamuli with irregular spacing pattern. prong. Basal area of propodeum with weak basal REVISIONS: The genus has not been re- striae. Male. Mandible simple. Labrum with- vised. There are presently two named spe- out distal process; basal area not notched. cies, Andinaugochlora joannisi (Vachal) and Antennae extending posterad to scutellum; A. micheneri Eickwort, along with at least F1 longer than F2. Inner hind tibial spur ser- four undescribed species (personal obs.). rate. Metasoma oval. S4 unmodified. Apical DISTRIBUTION: The genus is found in Co- margin of S5 weakly emarginate. Apical lombia, Ecuador, and Peru. Based on the margin of S6 emarginate. Apical margins of available collection information, species S7 and S8 unmodified; spiculum narrow. seem to be restricted to montane regions. An Proctiger with anal filaments. Gonobasal undescribed species has recently been iden- bridge narrow; dorsal lobes strong. Parapen- tified from high altitude localities in Costa ial lobe of gonostylus present; basal process Rica (personal obs.). This species will be de- absent; dorsal process large and membra- scribed at a later date by the author. nous. Ventral surface of penis valve with prong. Genus Ariphanarthra Moure REVISIONS: At present Ariphanarthra contains only the type species. Moure (1951) Ariphanarthra Moure, 1951: 137. Type species: presents a thorough description of the spe- Ariphanarthra palpalis Moure, 1951, monobasic and original designation. cies. BIOLOGY: Moure (1951) suggested that Ar- DIAGNOSIS: This monotypic genus resem- iphanarthra may be crepuscular even though bles the nocturnal genus Megaloptidia in its specimens have been captured only during head structure, both having the face slightly the day and nothing about its morphology concave around the antennal sockets and the suggests this habit. Nothing is presently greatly narrowed and elongate prementum. known of Ariphanarthra biology. Ariphanarthra, however, is not nocturnal, DISTRIBUTION: Ariphanarthra has an exten- with dark integumental pigmentation and sive range along the western half of South normal-sized ocelli. The genus is remarkable America. Individuals of A. palpalis Moure for the elongate maxillary palpi that in re- are recorded from northern Argentina, south- 2000 ENGEL: BEE TRIBE AUGOCHLORINI 31 ern Brazil, Paraguay, Peru, as well as south- Michener (1954b) gave a key to the Pana- ern Colombia (Engel, 1996c). manian species and Cockerell (1897b) provided a key to the Mexican species; however, Genus Augochlora Smith this latter key is somewhat out of date and should be used with caution. Neither of these DIAGNOSIS: This large, diverse group is keys utilize the currently recognized subge- second only to the genus Augochloropsis in neric groupings and species identities should, number of recorded species. Augochlora is therefore, be cross-referenced with Moure most similar to the genera Augochlorella, and Hurd (1987) for subgeneric associations Ceratalictus, and Pereirapis. It can be distin- and appropriate synonymies. guished from them all, however, by the acute epistomal sulcus that forms a lobe protruding Subgenus Augochlora Smith s.s. into the basal margin of the clypeus. Figures 17, 41, 43, 48, 58 DESCRIPTION: Female. Lateral margins of labral distal process with strong teeth. Pre- Augochlora Smith, 1853: 73. Type species: Ha- mentum not greatly elongate. Galeal apex lictus purus Say, 1837, designated by Cockerell rounded; galeal comb absent. Hypostomal (1923). Oxystoglossa Smith, 1853: 83. Type species: Oxy- ridge carinate; anterior angle pointed or pro- stoglossa decorata Smith, 1853, monobasic. duced into a short tubercle. Length of malar Angochlora Schrottky, 1901: 212. Lapsus calami. space less than basal mandibular width. Ep- Odontochlora Schrottky, 1909a: 141. Type spe- istomal sulcus acute, protruding into clypeus. cies: Augochlora mulleri Cockerell, 1900 [ϭ Ocelli not greatly enlarged; ocellar furrow Augochlora muelleri Schrottky, 1909a, nomen absent. Vertex not ridged behind ocelli. Pre- emendatum (unjustified)], original designation. occipital ridge carinate among living species Oxystoglosss Moure, 1940: 57. Lapsus calami. (rounded in one extinct species: see below Odontochlor Mitchell, 1960: 456. Lapsus calami. under subgenus Electraugochlora). Pronotal Auglochlora Dodson, 1967: 6. Lapsus calami. lateral angle produced, acute to obtuse; dor- Augochlora (Mycterochlora) Eickwort, 1969a: 423. Type species: Halictus repandirostris sal ridge carinate; lateral ridge angled, but Vachal, 1911, original designation. NEW SYN- distinctly not carinate. Mesoscutal anterior ONYMY. border rounded; mesoscutal lip rounded. Tegula oval. Anterior basitarsal brush pre- DIAGNOSIS: The nominate subgenus is sent. Basitibial plate with well-developed most easily separated from the other subgen- borders. Inner hind tibial spur serrate. Apex era by the strongly bidentate mandible and of marginal cell truncate. Distal hamuli with the transverse basal elevation on the labrum. irregular spacing pattern. Basal area of pro- DESCRIPTION: As for the genus with the podeum striate or rugose. S1 frequently bear- following additions: Female. Mandible ing anterior median ridge or spine. Male. strongly bidentate. Labral distal process usu- Mandible simple. Labrum without distal pro- ally narrowly triangular (a few species have cess; basal area not notched. Antennae of broadly triangular processes); basal elevation moderate length, extending posterad to scu- transverse, protuberant. Angle of epistomal tellum; F2 length approximately equal to F1. sulcus strongly protruding into clypeus. Pre- Inner hind tibial spur serrate. Metasoma oval. occipital ridge carinate. Male. Ridge border- Apical margins of S4 and S5 unmodified. ing ventral gonostylar process with short se- Apical margin of S6 usually emarginate (a tae, not surpassing gonostylar apex. few species are unmodified). Apical margins REVISIONS: There has been no revision of of S7 and S8 with median projection; spic- Augochlora s.s. Moure and Hurd (1987) list- ulum narrow. Proctiger with anal filaments. ed 85 described species, although one, Au- Gonobasal bridge narrow; dorsal lobes gochlora essequibensis Cockerell, is newly strong. Parapenial lobe absent; basal process synonymized with A. nigrocyanea Cockerell present; dorsal process reduced to strong below (appendix 1). Three previously un- ridge with setae. Ventral surface of penis placed species have been recently transferred valve with prong. to this subgenus (Moure, 1999). REVISIONS: Refer to subgeneric treatments. BIOLOGY: Species of Augochlora are soli- 32 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250 tary and form nests in rotting wood (Eick- Cockerell, 1914, monobasic and original des- wort and Eickwort, 1973b; Stockhammer, ignation. NEW SYNONYMY. 1966). Immature stages have been described DIAGNOSIS: Oxystoglossella differs from by Michener (1953, 1954a) and Eickwort Augochlora s.s. by the less bidentate man- and Eickwort (1973b) (table 2). dibular apex and orbicular basal elevation on DISTRIBUTION: This subgenus has the great- the labrum. From Electraugochlora it differs est distribution of any augochlorine group. in the presence of a preoccipital carina and Species range from northern Argentina to a strong epistomal lobe. southern Canada and into the West Indies. DESCRIPTION: As for the genus with the following additions: Female. Mandible with Electraugochlora, new subgenus moderate to strong subapical tooth. Labral Figures 79Ð81 distal process narrowly triangular; basal ele- TYPE SPECIES: Augochlora (Electraugo- vation orbicular, protuberant. Angle of epi- chlora) leptoloba, new species (described in stomal sulcus strongly protruding into clyp- appendix 1). eus. Preoccipital ridge carinate. Male. Ridge DIAGNOSIS: This monotypic subgenus is bordering ventral gonostylar process with unique among Augochlora species in the ab- long setae, greatly surpassing gonostylar sence of a preoccipital carina, instead having apex. an entirely rounded preocciptal area (a de- REVISIONS: The species of Oxystoglossella rived feature among the enitre Augochlora- have not been revised. Moure and Hurd group of genera). The acute epistomal sulcus (1987) listed 28 described species, although places it among Augochlora, but the lobe is two of these have since been synonymized very small, in this way differing from living (Engel, 1996c) and a new species is de- species of the genus. The orbicular basal el- scribed in appendix 1. evation on the labrum is similar to Oxysto- BIOLOGY: Oxystoglossella species are glossella, but in this subgenus the elevation primitively eusocial and excavate nests in the is strongly protuberant. Further details of this soil (Eickwort and Eickwort, 1972). The ma- groups morphology are presented below as ture larva and pupa have been described for well as in appendix 1, where the type species A. cordiaefloris Cockerell and A. nominata is described. Michener (Eickwort and Eickwort, 1972) (ta- DESCRIPTION: As for the genus with the ble 2). following additions: Female. Mandible with DISTRIBUTION: Oxystoglossella has a wide moderate subapical tooth. Labral basal ele- distribution; species occur from Argentina to vation orbicular, low and not protuberant the southwestern United States and east into (figs. 80, 81). Angle of epistomal sulcus only the West Indies. weakly protruding into clypeus (figs. 80, 81). Preoccipital ridge rounded. S1 without me- Genus Augochlorella Sandhouse dian ridge or spine. Male. Unknown. Augochlorella Sandhouse, 1937: 66. Type spe- ETYMOLOGY: The new generic name is a cies: Augochlora gratiosa Smith, 1853, original combination of electrum (L. amber) and the designation. genus Augochlora, type genus of the tribe. Oxystoglossidia Moure, 1943a: 473. Type species: REVISIONS: There is only the one included Oxystoglossidia uraniella Moure, 1943a [ϭ species (described in appendix 1). Oxystoglossa ephyra Schrottky, 1911], original DISTRIBUTION: Presently known only from designation. Miocene Dominican amber. DIAGNOSIS: This genus can be separated from other members of the Augochlora- Subgenus Oxystoglossella Eickwort group (Augochlora, Ceratalictus, and Perei- Augochlora (Oxystoglossella) Eickwort, 1969a: rapis) by the combination of an orthogonal 422. Type species: Augochlora cordiaefloris epistomal sulcus, the acute marginal cell Cockerell, 1907, original designation. apex, and the absence of a large basal lobe Augochlora (Aethechlora) Moure and Hurd, 1987: on the inner metatibial spur. 275. Type species: Augochlora matucanensis DESCRIPTION: Female. Mandible with 2000 ENGEL: BEE TRIBE AUGOCHLORINI 33 weak subapical tooth. Labral distal process concerning nestmate relatedness (Mueller, narrowly triangular; basal elevation orbicular 1991, 1996; Mueller et al., 1994) and social to transverse; teeth weak. Prementum not development (Ordway, 1965; Packer, 1990). greatly elongate. Galeal apex rounded; galeal The mature larva and pupa have been de- comb absent. Hypostomal ridge carinate; an- scribed for A. edentata Michener by Eick- terior angle rounded. Length of malar space wort and Eickwort (1973a) (table 2). less than basal mandibular width. Epistomal DISTRIBUTION: Species of Augochlorella sulcus orthogonal. Ocelli not greatly en- are distributed from northern Argentina to larged; ocellar furrow absent. Vertex neither southern Canada. No species occur in the expanded nor ridged behind ocelli. Preoccip- West Indies. ital ridge carinate. Pronotal lateral angle not produced, orthogonal to obtuse; dorsal ridge Genus Augochlorodes Moure carinate; lateral ridge angled, but distinctly not carinate. Mesoscutal anterior border Augochlorodes Moure, 1958a: 53. Type species: rounded; mesoscutal lip rounded. Tegula Augochlorodes turrifaciens Moure, 1958a, oval. Anterior basitarsal brush present. Ba- monobasic and original designation. Augochlorocles Sakagami, 1979: 83. Lapsus cal- sitibial plate with well-developed borders. ami. Inner hind tibial spur serrate. Apex of marginal cell apex acute. Distal hamuli with ir- DIAGNOSIS: Augochlorodes is similar to the regular spacing pattern. Basal area of pro- genera of the Augochlora-group (Augochlo- podeum striate, rugose, or granular. Male. ra, Augochlorella, Ceratalictus, and Perei- Mandible simple. Labrum with distal pro- rapis) but differs by the pectinate inner me- cess; basal area not notched. Antenna ex- tatibial spur and the rounded preoccipital tending back to scutellum; F2 shorter than ridge. F1. Inner hind tibial spur serrate. Metasoma DESCRIPTION: Female. Mandible with oval. Apical margins of S4 and S5 unmodi- moderately developed subapical tooth. La- fied. Apical margin of S6 emarginate. Apical bral distal process narrowly triangular; basal margins of S7 and S8 with median processes; elevation orbicular; teeth weak. Prementum spiculum narrow. Proctiger with anal fila- not greatly elongate. Galeal apex rounded; ments. Gonobasal bridge narrow; dorsal galeal comb absent. Hypostomal ridge cari- lobes variously produced. Parapenial lobe nate; anterior angle rounded. Length of malar and basal process of gonostylus absent; ven- space less than basal mandibular width. Ep- tral process divided; dorsal process variously istomal sulcus obtuse. Ocelli not greatly en- constructed, a large partially membranous or larged; ocellar furrow absent. Vertex not ex- sclerotized flange. Ventral surface of penis panded or ridged behind ocelli. Preoccipital valve with prong. ridge rounded. Pronotal lateral angle not pro- REVISIONS: The species of Augochlorella duced, obtuse; dorsal ridge carinate; lateral occurring in the United States and Mexico ridge rounded. Mesoscutal anterior border have been revised by Ordway (1966b). A rounded; lip rounded. Tegula oval. Anterior new revision of the genus, including the basitarsal brush present. Basitibial plate with South American species, is currently under well-developed borders. Inner hind tibial preparation (B. Coelho, in prep.). Moure and spur pectinate. Apex of marginal cell acute. Hurd (1987) listed sixteen extant species; Distal hamuli with irregular spacing pattern. however, their Augochlorella cladopyga Basal area of propodeum weakly striate ba- (Cockerell) is a junior synonym of Perei- sally; propodeal pit narrow. Male. Mandible rapis semiaurata (Spinola) (see appendix 1). simple. Labrum with distal process; basal BIOLOGY: Species of Augochlorella are area not notched. Antenna extending back to primitively eusocial and nest in the soil scutellum; F2 length approximately equal to (Eickwort and Eickwort, 1973a; Knerer, F1. Inner hind tibial spur serrate. Metasoma 1968; Knerer and Atwood, 1962, 1966; Ord- elongate. Apical margin of S4 produced lat- way, 1961, 1964, 1966a; Sakagami and Mou- erally, with long setae. Apical margin of S5 re, 1967). One species, A. striata (Provanch- unmodified, with dense medial patch of se- er), has been the focus of a number of studies tae. Apical margin of S6 weakly emarginate. 34 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

Apical margin of S7 with median process. broad. Proctiger unmodified. Gonobasal Apical margin of S8 unmodified; spiculum bridge narrow, or completely membranous; broad. Proctiger with anal filaments. Gono- dorsal lobes strong. Basal process of gonobasal bridge narrow; dorsal lobes weak. Par- stylus and parapenial lobe absent; ventral apenial lobe and basal process of gonostylus process with long, thin process. Ventral sur- absent; dorsal process large and weakly face of penis valve with keel. Volsella with sclerotized. Ventral surface of penis valve inner apical angle produced into thin projec- with prong. tion. REVISIONS: At present there is only the one REVISIONS: There is no revision of any Au- included species. gochloropsis subgenus, and many species are BIOLOGY: The biology of the type species not placed to subgenus. Excluding the four was studied by Michener and Seabra (1959). species placed in the subgenus Glyptochlora This species nests in the soil and is appar- (although one placed there only provision- ently semisocial. ally), there are 134 species listed by Moure DISTRIBUTION: Augochlorodes is currently and Hurd (1987) as belonging to Augochlo- known only from southern Brazil. ropsis and Paraugochloropsis. Since the time of their catalog two species of Parau- Genus Augochloropsis Cockerell gochloropsis have been synonymized (Engel, 1996c) and a third has been newly trans- DIAGNOSIS: This is the most speciose genus ferred to the genus bringing the total specific of the tribe. Augochloropsis s.l. can be quick- diversity of the genus to 133. ly identified by the presence of a notch on BIOLOGY: The nesting biology of species the inner, posterior margin of the tegula as in this genus have been studied by several well as the pectinate inner metatibial spur authors (Gimenes et al., 1991; Michener and and the lamellate pronotal dorsal ridge. Lange, 1959; Michener and Seabra, 1959; DESCRIPTION: Female. Mandible with Sakagami and Moure, 1967; Smith, 1901). moderate to strong subapical tooth. Labral All are soil nesters and are presumably com- distal process quadrate; basal elevation bi- munal. The peculiar quadrate pollen masses lobed; teeth absent. Prementum not greatly serve as an ethological synapomorphy for the elongate. Galeal apex rounded; galeal comb genus. absent; galeal base extending to base of sti- DISTRIBUTION: The genus ranges from Ar- pes. Hypostomal ridge carinate; anterior an- gentina to southern Canada, although the gle rounded. Length of malar space less than three species occurring in the United States basal mandibular width. Epistomal sulcus and Canada belong to the subgenus Parau- obtuse. Ocelli not greatly enlarged; ocellar gochloropsis. Species are known from the is- furrow absent. Pronotal lateral angle pro- land of Trinidad, but do not occur in the West duced, orthogonal to obtuse; dorsal ridge la- Indies. mellate; lateral ridge rounded to angled. Teg- ula with inner posterior border notched. An- Subgenus Augochloropsis Cockerell s.s. terior basitarsal brush present. Basitibial plate bordered on all sides, extremely short. Augochlora (Augochloropsis) Cockerell, 1897b: 4. Type species: Augochlora (Augochloropsis) Inner hind tibial spur pectinate. Apex of mar- subignita Cockerell, 1897a [ϭ Augochlora ig- ginal cell truncate. Distal hamuli with irreg- nita Smith, 1861], original designation. ular spacing pattern. Male. Mandible simple. Angochlora (Angochloropsis) Schrottky, 1901: Labrum without distal process; basal area not 213. Lapsus calami. notched. Antennae extending back to scutel- Augochlora (Auhochloropsis) Moure, 1940: 45. lum; F2 approximately equal in length to F1. Lapsus calami. Inner hind tibial spur serrate. Metasoma oval. Augochlora (Autochloropsis) Moure, 1940: pl. 2. Apical margin of S4 produced laterally, with Lapsus calami. setae at apices of projections, also produced Augoschloropsis Moure, 1943b: 197. Lapsus cal- medially. Apical margin of S6 emarginate. ami. Auochloropsis Roubik, 1989: 392. Lapsus calami. Apical margin of S7 and S8 with median process, that of S8 usually bilobed; spiculum DIAGNOSIS: Among the subgenera of Au- 2000 ENGEL: BEE TRIBE AUGOCHLORINI 35 gochloropsis, the nominate subgenus is most striking differences from Paraugochloropsis similar to Paraugochloropsis from which it and Augochloropsis s.s. can be separated by the presence of striae on DISTRIBUTION: All three species are known the basal area of the propodeum. only from northwestern Brazil (Amazonas, DESCRIPTION: As for the genus with the Saõ Paulo de Olivença), while the uncertain following additions: Female. Vertex not species, A. cyclis, is from Peru. Unidentified greatly shortened, longer than diameter of material of Glyptochlora has recently been median ocellus. Preoccipital ridge carinate. collected in French Guiana (R. W. Brooks, Mesoscutal anterior border rounded; meso- personal commun.). scutal lip rounded. Basal area of propodeum striate. Male. Apical margin of S5 emargin- KEY TO THE SPECIES OF AUGOCHLOROPSIS ate. (GLYPTOCHLORA) REVISIONS: See the account of the genus. 1. Metasomal T1ÐT2 lacking apical fringe of se- BIOLOGY: See the account of the genus. tae ...... 2 DISTRIBUTION: See the account of the ge- — Metasomal T1ÐT2 with conspicuous apical nus. fringe of setae ...... atropos (Smith) 2. Basal area of propodeum with strong carina Subgenus Glyptochlora Moure, revised on dorsal ridge and on margins with pro- status podeal lateral surfaces, strong medial carina running between basal and apical margins Augochloropsis (Glyptochlora) Moure, 1958b: ...... ornata (Smith) 188. Type species: Megalopta ornata Smith, — Basal area of propodeum not encircled by a 1879, original designation. strong carina ...... refulgens (Smith)

DIAGNOSIS: The species of Glyptochlora Subgenus Paraugochloropsis Schrottky are the most distinctive among all Augochlo- Figures 34, 68 ropsis species. The coarse punctation of the body resembles to some degree that of the Augochloropsis (Paraugochloropsis) Schrottky, cleptoparasitic genus Temnosoma. The ex- 1906: 312. Type species: Augochloropsis (Par- augochloropsis) lycorias Schrottky, 1906 [ϭ tremely short vertex, sharply carinate preoc- Augochlora epipyrgitis Holmberg, 1903], cipital ridge, and the strongly narrowed and monobasic. lamellate anterior border of the mesoscutum Augochloropsis (Pseudaugochloropsis) Schrottky, separate Glyptochlora from all other Augo- 1906: 313. Type species: Augochloropsis chloropsis. (Pseudaugochloropsis) sthena Schrottky, 1906, DESCRIPTION: As for the genus with the designation of Sandhouse (1943). The desig- following additions: Female. Head and me- nation of Halictus nigromarginatus Spinola, sosoma coarsely punctured. Vertex short, 1841 [ϭ Megilla graminea Fabricius, 1804], as barely one ocellar diameter in length. Pre- the type by Schrottky (1909b) and supported by occipital ridge strongly carinate and coming Moure (1944) is erroneous as it was not an originally included species (Sandhouse, 1943; to a sharp angle. Mesoscutal anterior border Michener, 1954b, 1994). narrowed; mesoscutal lip lamellate. Basal Augochlora (Tetrachlora) Schrottky, 1909b: 481. area of propodeum strongly striate. Male. Type species: Halictus multiplex Vachal, 1903, Unknown. monobasic. REVISIONS: At present Glyptochlora con- Paraugochlora Schrottky, 1910: 540. Type spe- tains three species, A. atropos (Smith), A. or- cies: Augochlora spinolae Cockerell, 1900, nata (Smith), and A. refulgens (Smith), original designation. which are identiﬁed by the key provided be- Rivalisia Strand, 1921: 270. Type species: Rival- ϭ low. A fourth species, A. cyclis (Vachal), has isia metallica Strand, 1921 [ Augochloropsis been referred to this subgenus by Moure and (Paraugochloropsis) aenigma Engel, 1996c], monobasic. Hurd (1987). This last species is included Augochlora (Glyptobasis) Moure, 1940: 48. Type here only provisionally until further work species: Augochlora (Glyptobasis) chloe¬ra can be done on the subgenus. Glyptochlora Moure, 1940, original designation. Nomen might eventually be recognized at the generic praeoccupatum [nec Glyptobasis M’Lachlan, level, should the male exhibit additional 1871 (Neuroptera: Ascalaphidae) and others]. 36 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

Glyptobasia Moure, 1941: 98. Nomen novum pro angular setal patch on S4. Apical margins of Glyptobasis Moure, 1940. Type species: auto- S5 and S6 emarginate. Apical margin of S7 basic with Glyptobasis Moure, 1940. with median projection. Spiculum narrow. Tetrachlor Mitchell, 1960: 463. Lapsus calami. Proctiger unmodified. Gonobasal bridge nar- DIAGNOSIS: See Diagnosis for Augochlo- row; dorsal lobes weak. Basal process of ropsis s.s. gonostylus and parapenial lobe absent; dorsal DESCRIPTION: As for the genus with the process partly membranous. Ventral surface following additions: Female. Vertex not of penis valve with prong. greatly shortened, longer than diameter of median ocellus. Preoccipital ridge carinate. Subgenus Caenaugochlora Michener s.s. Mesoscutal anterior border rounded; meso- Figure 60 scutal lip rounded. Basal area of propodeum Caenaugochlora Michener, 1954b: 76. Type spe- rugulose or granular. Male. Apical margin of cies: Caenaugochlora macswaini Michener, S5 unmodified. 1954b, original designation. REVISIONS: See the account of the genus. Coenaugochlora Michener, 1954b: 85. Lapsus BIOLOGY: See the account of the genus. calami. DISTRIBUTION: See the account of the ge- DIAGNOSIS: Caenaugochlora s.s. is distin- nus. guished from Ctenaugochlora by the normal pectination of the inner metatibial spur, fre- Genus Caenaugochlora Michener quent presence of long compound eye hairs, strong anterior border of the basitibial plate, DIAGNOSIS: Species of Caenaugochlora are similar in general appearance to those of and propodeal striae not reaching to the api- the genus Pseudaugochlora; the later genus cal margin. has, however, a strong ridge on the vertex DESCRIPTION: As for the genus with the and a pointed galeal apex. From the related following additions: Female. Compound genus Augochloropsis, Caenaugochlora dif- eyes usually with long hairs. Pronotal lateral fers in the absence of a notch on the tegula, angle usually produced, angle slightly acute the nonlamellate pronotal dorsal ridge, and to obtuse. Basitibial plate with well-devel- the orthogonal epistomal sulcus. oped borders. Inner hind tibial spur pectinate. Basal area of propodeum with basal striae, DESCRIPTION: Female. Mandible with subapical tooth variously defined. Labral distal not reaching apical margin. Male. Apical process narrowly triangular; basal elevation margin of S4 concave, with dense setal orbicular; teeth absent. Prementum not great- patches raised on tubercles toward apex. Api- ly elongate. Galeal apex rounded; galeal cal margin of S8 with median projection, comb absent; galeal base extending to stipital sometimes bilobed at apex. Volsella indented base. Hypostomal ridge carinate; anterior an- on inner margin. gle rounded. Length of malar space less than REVISIONS: There has been no revision of basal mandibular width. Epistomal sulcus or- Caenaugochlora species. At the present time, thogonal. Ocelli not greatly enlarged; ocellar there are 15 described species in the subge- furrow absent. Vertex not expanded or ridged nus, but many more await study. behind ocelli. Preoccipital ridge angled or BIOLOGY: The soil-nesting biology of Cae- carinate. Pronotal dorsal ridge carinate; lat- naugochlora costaricensis (Friese) was stud- eral ridge rounded to carinate. Mesoscutal ied by Michener and Kerfoot (1967: as a spe- anterior border rounded; mesoscutal lip cies of Pseudaugochloropsis), this species is rounded. Tegula oval. Anterior basitarsal possibly semisocial. brush present. Apex of marginal cell trun- DISTRIBUTION: Caenaugochlora ranges cate. Distal hamuli with irregular spacing from Colombia to Mexico. No species are pattern. Propodeal pit narrow. Male. Man- known to occur in the West Indies. dible simple. Labrum with distal process; basal area notched. Antennae extending back Subgenus Ctenaugochlora Eickwort to scutellum; F2 about as long as F1. Inner Caenaugochlora (Ctenaugochlora) Eickwort, hind tibial spur serrate. Metasoma oval. Tri- 1969a: 435. Type species: Neocorynura perpec- 2000 ENGEL: BEE TRIBE AUGOCHLORINI 37

tinata Michener, 1954b, monobasic and original ridge angled. Mesoscutal anterior border designation. rounded; mesoscutal lip rounded. Tegula DIAGNOSIS: Refer to Diagnosis for Cae- oval. Anterior basitarsal brush present. Ba- naugochlora s.s. sitibial plate bordered posteriorly, margin ob- DESCRIPTION: As for the genus with the solete anteriorly. Inner hind tibial spur ser- following additions: Female. Compound rate. Apex of marginal cell acute. Distal eyes with minute hairs. Pronotal lateral angle hamuli with irregular spacing pattern. Basal not produced, obtuse. Basitibial plate bor- area of propodeum striate; propodeal pit nar- dered posteriorly, margin obsolete anteriorly. row. Male. Mandible simple. Labrum with Inner hind tibial spur densely pectinate, with distal process; basal area not notched. Anten- more than 10 long teeth. Basal area of pro- nae extending back to scutellum; F1 about as podeum with strong striae radiating from long as F2. Inner hind tibial spur serrate. Me- basal margin to apex. Male. Apical margin tasoma oval. Apical margins of S4 and S5 of S4 concave, dense triangular setal patch, unmodified. Apical margin of S6 emarginate. patch not raised on tubercles. Apical margin Apical margin of S7 unmodified. Apical of S8 unmodified. Inner margin of volsella margin of S8 with median process; spiculum rounded. narrow. Proctiger unmodified. Gonobasal REVISIONS: There has been no revision of bridge narrow; dorsal lobes strong. Parapen- Ctenaugochlora, although Engel (1995a) ial lobe present; basal process of gonostylus provided a key to the four described species. present; ventral process divided. Ventral sur- DISTRIBUTION: Ctenaugochlora is currently face of penis valve with prong. known from Costa Rica and Panama al- REVISIONS: The genus has not been re- though undescribed species occur in Trinidad vised. At present there are six recognized and Mexico (personal obs.). species (Moure and Hurd, 1987; Moure, 1999). Moure (1999) recently synonymized Genus Ceratalictus Moure the type species, Ceratalictus theius (Schrott- ky), with C. clonius (Bre`thes). Ceratalictus Moure, 1943a: 463. Type species: DISTRIBUTION: The genus occurs in south- Oxystoglossa theia Schrottky, 1911 [ϭ Augo- ern Brazil, Bolivia, Paraguay, and southeast- chlora clonia Bre`thes, 1909], original designa- ern Peru. tion. Ceratilictus Roubik, 1989: 392. Lapsus calami. Genus Chlerogas Vachal DIAGNOSIS: This genus is most similar to Augochlorella and it may be prudent in the Chlerogas Vachal, 1904: 127. Type species: Ha- lictus chlerogas Vachal, 1904, monobasic and future to consider Ceratalictus a junior syn- absolute tautonymy. onym of the former. Ceratalictus can be separated from Augochlorella by the obtuse ep- DIAGNOSIS: The elongate heads produced istomal sulcus and the obsolescent anterior by a greatly lengthened malar space serves border of the basitibial plate. to separate Chlerogas from other Augochlo- DESCRIPTION: Female. Mandible with rini except the genera Chlerogella and Is- moderately developed subapical tooth. La- chnomelissa. The genera, however, are not bral distal process narrowly triangular; basal related and although Chlerogas resembles elevation orbicular; teeth weak. Prementum the latter two in the structure of the head, this not greatly elongate. Galeal apex rounded; genus is unique for the pectinate inner hind galeal comb absent; galeal base extending to tibial spur of the male, the reduced flagel- base of stipes. Hypostomal ridge carinate; lomere count, the shorter propodeal basal anterior angle rounded. Length of malar area, and the larger body sizes. space less than basal mandibular width. Ep- DESCRIPTION: Female. Mandible with istomal sulcus obtuse. Ocelli not greatly en- moderately defined subapical tooth. Labral larged; ocellar furrow absent. Vertex not ex- distal process narrowly triangular; basal ele- panded or ridged behind ocelli. Preoccipital vation orbicular; teeth absent. Prementum ridge carinate. Pronotal lateral angle pro- not greatly elongate. Galeal apex rounded; duced, acute; dorsal ridge carinate; lateral galeal comb absent; galeal base extending to 38 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

base of stipes. Hypostomal ridge carinate; DIAGNOSIS: The genus Chlerogella is most anterior angle rounded. Malar space much similar and closely related to the genera longer than basal mandibular width. Episto- Chlerogelloides and Ischnomelissa, sharing mal sulcus very slightly acute, only a little with these genera the elongate propodeum. less than 90Њ. Flagellum with only 9 flagel- From the former Chlerogella differs in the lomeres. Ocelli not greatly enlarged; ocellar elongate malar space, the pectinate inner me- furrow absent. Vertex not expanded or ridged tatibial spur, strong basitibial plate, and in the behind ocelli. Preoccipital ridge angled. structure of the epistomal sulcus; while from Pronotal lateral angle not produced, obtuse; the later genus Chlerogella differs in the nor- dorsal ridge angled; lateral ridge rounded. mal pectinations of the inner metatibial spur, Mesoscutal anterior border rounded; meso- the acute epistomal sulcus, and the inflated scutal lip rounded. Tegula oval. Anterior bas- dorsal surface of the pronotum. itarsal brush present. Basitibial plate border DESCRIPTION: Female. Mandible with on all sides. Inner hind tibial spur pectinate. moderately developed subapical tooth. La- Apex of marginal cell acute. Distal hamuli bral distal process narrowly triangular; basal with irregular spacing pattern. Basal area of elevation orbicular to bilobed; teeth absent. propodeum about as long as scutellum, gran- Prementum not greatly elongate. Galeal apex ular or striate; propodeal pit narrow. Male. rounded; galeal comb absent; galeal base ex- Mandible simple. Labrum with short distal tending to base of stipes. Hypostomal ridge process; basal area not notched. Antennae carinate; anterior angle rounded. Length of long, extending back to propodeum; flagel- malar space as long as or frequently longer lum with only 10 flagellomeres. Inner hind than basal mandibular width. Epistomal sul- tibial spur pectinate. Metasoma elongate. cus acute. Ocelli not greatly enlarged; ocellar Apical margins of S4ÐS5 concave. Apical furrow absent. Vertex not expanded or ridged margin of S6 emarginate. Apical margin of behind ocelli. Preoccipital ridge rounded. S7 with median projection. Apical margin of Pronotal dorsal surface frequently inflated, S8 with median projection; spiculum broad. rendering lateral ridge and angle obsolete; Proctiger unmodified. Gonobasal bridge nar- dorsal ridge rounded. Mesoscutal anterior row; dorsal lobes moderately developed. Par- border rounded; mesoscutal lip absent. Teg- apenial lobe and basal process of gonostylus ula oval. Anterior basitarsal brush present. absent; dorsal process a large lightly mela- Basitibial plate bordered on all sides. Inner nized mass. Volsella indented on inner mar- hind tibial spur pectinate. Apex of marginal gin at base of digitus; digitus sometimes cell acute. Distal hamuli with irregular spac- elongated apically. Ventral surface of penis ing pattern. Basal area of propodeum longer valve unmodified. than scutellum, smooth to granular; propo- REVISIONS: The genus has recently been re- deal pit narrow. Male. Mandible simple. La- vised by Brooks and Engel (1999) who recognize nine species. brum without distal process; basal area not notched. Antennae long, extending back to BIOLOGY: Nothing is known of Chlerogas biology, although Brooks and Engel (1999) posterior border of propodeum; F2 longer speculated that the elongate heads are an ad- than F1. Inner hind tibial spur serrate. Me- aptation for tubular flowers. tasoma elongate. Apical margin of S4 emar- DISTRIBUTION: Species of Chlerogas are ginate. Apical margin of S5 unmodified. Api- found throughout northern South America, cal margin of S6 emarginate. Apical margin although the range of any single species is of S7 with median projections. Apical mar- seemingly restricted. Species are known gin of S8 with median projection; spiculum from the mountains of Bolivia, Colombia, narrow. Proctiger unmodified. Gonobasal Ecuador, Peru, and Venezuela. bridge narrow; dorsal lobes weak. Parapenial lobe and basal process of gonostylus absent. Genus Chlerogella Michener Ventral surface of penis valve with prong. Figures 4, 37, 39, 46, 50 REVISIONS: A revision of Chlerogella is Chlerogella Michener, 1954b: 75. Type species: being completed by Brooks and Engel (in Chlerogella elongaticeps Michener, 1954b, prep.) and currently recognizes 17 species monobasic and original designation. (15 new to science), although one, Chlero- 2000 ENGEL: BEE TRIBE AUGOCHLORINI 39 gella bouyssoni (Vachal), is uncertainly basitarsal brush absent. Basitibial plate rep- placed in the absence of the holotype (Engel, resented by slightly elevated glabrous region. personal obs.). Inner hind tibial spur serrate. Apex of mar- BIOLOGY: The biology of Chlerogella re- ginal cell acute. Distal hamuli with irregular mains undiscovered. The elongate heads may spacing pattern. Basal area of propodeum indicate an association with flowers possess- longer than scutellum and metanotum coming deep corollas similar to that hypothesized bined, smooth; propodeal pit narrow. Male. for species of Chlerogas (Brooks and Engel, Mandible simple, with strong double curve 1999). towards apex. Labral basal elevation absent; DISTRIBUTION: Species of Chlerogella are distal process absent. Antenna short, extend- distributed throughout southern Central ing posteriorly to mesoscutum; F1 longer America and northern South America. Their than F2. Inner hind tibial spur serrate. Me- range includes mountainous regions of the tasoma elongate. S3ÐS5 unmodified. Apical following countries: Bolivia, Colombia, Cos- margin of S6 emarginate. Apical margin of ta Rica, Ecuador, Panama, Peru, and Vene- S7 with bilobed projection. Apical margin of zuela. S8 with median projection; spiculum narrow. Proctiger unmodified. Gonobasal bridge nar- Genus Chlerogelloides row; dorsal lobes weak. Gonostylus without Engel, Brooks, and Yanega basal process; parapenial lobe present. Venter of penis valve with prong. Chlerogelloides Engel, Brooks, and Yanega, REVISIONS: The genus presently contains 1997: 3. Type species: Chlerogelloides femo- two species that have been revised by Engel ralis Engel, Brooks, and Yanega, 1997, monobasic and original designation. and Brooks (1999b). BIOLOGY: Nothing is known of Chlerogel- DIAGNOSIS: This genus is remarkably sim- loides biology, although the peculiarly mod- ilar to Chlerogella; both having the peculiar ified midlegs of the type species, C. femo- inflation of the pronotal dorsal surface. ralis, are likely involved in mating. Chlerogelloides, however, has a deeply acute DISTRIBUTION: Individuals of the type spe- epistomal sulcus protruding into the basal cies have been collected in Colombia, Ec- margin of the clypeus such that the epistomal uador, and Peru, whereas the second species lobe nearly reaches to the clypeal apex. In occurs in French Guiana (Engel and Brooks, addition, Chlerogelloides has a serrate inner 1999b). metatibial spur and a short malar space. DESCRIPTION: Female. Mandible with Genus Ischnomelissa Engel strong subapical tooth. Labral distal process narrowly triangular; basal elevation orbicular Ischnomelissa Engel, 1997b: 42. Type species: Is- chnomelissa zonata Engel, 1997b, monobasic and weak; lateral teeth absent. Prementum and original designation. greatly elongate. Galeal apex rounded; galeal comb absent; galeal base extending to base DIAGNOSIS: This group is similar to the of stipes. Hypostomal ridge feebly carinate more diverse genus Chlerogella, both genera on anterior half, carina disappearing on pos- having an elongate propodeum and a fre- terior half; anterior angle rounded. Malar quently elongate malar space that makes the space less than basal mandibular width. Ep- head much longer than wide. Ischnomelissa istomal sulcus acute, projecting deeply into differs by a densely pectinate inner metatibial clypeus as a thin line nearly reaching clypeal spur, an orthogonal epistomal sulcus, and a apex. Ocelli not greatly enlarged; ocellar fur- pronotum that is not dorsally inflated. row absent. Vertex short, approximately one DESCRIPTION: Female. Mandible with ocellar diameter in length. Preoccipital ridge moderately developed subapical tooth. La- rounded. Pronotal dorsal surface inflated, lat- bral distal process narrowly triangular; basal eral angle feebly present and obtuse; lateral elevation orbicular; teeth absent. Prementum ridge absent; dorsal ridge rounded. Meso- not greatly elongate. Galeal apex rounded; scutal anterior border weakly narrowed; me- galeal comb absent; galeal base extending to soscutal lip absent. Tegula oval. Anterior stipital base. Hypostomal ridge carinate; an- 40 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250 terior angle rounded. Length of malar space apex. The pectinate inner metatibial spur and usually less than basal mandibular width, ex- densely packed hamuli can also separate Me- cept in some species in which the length is galopta from other nocturnal augochlorines. equal to the basal mandibular width. Epis- DESCRIPTION: Female. Labral distal pro- tomal sulcus orthogonal. Ocelli not greatly cess broadly triangular; keel expanded basal- enlarged; ocellar furrow absent. Vertex not ly; basal elevation bilobed; teeth absent. Pre- expanded or ridged behind ocelli. Preoccip- mentum not greatly elongate. Galeal apex ital ridge rounded. Pronotal lateral angle not rounded; galeal comb absent; galeal base ex- produced, obtuse; dorsal ridge angled, but tending to base of stipes. Hypostomal ridge distinctly not carinate; lateral ridge rounded. carinate to weakly lamellate; anterior angle Mesoscutal anterior border rounded; meso- rounded. Malar space less than basal man- scutal lip rounded. Tegula oval. Anterior bas- dibular width. Epistomal sulcus acute, gently itarsal brush present. Basitibial plate border projecting into clypeus. Ocelli greatly en- posteriorly, margin obsolete anteriorly. Inner larged; ocellar furrow present. Vertex ex- hind tibial spur densely pectinate, with more panded behind ocelli. Preoccipital ridge than 10 teeth. Apex of marginal cell acute. rounded. Pronotal lateral angle not produced; Distal hamuli with irregular spacing pattern. dorsal ridge rounded; lateral ridge carinate. Basal area of propodeum as long as or longer Mesoscutal anterior border rounded; meso- than scutellum and metanotum combined, scutal lip rounded. Tegula oval. Anterior bas- nonstriate, smooth; propodeal pit narrow. itarsal brush present. Inner hind tibial spur Male. Mandible simple. Labrum without dis- pectinate. Apex of marginal cell truncate. tal process; basal area not notched. Antenna Distal hamuli closely and evenly spaced. long, extending beyond posterior border of Basal area of propodeum smooth or striate, propodeum; F1 much shorter than F2. Inner strongly declivitous; propodeal pit narrow. hind tibial spur serrate. Metasoma elongate. Male. Mandible simple. Labrum with distal S3ÐS5 unmodified. Apical margin of S6 process; basal area not notched. Antennae cleft. Apical margin of S7 with short median extending back to scutellum, metanotum, or process. Apical margin of S8 with median propodeal triangle; F2 much longer than F1. process; spiculum narrow. Proctiger unmod- Inner hind tibial spur serrate. Apical margin ified. Gonobasal bridge narrow; dorsal lobes of S4 emarginate medially, with lateral weak. Basal process of gonostylus and par- notches, setal patches bordering emargina- apenial lobes absent; gonostylus not divided tion. Apical margin of S5 emarginate, medial into ventral and dorsal gonostylar processes. surface with tubercle fitting into emargina- Ventral surface of penis valve with prong. tion of S4. Apical margin of S6 emarginate. REVISIONS: The genus has recently been re- Apical margin of S7 with median process bi- vised by Brooks and Engel (1998) who rec- lobed. Apical margin of S8 unmodified; spic- ognize six species. ulum narrow. Proctiger with anal filaments. DISTRIBUTION: Species are known from Gonobasal bridge narrow; dorsal lobes weak. Colombia and Ecuador. Parapenial lobe present; basal process of gonostylus absent. Ventral surface of penis Genus Megalopta Smith valve with prong.

DIAGNOSIS: This is a large genus of noc- Subgenus Megalopta Smith s.s. turnal bees exhibiting the typical characters Figures 6, 11, 16, 19, 36, 59, 66, 70, 73 of large ocelli and pale integumental pigmentation. Megalopta is related to the genus Megalopta Smith, 1853: 83. Type species: Me- ϭ Xenochlora, but this group has normal-sized galopta idalia Smith, 1853 [ Halictus amoen- ocelli, lacks the densely packed, single series us Spinola, 1853], designation of I.C.Z.N. (1966), petition by Michener and Moure of distal hamuli, and has stiff, black setae on (1964). Designation of Megalopta bituberculata the hindlegs. Among the other nocturnal gen- Smith, 1853, as the type species by Cockerell era, Megaloptidia and Megommation, Me- (1900) and Meade-Waldo (1916) is invalid. galopta lacks the strongly narrowed and Megaloptera Ashmead, 1899: 92. Lapsus calami. elongate prementum and the pointed galeal Megalopta (Megaloptella) Schrottky, 1906: 312. 2000 ENGEL: BEE TRIBE AUGOCHLORINI 41

Type species: Halictus ochrias Vachal, 1904, Subgenus Noctoraptor monobasic and original designation. Engel, Brooks, and Yanega Tmetocoelia Moure, 1943a: 481. Type species: Megalopta sulciventris Friese, 1926, original Megalopta (Noctoraptor) Engel, Brooks, and designation. Yanega, 1997: 12. Type species: Megalopta (Noctoraptor) byroni Engel, Brooks, and Yane- DIAGNOSIS: The nominate subgenus differs ga, 1997, original designation. from the cleptoparasitic species of Noctorap- DIAGNOSIS: See Diagnosis for Megalopta tor by the presence of scopal hairs, a basiti- s.s. (above). bial plate, and the normally developed man- DESCRIPTION: As for the genus with the dibles. The apical margin of the male clypeus following additions: Female. Mandible long, is white and F2 is about as long as F3 in scythe-shaped, without supplementary teeth. Megalopta s.s., while the clypeus is black Scopa absent. Basitibial plate absent. Male. and F2 is much shorter than F3 in Noctorap- Apical margin of clypeus black. Scape an- tor. teriorly black; F2 approximately two-thirds DESCRIPTION: As for the genus with the length of F3. Metasoma elongate. Apical following additions: Female. Mandible mul- margin of S5 broadly emarginate. tidentate, with supplementary teeth. Scopa REVISIONS: At present there are two spe- present. Basitibial plate bordered posteriorly, cies included in Noctoraptor; the type spe- margin obsolete anteriorly. Male. Apical cies and M.(Noctoraptor) noctifurax Engel margin of clypeus white. Scape anteriorly et al. white; F2 approximately equal in length to BIOLOGY: Noctoraptor species are clepto- F3. Metasoma oval to elongate. Apical mar- parasitic, based on their anatomy, presum- gin of S5 narrowly emarginate. ably on other species of Megalopta. REVISIONS: Friese (1926) revised the spe- DISTRIBUTION: The two known species oc- cies of Megalopta, although he included spe- cur in Panama (M. byroni) and Ecuador (M. cies of Megaloptidia and Megommation. noctifurax). There has been no modern treatment of the genus. Moure and Hurd (1987) listed 28 spe- Genus Megaloptidia Cockerell cies, although their Megalopta ianthina and Megalopta (Megaloptidia) Cockerell, 1900: 373. M. nigrofemorata are species of Xenochlora Type species: Megalopta (Megaloptidia) con- and were transferred to that genus by Engel tradicta Cockerell, 1900, monobasic and origi- et al. (1997). Megalopta intermedia, listed by nal designation. Sakagami (1979) is a nomen nudum (see ap- pend. 1). DIAGNOSIS: This is a genus of moderately large, robust, nocturnal bees generally resem- BIOLOGY: Species of Megalopta are nocturnal. Individuals can be readily captured at bling the larger nocturnal genus Megalopta; they were at one time included as a subge- lights just after dusk until just before dawn nus. The genus can be readily separated from (personal obs.). Brief accounts of the nesting Megalopta by the narrowed, elongate pre- biology of two Megalopta species have been mentum, the serrate inner metatibial spur, the given by Sakagami (1964) and Janzen enlarged compound eyes, and the irregularly (1968), while accounts of their seasonal spaced distal hamuli. Like most nocturnal abundance and nocturnal activity are given bees, species are generally pale in coloration by Wolda and Roubik (1986) and Roulston and have greatly enlarged ocelli. (1997) respectively. Immatures for two Pan- DESCRIPTION: Female. Mandible with amanian species are currently being studied strong subapical tooth. Labral distal process (Engel and Wcislo, in prep.), as are ﬁner de- narrowly triangular; basal elevation orbicu- tails of the nesting and social biology of lar; teeth absent. Prementum greatly elon- these species (W. T. Wcislo, personal com- gate. Galeal apex acute; galeal comb absent; mun.). galeal base extending to base of stipes. Hy- DISTRIBUTION: Megalopta s.s. ranges from postomal ridge carinate; anterior angle southern Brazil to Mexico. rounded. Length of malar space less than 42 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

basal mandibular width. Epistomal sulcus or- DIAGNOSIS: The genus Megaloptilla is thogonal. Compound eyes greatly enlarged, most similar to as well as most closely re- reaching above vertex in profile. Ocelli great- lated to Paroxystoglossa. Megaloptilla can ly enlarged; ocellar furrow absent. Vertex be separated from Paroxystoglossa, however, short, barely one ocellar diameter in length. by a rounded preoccipital ridge, a transverse Preoccipital ridge rounded. Pronotal lateral labral basal elevation, an acute marginal cell angle not produced, obtuse; dorsal ridge apex, and a very weakly narrowed anterior rounded; lateral ridge rounded. Mesoscutal border of the mesoscutum. anterior border rounded; mesoscutal lip DESCRIPTION: Female. Mandible bidentate, rounded. Tegula oval. Anterior basitarsal with weak supplementary teeth. Labral distal brush present. Basitibial plate bordered on all process narrowly triangular; basal elevation sides. Inner hind tibial spur serrate. Apex of transverse; teeth weak. Prementum not great- marginal cell truncate. Distal hamuli with ir- ly elongate. Galeal apex rounded; galeal regular spacing pattern. Basal area of pro- comb absent; galeal base extending to stipital podeum granular, nonstriate; propodeal pit base. Hypostomal ridge carinate; anterior an- narrow. Male. Mandible simple. Labrum gle rounded. Length of malar space less than without distal process; basal elevation ab- basal mandibular width. Epistomal sulcus or- sent. Antenna extending to posterior border thogonal. Ocelli not greatly enlarged; ocellar of mesoscutum; F2 shorter than F1. Inner furrow absent. Vertex not expanded or ridge hind tibial spur serrate. Metasoma oval. Api- behind ocelli. Preoccipital ridge rounded. cal margins of S4 and S5 concave. Apical Pronotal lateral angle not produced, obtuse; margin of S6 emarginate. Apical margin of dorsal ridge carinate, weakly lamellate in S7 with bilobed median process. Apical mar- some places; lateral ridge rounded. Meso- gin of S8 with short median projection; spic- scutal anterior border slightly narrowed; me- ulum narrow. Proctiger with anal filaments. soscutal lip rounded. Tegula oval. Anterior Gonobasal bridge narrow; dorsal lobes basitarsal brush present. Basitibial plate bor- strong. Basal process of gonostylus absent; dered on all sides. Inner hind tibial spur ser- parapenial lobe present. Ventral surface of rate. Apex of marginal cell acute. Distal penis valve with prong. hamuli with irregular spacing pattern. Basal REVISIONS: Megaloptidia was revised by area of propodeum subequal to scutellum, ru- Engel and Brooks (1998) who recognized gulose; propodeal pit narrow. Male. Mandi- three species; Megaloptidia contradicta (Cockerell), M. nocturna (Friese), and M. ble simple. Labrum without distal process; saulensis Engel and Brooks. no elevation. Antenna extending back to scutellum; F2 longer than F1. Inner hind tibial BIOLOGY: Species of Megaloptidia are nocturnal and can be captured at lights. spur serrate. Metasoma oval. Apical margin Nothing is known of their nesting or social of S4 unmodified. Apical margin of S5 biology. A female of M. nocturna has been weakly emarginate. Apical margin of S6 captured at flowers of the monocotyledon Di- emarginate. Apical margin of S7 with me- chorisandra ulei (Commelinaceae) (Engel dian projection. Apical margin of S8 with bi- and Brooks, 1998). lobed median projection; spiculum narrow. DISTRIBUTION: The genus occurs in north- Proctiger unmodified. Gonobasal bridge nar- ern South America. Individuals are known row; dorsal lobes weak. Parapenial lobe and from Brazil, Colombia, Ecuador, French Gui- basal process of gonostylus absent. Ventral ana, Peru, and Venezuela. surface of penis valve unmodified. REVISIONS: Engel and Brooks (1999a) have Genus Megaloptilla Moure and Hurd recently revised Megaloptilla and recognized two species: Megaloptilla callopis (Vachal) Megommation (Megaloptilla) Moure and Hurd, and M. byronella Engel and Brooks. 1987: 241. Type species: Halictus callopis Vachal, 1911, monobasic and original designa- DISTRIBUTION: Megaloptilla callopis is re- tion. corded from Colombia, Ecuador, and Peru Emgaloptilla Moure and Hurd, 1987: vi. Lapsus while the second species is presently known calami. only from Panama. 2000 ENGEL: BEE TRIBE AUGOCHLORINI 43

Genus Megommation Moure also distinguishes Cleptommation from Me- galoptina. DIAGNOSIS: This is a heterogenous group DESCRIPTION: As for the genus with the of taxa with strongly narrowed mouthparts following additions: Female. Mandible sim- (e.g., Megaloptidia, Ariphanarthra). Megom- ple, bladelike (fig. 15). Labral basal elevation mation in the broad sense can be most easily absent; teeth absent. Clypeal apex relatively separated from these other genera by normal- straight. Hypostomal ridge carinate. Ocelli sized compound eyes (greatly enlarged in not greatly enlarged. Pronotal dorsal ridge Megaloptidia) and normal maxillary palpi rounded. Mesoscutal anterior border round- (greatly lengthened in Ariphanarthra). See ed; mesoscutal lip rounded. Scopa absent. also the diagnoses for Megaloptidia, Ari- Basitibial plate absent. Inner hind tibial spur phanarthra, and Micrommation. serrate. Male. Labrum without distal process; DESCRIPTION: Female. Labral distal pro- basal area not notched. Antenna extending cess broadly triangular. Prementum greatly back to scutellum or metanotum; F1 longer elongate. Galeal apex acute; galeal comb ab- than F2. Dense patch of plumose setae at sent; galeal base extending to stipital base. propodeal spiracle. Metasoma slightly elon- Hypostomal ridge carinate to weakly lamel- gate. Apical margins of S2 and S3 with slight late; anterior angle rounded. Length of malar median projection. Apical margins of S4 and space less than basal mandibular width. Ep- S5 unmodified. Apical margins of S7 and S8 istomal sulcus orthogonal. Ocellar furrow ab- with median process. sent. Vertex not expanded or ridged behind REVISIONS: There is only one included spe- ocelli. Preoccipital ridge rounded. Pronotal cies, although there may be a second species lateral angle not produced, obtuse; lateral (R. W. Brooks, personal commun.; D. Yane- ridge rounded. Mesoscutal anterior border ga, personal commun.). rounded; mesoscutal lip rounded. Tegula BIOLOGY: Based on the adult anatomy, oval. Anterior basitarsal brush absent. Apex Cleptommation is cleptoparasitic, perhaps on of marginal cell acute. Distal hamuli with ir- other species of Megommation (most likely regular spacing pattern. Male. Mandible sim- Megaloptina or Stilbochlora). ple. Inner hind tibial spur serrate. Gonobasal DISTRIBUTION: Cleptommation occurs in bridge narrow; dorsal lobes strong. Apical Bolivia, Brazil, Costa Rica, Ecuador, Pana- margin of S6 emarginate. Proctiger with anal ma, and Peru. filaments. Basal process of gonostylus absent; parapenial lobe present; dorsal process of gonostylus partially membranous. Ventral Subgenus Megaloptina Eickwort surface of penis valve with prong. Figures 5, 12Ð13 Megommation (Megaloptina) Eickwort, 1969a: Subgenus Cleptommation 441. Type species: Augochlora (Pseudaugo- Engel, Brooks, and Yanega chloropsis) ogilviei Cockerell, 1930, original Figures 3, 12, 13, 15, 32, 38, 45, 51 designation.

Megommation (Cleptommation) Engel, Brooks, DIAGNOSIS: This subgenus is most similar and Yanega, 1997: 19. Type species: Megalopta to Cleptommation, particularly in the male minuta Friese, 1926, monobasic and original sex where both share the presence of dense, designation. plumose setae surrounding the propodeal spi- DIAGNOSIS: This cleptoparasitic subgenus racle. Cleptommation, however, lacks a sco- can be quickly identified by the simple, pa, has a bladelike mandible, lacks a basiti- bladelike mandibles (fig. 15), the absence of bial plate, and differs in the structure of the a basitibial plate, and the absence of a scopa. male metasomal sterna. Megaloptina differs It is most similar to Megaloptina, both shar- from Stilbochlora in the presence of a pro- ing the dense, plumose setae surrounding the podeal tuft of setae in males and the short propodeal spiracle in males. Megaloptina, basitibial plate with weakly developed bor- however, is not parasitic and has a well-de- ders. veloped scopa, a basitibial plate, and normal DESCRIPTION: As for the genus with the mandibles. The serrate inner metatibial spur following additions: Female. Mandible with 44 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250 moderate subapical tooth. Labral basal ele- Metasoma oval. Apical margin of S2 unmod- vation orbicular; teeth weak. Clypeal apex ified. Apical margin of S3 with median pro- concave (fig. 12). Hypostomal ridge carinate. jection. Apical margin of S4 with median Ocelli not greatly enlarged. Pronotal dorsal projection. Apical margin of S5 emarginate. ridge rounded. Scopa present. Basitibial plate Apical margin of S7 with bilobed median extremely short, borders faint. Inner hind tib- projection. Apical margin of S8 unmodified. ial spur pectinate. Male. Labrum without dis- REVISIONS: Megommation proper contains tal process. Antenna extending back to scu- only the type species. tellum; F1 longer than F2. Dense patch of BIOLOGY: The nesting biology of M. insig- plumose setae at propodeal spiracle. Meta- ne (Smith) was studied by Jo¬rgensen (1912), soma oval. Apical margin of S2 unmodified. Michener and Lange (1958b), and Sakagami Apical margin of S3 with median projection. and Moure (1967). Apical margin of S4 unmodified. Apical DISTRIBUTION: Megommation insigne oc- margin of S5 emarginate. Apical margin of curs in northern Argentina, southern Brazil, S7 with median projection. Apical margin of and Paraguay. S8 unmodified. REVISIONS: There are presently only two Subgenus Stilbochlora described species, Megommation festivagum Engel, Brooks, and Yanega (Dalla Torre) and M. ogilviei (Cockerell), and Megommation (Stilbochlora) Engel, Brooks, and at least one undescribed species (personal Yanega, 1997: 15. Type species: Megommation obs.). (Stilbochlora) eickworti Engel, Brooks, and DISTRIBUTION: Megaloptina is known from Yanega, 1997, monobasic and original desig- Ecuador, Guyana, and Brazil. nation. DIAGNOSIS: Refer to Diagnosis for subge- Subgenus Megommation Moure s.s. nus Megaloptina (above). Megommation Moure, 1943a: 479. Type species: DESCRIPTION: As for the genus with the Halictus insignis Smith, 1853, monobasic and following additions: Female. Mandible with original designation. strong subapical tooth. Labral basal elevation Megammation Sakagami and Michener, 1962: 88. orbicular; teeth weak. Clypeal apex relatively Lapsus calami. straight. Hypostomal ridge carinate. Ocelli not greatly enlarged. Pronotal dorsal ridge DIAGNOSIS: This monobasic subgenus is carinate. Scopa present. Basitibial plate not nocturnal in habit and exhibits the typical shortened, bordered posteriorly. Inner hind features of nocturnal bees: pale integumental tibial spur pectinate. Male. Labrum with dis- pigmentation and greatly enlarged ocelli. The tal process, without basal elevation. Antenna nocturnal characters can readily distinguish extending back to scutellum; F2 longer than Megommation proper from the other subgen- F1. Metasoma oval. Apical margins of S2 era. This group can be easily confused with and S3 unmodified. Apical margin of S4 with the related nocturnal genus Megaloptidia, but median projection. Apical margin of S5 un- Megaloptidia has the compound eyes greatly modified. Apical margin of S7 unmodified. enlarged and the marginal cell apex feebly Apical margin of S8 with median projection. truncate and appendiculate. REVISIONS: There is only the one included DESCRIPTION: As for the genus with the species. following additions: Female. Mandible with DISTRIBUTION: Megommation eickworti is moderate subapical tooth. Labral basal ele- found in Brazil, Bolivia, Colombia, Ecuador, vation orbicular; teeth weak. Clypeal apex and Peru. concave. Hypostomal ridge weakly lamellate on posterior half. Ocelli greatly enlarged. Genus Micrommation Moure Pronotal dorsal ridge rounded. Scopa present. Basitibial plate extremely short, borders Micrommation Moure, 1969: 247. Type species: faint. Inner hind tibial spur serrate. Male. La- Micrommation larocai Moure, 1969, monoba- brum without distal process. Antenna extend- sic and original designation. ing back to scutellum; F1 longer than F2. DIAGNOSIS: This genus appears to be sim- 2000 ENGEL: BEE TRIBE AUGOCHLORINI 45 ilar to the nocturnal genus Megaloptidia; mun.) who has examined the as of yet un- however, Micrommation does not have en- described male of Micrommation. For further larged ocelli nor a feebly truncate marginal details on Micrommation morphology refer cell apex. Likewise, Micrommation is not to Yanega (in prep.). nocturnal and has brilliant metallic-green integument, whereas Megaloptidia has the typ- Genus Neocorynura Schrottky ically pale integument pigmentation of noc- Figure 64 turnal bees. Cacosoma Smith, 1879: 39. Type species: Caco- DESCRIPTION: Female. Mandible with soma discolor Smith, 1879, designation of strong subapical tooth. Labral distal process Sandhouse (1943). Nomen praeoccupatum (nec narrowly triangular; basal elevation orbicu- Cacosoma Felder In Felder and Rogenhofer, lar; teeth absent. Prementum greatly elon- 1874 [Lepidoptera: Zygaenidae]). gate. Galeal comb absent. Hypostomal ridge Neocorynura Schrottky, 1910: 540. Nomen novum carinate; anterior angle rounded. Length of pro Cacosoma Smith, 1879. Type species: au- malar space less than basal mandibular tobasic with Cacosoma Smith, 1879. width. Epistomal sulcus orthogonal. Ocelli Neocorynura (Neocorynuroides) Eickwort, 1969a: not greatly enlarged; ocellar furrow absent. 404. Type species: Halictus rhytis Vachal, Vertex not expanded or ridged behind ocelli. 1904, monobasic and original designation. NEW SYNONYMY. Preoccipital ridge rounded. Tegula oval. An- terior basitarsal brush present. Basitibial DIAGNOSIS: The speciose genus Neocory- plate strongly rimmed on all borders. Inner nura contains a heterogenous assemblage of hind tibial spur serrate. Apex of marginal cell primitive Augochlorina. The combination of acute. Distal hamuli with irregular spacing an obtuse epistomal sulcus, frequently nar- pattern. Basal area of propodeum as long as rowed mesoscutal anterior margin, carinate scutellum, smooth; propodeal pit narrow. preoccipital ridge, pectinate inner metatibial Male. Mandible simple. Labral distal process spur, and strongly bordered basitibial plate absent. Antenna extending to posterior bor- distinguishes the genus from similar genera der of mesoscutum; F1 longer than F2. Inner such as Andinaugochlora and Neocorynurel- hind tibial spur serrate. Metasoma oval. Api- la. cal margins of S4 and S5 concave. Apical DESCRIPTION: Female. Mandible with margin of S6 emarginate. Apical margin of moderate to strong subapical tooth. Labral S7 with bilobed median process. Apical mar- distal process narrowly triangular; basal ele- gin of S8 with short median projection; spic- vation transverse; teeth absent. Prementum ulum narrow. Proctiger with anal filaments. not greatly elongate. Galeal apex rounded; Gonobasal bridge narrow; dorsal lobes galeal comb absent; galeal base extending to strong. Basal process of gonostylus absent; base of stipes. Hypostomal ridge carinate; parapenial lobe present. Ventral surface of anterior angle rounded. Length of malar penis valve with prong. space less than basal mandibular width. Ep- REVISIONS: Only the type species is cur- istomal sulcus obtuse. Ocelli not greatly en- rently recognized in Micrommation. Douglas larged; ocellar furrow absent. Vertex not ex- Yanega is preparing a new description of panded or ridged behind ocelli. Preoccipital Moure’s type as well as of the only known ridge rounded to carinate. Pronotal lateral an- male (Yanega, in prep.). gle variable; dorsal ridge carinate; lateral DISTRIBUTION: Micrommation is only ridge carinate to lamellate. Mesoscutal ante- known from the type locality in Parana«, Bra- rior border frequently narrowed; mesoscutal zil (at 900 m). lip rounded to angled. Tegula oval. Anterior COMMENTS: I have not seen specimens of basitarsal brush present. Basitibial plate bor- Micrommation, which is known only on the dered on all sides. Inner hind tibial spur pec- basis of the female holotype, a single female tinate. Apex of marginal cell acute. Distal paratype, and one nontype male. My generic hamuli with irregular spacing pattern. Basal diagnosis is taken from Moure’s original de- area of propodeum granular, striate, or ru- scription and figures as well as information gose; propodeal pit narrow. Male. Mandible provided to me by D. Yanega (personal com- simple. Labrum with distal process; basal 46 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250 area not notched. Antenna extending back to obtuse. Ocelli not greatly enlarged; ocellar propodeum or beyond; F2 longer than F1. furrow absent. Vertex slightly expanded be- Inner hind tibial spur serrate. Metasoma fre- hind ocelli. Preoccipital ridge rounded or quently petiolate. Apical margins of S4 and weakly carinate. Pronotal lateral angle not S5 unmodified. Apical margin of S6 emar- produced, obtuse; dorsal ridge rounded; lat- ginate. Apical margin of S7 unmodified. Api- eral ridge angled. Mesoscutal anterior border cal margin of S8 with median projection; rounded; mesoscutal lip rounded. Tegula spiculum narrow. Proctiger unmodified. oval. Anterior basitarsal brush present. Ba- Gonobasal bridge narrow; dorsal lobes sitibial plate with well-developed borders. strong. Parapenial lobe and basal process of Inner hind tibial spur pectinate. Apex of mar- gonostylus absent. Ventral surface of penis ginal cell truncate. Distal hamuli with irreg- valve with or without prong, never with keel. ular spacing pattern. Basal area of propo- REVISIONS: There has been no revision of deum as long as metanotum, striate; propo- Neocorynura. There are presently 67 valid deal pit narrow. Male. Mandible simple. La- species. brum with short distal process; basal BIOLOGY: The biology of Neocorynura is elevation not notched. Antennae extending varied with species nesting in the soil (Mich- back to propodeum; F2 longer than F1. Inner ener, 1977; Michener and Lange, 1958b; hind tibial spur serrate. Metasoma elongate. Michener et al., 1966; Sakagami and Moure, Apical margins of S4 and S5 weakly de- 1967) or rotten wood (Lu¬derwaldt, 1911; pressed. Apical margin of S6 deeply emar- Schremmer, 1979). Neocorynura colombiana ginate. Apical margin of S7 with broad me- Eickwort and N. erinnys (Schrottky) may be dian projection. Apical margin of S8 with semisocial (Lu¬derwaldt, 1911; Schremmer, narrow median projection; spiculum broad. 1979). Immatures have been described for N. Proctiger unmodified. Gonobasal bridge nar- colombiana by Eickwort (1979a). row; dorsal lobes strong. Parapenial lobe and DISTRIBUTION: The genus ranges from Ar- basal process of gonostylus absent; ventral gentina to Mexico. Although Neocorynura is process twisted apically. Ventral surface of unknown from the West Indies today, one penis valve unmodified. fossil species, N. electra Engel, has been REVISIONS: The genus currently contains found as a Miocene amber inclusion from the three species; Neocorynurella cosmetor Dominican Republic (Engel, 1995c). (Vachal) (see appendix 1), N. seeleyi, and N. virida Engel and Klein (nomen emendatum, Genus Neocorynurella Engel from N. viridis). DISTRIBUTION: The genus occurs in moun- Neocorynurella Engel in Engel and Klein, 1997: tainous areas of Colombia and Venezuela. 156. Type species: Neocorynurella seeleyi En- COMMENTS: Thanks to Frank Koch gel and Klein, 1997, original designation. (ZMHB) I have had the opportunity to ex- Vachalius Moure, 1999: 74. Type species: Ha- lictus cosmetor Vachal, 1911, monobasic and amine the lectotype of Halictus cosmetor original designation. NEW SYNONYMY. designated by Moure (1999) and to confirm the synonymy of Vachalius with Neocory- DIAGNOSIS: The genus is most similar to nurella. Moure did not have a male, nor is Andinaugochlora. Refer to the diagnosis of one present in the original series; however, Andinaugochlora for features separating the several females that are clearly conspecific two genera. with the types are in the AMNH and SEMC DESCRIPTION: Female. Mandible with (see appendix 1). weak subapical tooth. Labral distal process narrowly triangular; basal elevation orbicu- Genus Oligochlora Engel lar; teeth absent. Prementum not greatly elongate. Galeal apex rounded; galeal comb DIAGNOSIS: This fossil genus is not without absent; galeal base extending to base of sti- affinities to Neocorynura but differs by the pes. Hypostomal ridge carinate; anterior an- broadly rounded mesoscutum, the rounded gle rounded. Length of malar space less than preoccipital ridge, the unproduced pronotal basal mandibular width. Epistomal sulcus lateral angle, and in some species (Oligo- 2000 ENGEL: BEE TRIBE AUGOCHLORINI 47 chlora s.s.) by the obsolescent anterior bor- riorly. Anterior surface T1 weakly modified der to the basitibial plate and the presence of into an acarinarium. Male. Unknown. a weak acarinarium on T1. REVISIONS: At present this group contains DESCRIPTION: Female. Mandible with three species. The subgenus was newly di- strong subapical tooth. Labral distal process agnosed by Engel (1997a) based on the dis- narrowly triangular; lateral teeth strong. Pre- covery of a third species. mentum not greatly elongate. Galeal apex PALEOBIOLOGY: Two of the three species of rounded. Hypostomal ridge carinate; anterior Oligochlora are associated with astigmatid angle rounded. Length of malar space less mites (Engel, 1996b; Fain et al., 1999). Mite- than basal mandibular width. Epistomal sul- bee associations are common, although poor- cus obtuse. Ocelli not greatly enlarged; ocel- ly understood (Eickwort, 1979b, 1994; Fain lar furrow absent. Vertex not expanded nor et al., 1999). Oligochlora, like the genus ridged behind ocelli. Preoccipital ridge Thectochlora (see below), possesses an aca- rounded. Pronotal lateral angle not produced, rinarium on the first metasomal tergum sug- acute to obtuse; dorsal ridge carinate; lateral gesting a mutualistic association between ridge rounded or carinate. Mesoscutal ante- these ancient bees and mites. rior border rounded; mesoscutal lip rounded. Tegula oval. Anterior basitarsal brush pre- Soliapis, new subgenus sent. Inner hind tibial spur pectinate. Apex Figure 82 of marginal cell feebly truncate. Distal ham- TYPE SPECIES: Oligochlora (Soliapis) roz- uli with irregular spacing pattern. Basal area eni, new species (appendix 1). of propodeum smooth to weakly striate and DIAGNOSIS: See Diagnosis for Oligochlora granular, not elongate; propodeal pit narrow. s.s. (above). Male. Unknown. DESCRIPTION: As for the genus with the DISTRIBUTION: The genus is presently following additions: Female. Basitibial plate known only from the Miocene amber depos- strongly bordered on all sides. Anterior sur- its of the Dominican Republic (Iturralde-Vi- face of T1 unmodified, not developed into an nent and MacPhee, 1996). The Eocene dates acarinarium. Male. Unknown. which have been estimated for some of these ETYMOLOGY: The subgeneric name is a mines based on NMR studies (Lambert et al., combination of the Latin words sola (mean- 1985) have not been substantiated. ing ‘‘alone’’) and apis (meaning ‘‘bee’’) and is a reference to the lack of an acarinarium Subgenus Oligochlora Engel s.s. and thereby the mutualism with mites in these species. Oligochlora Engel, 1996b: 336. Type species: REVISIONS: Two species are included in Oligochlora eickworti Engel, 1996b, original this subgenus. The first is described in ap- designation. pendix 1 as the type species of the group DIAGNOSIS: This subgenus differs from So- while a second is treated by Engel and Right- liapis by the presence of an acarinarium and myer (in press). the obsolescent anteior border of the basitibial plate. The only other augochlorine with Genus Paroxystoglossa Moure an acarinarium is the South American genus Paroxystoglossa Moure, 1940: 59. Type species: Thectochlora. Oligochlora is distinguished Oxystoglossa jocasta Schrottky, 1911, original from Thectochlora by an unproduced pron- designation. otal lateral angle, absence of a lamella on the Paraoxystoglossa Roubik, 1989: 392. Lapsus cal- pronotal dorsal ridge, a broadly rounded me- ami. soscutum, absence of a dorsal hook on the DIAGNOSIS: Refer to the Diagnosis for Me- mesotrochanter, and abence of dense tomen- galoptilla (above). tum on the basal area of the propodeum. DESCRIPTION: Female. Mandible with DESCRIPTION: As for the genus with the moderate subapical tooth. Labral distal pro- following additions: Female. Basitibial plate cess narrowly triangular; basal elevation or- bordered posteriorly, margin obsolete ante- bicular; teeth weak. Prementum not greatly 48 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250 elongate. Galeal apex rounded; galeal comb marginal cell apex. Pereirapis differs from absent; galeal base extending to stipital base. Augochlorella in the presence of a large bas- Hypostomal ridge carinate; anterior angle al lobe on the inner metatibial spur. Perei- rounded. Length of malar space less than rapis are small, brilliant metallic bees that basal mandibular width. Epistomal sulcus are frequently green, with some populations obtuse. Ocelli not greatly enlarged; ocellar slightly bluish and others more cupreous. furrow absent. Vertex not expanded or ridged DESCRIPTION: Female. Mandible with behind ocelli. Preoccipital ridge angled, dis- moderate to strong subapical tooth. Labral tinctly not carinate. Pronotal lateral angle distal process narrowly triangular; basal ele- variable; dorsal ridge carinate; lateral ridge vation orbicular; teeth strong. Prementum not rounded to angled. Mesoscutal anterior bor- greatly elongate. Galeal apex rounded; galeal der narrowed; mesoscutal lip angled. Tegula comb absent; galeal base extending to base oval. Anterior basitarsal brush present. Ba- of stipes. Hypostomal ridge carinate; anterior sitibial plate bordered on all sides. Inner hind angle rounded. Length of malar space less tibial spur serrate. Apex of marginal cell than basal mandibular width. Epistomal sul- truncate. Distal hamuli with irregular spacing cus orthogonal. Ocelli not greatly enlarged; pattern. Basal area of propodeum weakly ocellar furrow absent. Vertex not expanded striate to rugulose; propodeal pit narrow. or ridge behind ocelli. Preoccipital ridge car- Male. Mandible simple. Labrum with distal inate. Pronotal lateral angle not produced, or- process; basal area notched. Antenna long, thogonal to obtuse; dorsal ridge carinate; lat- extending back to propodeum; F2 longer eral ridge angled. Mesoscutal anterior border than F1. Inner hind tibial spur serrate. Me- rounded; mesoscutal lip rounded. Tegula tasoma oval to slightly elongate. Apical mar- oval. Anterior basitarsal brush absent. Basi- gins of S4ÐS6 emarginate. Apical margins of tibial plate bordered on all sides. Inner hind S7 and S8 with median projection. Proctiger tibial spur serrate, basally with an expanded unmodified. Gonobasal bridge narrow; dorsal tooth (fig. 49). Apex of marginal cell acute. lobes strong. Parapenial lobe and basal pro- Distal hamuli with irregular spacing pattern. cess of gonostylus absent. Ventral surface of Basal area of propodeum striate; propodeal penis valve with prong. pit narrow. Male. Mandible simple. Labrum REVISIONS: The genus was revised by with distal process; basal area not notched. Moure (1960), who recognized nine species. Antenna extending back to scutellum; F2 ap- BIOLOGY: The nesting biology of Parox- ystoglossa has been studied by Michener and proximately equal in length to F1. Inner hind Lange (1958a, 1958b) and Michener and tibial spur serrate. Metasoma oval. Apical Seabra (1959). margin of S4 concave. Apical margin of S5 unmodified. Apical margin of S6 emarginate. DISTRIBUTION: Species of the genus range in Argentina, southern Brazil, and Paraguay. Apical margin of S7 unmodified. Apical margin of S8 with median process; spiculum narrow. Proctiger unmodified. Gonobasal Genus Pereirapis Moure bridge narrow; dorsal lobes strong. Basal Figures 27, 49, 52, 62 process of gonostylus present; parapenial Pereirapis Moure, 1943a: 461. Type species: Pe- lobe absent. Ventral surface of penis valve reirapis rhizophila Moure, 1943a [ϭ Halictus unmodified. semiauratus Spinola, 1853], monobasic and REVISIONS: There is only the one included original designation. species (see appendix 1). Pereirapsis Alves dos Santos, 1997: 6. Lapsus BIOLOGY: The nesting biology, phenology, calami. and primitively eusocial behavior of Perei- DIAGNOSIS: Pereirapis generally resembles rapis was investigated by Oliveira Campos species of the more common genera Augo- (1980). McGinley (1989) cited the descrip- chlorella and Augochlora. It can be separated tion of immature stages for Pereirapis ed- from the latter genus by the absence of a entata (Michener), but this is a species of deeply acute epistomal sulcus that projects Augochlorella. into the base of the clypeus and by the acute DISTRIBUTION: Pereirapis ranges from 2000 ENGEL: BEE TRIBE AUGOCHLORINI 49

Mexico eastward into the West Indies and REVISIONS: The genus has never been re- south to southern Brazil. vised. Moure and Hurd (1987) list seven named species while an eighth species is pro- Genus Pseudaugochlora Michener posed in appendix 1. Figures 10, 22Ð23, 63, 65, 71 BIOLOGY: The nesting biology of P. graminea and P. sordicutis (Vachal) has been Caenaugochlora (Pseudaugochlora) Michener, investigated by Michener and Kerfoot [1967: 1954b: 77. Type species: Halictus nigromargin- as species of Pseudaugochloropsis, the latter atus Spinola, 1841 [ϭ Megilla graminea Fabri- cius, 1804], original designation. under the name P. nigerrima (Friese)]. DISTRIBUTION: Pseudaugochlora extends DIAGNOSIS: Pseudaugochlora is superfi- from the southwestern United States to Ar- cially most similar to Caenaugochlora. The gentina and east into the West Indies. genus can be separated from Caenaugochlo- COMMENTS: This group has generally been ra by the presence of a strong ridge on the known under the name Pseudaugochloropsis vertex, pointed galeal apex, and hooked apex by recent authors (e.g., Eickwort, 1967; of F11 in males. Michener, 1974; Eickwort and Sakagami, DESCRIPTION: Female. Mandible with 1979; Moure and Hurd, 1987; Roubik, 1989; strong subapical tooth. Labral distal process Michener et al., 1994; Radchenko and Pe- narrowly triangular; basal elevation orbicu- senko, 1994b; Griswold et al., 1995; Dan- lar; teeth weak. Prementum not greatly elon- forth and Eickwort, 1997). However, the type gate. Galeal apex acute; galeal comb absent; species of Pseudaugochloropsis is a species galeal base extending to stipital base. Hy- of Augochloropsis (Paraugochloropsis), and postomal ridge carinate; anterior angle thus the valid name for the genus is Pseu- rounded. Length of malar space less than daugochlora (see also discussions in Mich- basal mandibular width. Epistomal sulcus or- ener, 1954b, 1994). As if this confusion were thogonal. Ocelli not greatly enlarged; ocellar not enough, the type species of this genus has furrow absent. Vertex with ridge behind ocel- been consistently cited as Halictus nigro- li. Preoccipital ridge rounded. Pronotal lat- marginatus Spinola, 1851 (Eickwort, 1969a; eral angle not produced, obtuse; dorsal ridge Michener, 1994, 1997), which is obviously a carinate; lateral ridge rounded. Mesoscutal homonym of the actual type species of Pseu- anterior border rounded; mesoscutal lip daugochlora (namely, H. nigromarginatus rounded. Tegula oval. Anterior basitarsal Spinola, 1841). Spinola’s 1851 nigromargin- brush present. Basitibial plate bordered on all atus is in actuality a species of Caenohalictus sides. Inner hind tibial spur pectinate. Apex (ϭ Caenohalictus oblitus Moure and Hurd, of marginal cell truncate. Distal hamuli with 1987). irregular spacing pattern. Basal area of propodeum rugose; propodeal pit narrow. Male. Genus Temnosoma Smith Mandible simple. Labrum with distal pro- Figures 7Ð9, 20, 30, 40, 56Ð57, 67, 69, 76 cess; basal area notched. Antenna extending Temnosoma Smith, 1853: 38. Type species: Tem- back to scutellum or metanotum; F2 approx- nosoma metallicum Smith, 1853, monobasic. imately equal in length to F1; F11 hooked at Tamnosoma Taschenberg, 1883: 93. Lapsus cala- apex. Inner hind tibial spur serrate. Metaso- mi. ma oval. Apical margins of S4 and S5 emar- Micraugochlora Schrottky, 1909a: 138. Type spe- ginate; S4 with medio-apical setal patch; S5 cies: Micraugochlora sphaerocephala Schrott- with lateroapical setal patches. Apical margin ky, 1909a, monobasic. of S6 emarginate. Apical margin of S7 pro- Temnosoma (Temnosomula) Ogloblin, 1953: 2. Type species: Temnosoma (Temnosomula) pla- duced laterally. Apical margin of S8 with ϭ median process; spiculum broad. Proctiger tensis Ogloblin, 1953 [ Micraugochlora sphaerocephala Schrottky, 1909a], monobasic with anal filaments. Gonobasal bridge nar- and original designation. Proposed as new again row or membranous; dorsal lobes strong. in Ogloblin, 1954. Parapenial lobe present; basal process of Themnosoma Sakagami, 1979: 83. Lapsus calami. gonostylus absent. Ventral surface of penis Microaugochlora Schlindwein, 1998: 52. Lapsus valve unmodified. calami. 50 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

DIAGNOSIS: This is one of the most dis- expanded with long setae; dorsal process re- tinctive genera of Augochlorini. The coarse duced, small ridge with setae. Ventral surface sculpturing of the body gives these clepto- of penis valve unmodified, with large dorsal parasitic bees the general appearance of process. chrysidid wasps. Among augochlorine gen- REVISIONS: Friese (1925) revised the spe- era, Temnosoma’s appearance comes closest cies of Temnosoma known at the time rec- to species of Glyptochlora. Temnosoma can ognizing six species. Moure and Hurd (1987) be separated from Glyptochlora by an oval list seven species. There has been no modern tegula, absence of a scopa, the depressed api- treatment of the genus. cal margins of the first two metasomal terga, BIOLOGY: The biology of Temnosoma re- absence of a medio-apical cleft in the fifth mains unstudied, but the species are presum- metasomal tergum of females, and the pres- ably parasitic, perhaps on species of the ge- ence of a broad division in the seventh me- nus Augochloropsis or Augochlora (Miche- tasomal tergum of males, among other char- ner, 1978a). acters. DISTRIBUTION: Temnosoma occupies a DESCRIPTION: Female. Body coarsely wide distribution with species found from as punctured. Mandible simple. Labral distal far south as northern Argentina, throughout process quadrate; basal elevation weakly bi- South and Central America, north to Arizona lobed; keel absent; teeth absent. Prementum (Timberlake, 1958), as well as occurring in not greatly elongate. Galeal apex rounded; Cuba and Jamaica (Eickwort, 1988). One po- galeal comb absent; galeal base extending to tential host, Augochloropsis, does not occur base of stipes. Hypostomal ridge carinate; in the West Indies, but the other, Augochlora, anterior angle rounded. Length of malar does. space less than basal mandibular width. Ep- istomal sulcus obtuse. Ocelli not greatly en- Genus Thectochlora Moure larged; ocellar furrow absent. Vertex expand- Figures 1, 42, 47, 53Ð55, 72 ed behind ocelli. Preoccipital ridge sharply angled and weakly carinate. Pronotal lateral Thectochlora Moure, 1940: 51. Type species: Halictus alaris Vachal, 1904, monobasic and angle not produced, obtuse; dorsal ridge car- original designation. inate; lateral ridge rounded. Mesoscutal anterior border rounded; mesoscutal lip weak DIAGNOSIS: Thectochlora resembles to and rounded. Tegula oval; coarsely punc- some degree the genera of the Augochlora- tured. Anterior basitarsal brush absent. Me- group (e.g., Augochlorella, Pereirapis), but sofemoral brush absent. Scopa absent. Peni- differs most notably in the presence of a cillus absent. Basitibial plate absent. Inner well-developed acarinarium on the first me- hind tibial spur serrate. Apex of marginal cell tasomal tergum, presence of a strong dorsal acute. Distal hamuli with irregular spacing hook on the mesotrochanter, a lamellate dor- pattern. Basal area of propodeum strongly sal pronotal ridge, and the presence of dense striated. Apical margins of T1 and T2 strong- tomentum on the basal area of the propo- ly depressed and rimmed; apical margin of deum. The genus also differs in the combi- T5 without cleft. Male. Mandible simple. La- nation of an obtuse epistomal sulcus and a brum with quadrate distal process. Antenna pectinate inner metatibial spur. extending back to mesoscutum; F2 longer DESCRIPTION: Female. Mandible with than F1. Inner hind tibial spur serrate. Me- weak subapical tooth. Labral distal process tasoma oval. Apical margin of T7 with broad narrowly triangular; basal elevation orbicu- division. Apical margins of S4 and S5 un- lar; teeth strong. Prementum not greatly elon- modified. Apical margin of S6 emarginate. gate. Galeal apex rounded; galeal comb ab- Apical margin of S7 medially unsclerotized. sent; galeal base extending to base of stipes. Apical margin of S8 with median projection; Hypostomal ridge carinate; anterior angle spiculum narrow. Proctiger unmodified. rounded. Length of malar space less than Gonobasal bridge broad; dorsal lobes weak. basal mandibular width. Epistomal sulcus Gonostylus with parapenial lobe present; obtuse. Ocelli not greatly enlarged; ocellar basal process absent; ventral process greatly furrow absent. Vertex not expanded or ridged 2000 ENGEL: BEE TRIBE AUGOCHLORINI 51 behind ocelli. Preoccipital ridge rounded. ochlora, however, is not nocturnal and lacks Pronotal lateral angle produced, obtuse; dor- the greatly enlarged ocelli of Megalopta, has sal ridge lamellate; lateral ridge rounded. irregularly spaced distal hamuli along the an- Mesoscutal anterior border narrowed; meso- terior margin of the hind wing, and has stiff, scutal lip lamellate. Tegula oval. Anterior ba- black setae on the hindlegs. sitarsal brush absent. Mesotrochanter with DESCRIPTION: Female. Mandible bidentate, strong dorsal hook. Basitibial plate rimmed with supplementary teeth. Labral distal pro- on posterior edge, anterior border obsolete. cess broadly triangular; basal elevation bi- Inner hind tibial spur pectinate. Apex of mar- lobed; keel expanded basally; teeth absent. ginal cell truncate. Distal hamuli with irreg- Prementum not greatly elongate. Galeal apex ular spacing pattern. Basal area of propo- rounded; galeal comb absent. Hypostomal deum striate, extremely short, covered in ridge lamellate; anterior angle rounded. Ma- dense tomentum. Anterior surface T1 modi- lar space less than basal mandibular width. fied into an acarinarium (a depressed shiny Epistomal sulcus acute, gently protruding area bordered basally and partly on lateral into clypeus. Ocelli not greatly enlarged; margins by long plumose hairs). Male. Man- ocellar furrow present. Vertex expanded be- dible simple. Labrum with weak distal pro- hind ocelli. Preoccipital ridge rounded. Pron- cess; basal area not notched. Antennae ex- otal lateral angle not produced; dorsal ridge tending just beyond posterior margin of pro- angled to weakly carinate; lateral ridge cari- podeum; F2 much longer than F1. Inner hind nate. Mesoscutal anterior border rounded; tibial spur serrate. Metasoma oval. S4 with mesoscutal lip rounded. Tegula oval. Ante- lateral processes on apical margin possessing rior basitarsal brush present. Metatibia and strong setae at apices; central patch of dense metabasitarsus covered with stiff, black se- setae on disc. Apical margin of S5 emargin- tae. Basitibial plate rimmed posteriorly, an- ate. Apical margin of S6 emarginate. S7 un- terior border obsolescent. Inner hind tibial modified. Apical margin of S8 with medial spur pectinate. Apex of marginal cell acute. process; spiculum narrow. Proctiger unmod- Distal hamuli with irregular spacing pattern. ified. Gonobasal bridge narrow; dorsal lobes Basal area of propodeum about as long as strong. Gonostylus with parapenial lobe pre- metanotum and strongly declivitous. Male. sent; basal process absent. Penis valve with Unknown. prong on ventral surface. REVISIONS: There are currently four includ- REVISIONS: There is currently only one ed species, Xenochlora chalkeos Engel et al., species recognized in the genus. X. ianthina (Smith), X. nigrofemorata BIOLOGY: Nothing is known of Thecto- (Smith), and X. ochrosterna Engel et al., all chlora biology, aside from its association identified by the key provided in Engel et al. with a specific genus of acarid mites — (1997). Thectochloracarus (Fain et al., 1999). The BIOLOGY: A nest of X. ianthina was re- modified anterior surface of T1 for the trans- ported by Bates in a rotten twig (as a species port of mites suggests that this is perhaps a of Megalopta; Smith, 1861). Unfortunately, mutualistic relationship. no further details were given. DISTRIBUTION: Thectochlora occurs in DISTRIBUTION: Xenochlora species are dis- northern Argentina, southern Brazil, Para- tributed in Amazonian Brazil, Ecuador, and guay, and Guyana. Peru.

Genus Xenochlora Engel, Brooks, and CORYNURINA, NEW SUBTRIBE Yanega TYPE GENUS: Corynura Spinola, 1851, Xenochlora Engel, Brooks, and Yanega, 1997: 7. present designation. Type species: Xenochlora ochrosterna Engel, Brooks, and Yanega, 1997, original designation. DIAGNOSIS: This is the most primitive group of Augochlorini and is restricted to DIAGNOSIS: This group is most similar to southern South America. The species are the nocturnal genus Megalopta and most spe- generally small, although females of at least cies were at one time included therein. Xen- one species, R. inflaticeps, can be moderately 52 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250 large. Most species are dull metallic (except soma petiolate. Apical margins of S4 and S5 Callistochlora) or nonmetallic in coloration. unmodified. Apical margin of S6 emarginate. The presence of a strong galeal comb, the Apical margin of S7 unmodified. Apical short galea that extends posteriorly only half- margin of S8 with median projection; spic- way to the stipital base, and the short median ulum broad. Proctiger unmodified. Gonobas- process on the premental apex all serve to al bridge broad; dorsal lobes weak. Basal separate this group from the larger and more process of gonostylus present; parapenial diverse Augochlorina. lobe absent; dorsal gonostylus reduced, pres- DESCRIPTION: Female. Epistomal sulcus ent as a ridge bearing setae. Ventral surface forming an obtuse or nearly linear angle. Pre- of penis valve with keel. mental apex with median process as long as or shorter than lateral processes. Strong ga- Subgenus Callistochlora Michener leal comb present (figs. 24, 25); galeal apex Figures 28, 61, 77 rounded (fig. 24); galeal base extending only halfway to stipital base (fig. 21). Marginal Callochlora Moure, 1964: 269. Type species: cell apex acute. Integument frequently dull Halictus chloris Spinola, 1851, original designation. Nomen praeoccupatum (nec Callochlo- metallic. Male. Distal process of labrum fre- ra Packard, 1864 [Lepidoptera: Bombycidae]). quently absent (except in Rhinocorynura). Callistochlora Michener, 1997: 12. Nomen novum Apical margins of S4ÐS5 unmodified. pro Callochlora Moure, 1964. Type species: autobasic with Callochlora Moure, 1964. Genus Corynura Spinola DIAGNOSIS: Refer to Diagnosis for Cory- DIAGNOSIS: Species of Corynura s.l. can be nura s.s. (below). distinguished from other Corynurina by the DESCRIPTION: As for the genus with the combination of a broad hypostomal fossa, a following additions: Female. Compound nonproduced pronotal lateral angle, and a tri- eyes with long eye hairs. Basitibial plate angular recess surrouding the propodeal pit. rimmed posteriorly, anterior border obsolete. DESCRIPTION: Female. Mandible with Inner hind tibial spur pectinate. Basal area of moderate to strong subapical tooth. Labral propodeum striate. Male. Basal process of distal process broadly triangular; basal ele- gonostylus without setae. vation transverse; teeth absent. Prementum REVISIONS: Moure and Hurd (1987) list not greatly elongate. Galeal apex rounded; four species included in Callistochlora (as galeal comb present; galeal base extending Callochlora); however, two of these have about half of distance to stipital base. Hy- since been moved to other genera (Engel, postomal fossa as long as wide or wider; hy- 1996c). These taxonomic changes have left postomal ridge carinate; anterior angle the subgenus with only the two originally in- rounded. Malar space length less than man- cluded species, C. chloris (Spinola) and C. dibular width. Epistomal sulcus obtuse. Ocel- prothysteres (Vachal), both of which can be li not greatly enlarged; ocellar furrow absent. identified by Moure’s key (1964). Vertex not expanded or ridged behind ocelli. BIOLOGY: The nesting biology and mature Preoccipital ridge rounded. Pronotal lateral larva of C. chloris were studied by Claude- angle not produced, obtuse; dorsal ridge car- Joseph (1926, as a species of Halictus). inate; lateral ridge rounded. Mesoscutal an- DISTRIBUTION: Callistochlora occurs in terior border rounded; mesoscutal lip round- Chile and Argentina, with one record of C. ed. Tegula oval. Anterior basitarsal brush prothysteres from Peru. present. Apex of marginal cell acute. Distal hamuli with irregular spacing pattern. Pro- Subgenus Corynura Spinola s.s. podeal pit set into notch. Male. Mandible Figures 14, 21, 24Ð26 simple. Labrum without distal process; basal Corynura Spinola, 1851: 296. Type species: Cor- are not notched. Antennae long, extending ynura gayi Spinola, 1851 [ϭ Halictus rubellus back to propodeal triangle; F2 much longer Haliday, 1836], designation of Alfken (1926). than F1, usually longer than or as long as Corynogaster Sichel, 1867: 146. Type species: scape. Inner hind tibial spur serrate. Meta- Corynura gayi Spinola, 1851 [ϭ Halictus ru- 2000 ENGEL: BEE TRIBE AUGOCHLORINI 53

bellus Haliday, 1836], designation of Daly et al. group in Augochlorini. Species are minute, (1987). Designation of Halictus (Corynura) dark, and dull metallic blue-green. gayi Spinola, 1851 [meaningless combination] DESCRIPTION: Female. Mandible with by Dalla Torre (1896) and Sandhouse (1943) is moderate subapical tooth. Labral distal pro- invalid. This name was interpreted as a possible lapsus calami for Corynura Spinola, 1851, by cess narrowly triangular; basal elevation or- Eickwort (1969a). bicular; lateral margins of distal process ser- Rhopalictus Sichel, 1867: 146. Type species: Cor- rated. Prementum not greatly elongate. Ga- ynura flavofasciata Spinola, 1851 [ϭ Halictus leal apex rounded; galeal comb present; ga- chilensis Spinola, 1851], designation of Alfken leal base extending about half of distance to (1926). stipital base. Hypostomal ridge carinate; an- DIAGNOSIS: The dull metallic coloration, terior angle rounded. Malar space less than strongly bordered basitibial plate, and gran- mandibular width. Epistomal sulcus obtuse. ular propodeum separates species of Cory- Ocelli not greatly enlarged; ocellar furrow nura s.s. from Callistochlora. The monophy- absent. Vertex not expanded or ridged behind ly of this subgenus, however, could not be ocelli. Preoccipital ridge rounded. Pronotal confirmed. Those species with a pectinate in- lateral angle not produced, obtuse; dorsal ner metatibial spur may be more closely re- ridge carinate; lateral ridge rounded. Meso- lated to Callistochlora than to other Cory- scutal anterior border rounded; mesoscutal nura proper. Until this can be confirmed I lip rounded. Tegula oval. Basitibial plate bor- have not attempted to alter the classification dered posteriorly, margin obsolescent ante- of Corynura in the broad sense. riorly. Inner hind tibial spur pectinate. Apex DESCRIPTION: As for the genus with the of marginal cell acute. Distal hamuli with ir- following additions: Female. Basitibial plate regular spacing pattern. Basal area of pro- bordered on all sides. Inner hind tibial spur podeum granular; propodeal pit set into serrate or pectinate. Basal area of propodeum notch. Male. Mandible simple. Labrum with- granular. Male. Basal process of gonostylus out distal process; basal area not notched. with setae. Antennae long, extending beyond propo- REVISIONS: Alfken (1926) described and deum; F2 much longer than F1, usually lon- provided a key to the species known to him, ger than or as long as scape. Inner hind tibial and later updated this work (Alfken, 1931). spur serrate. Metasoma elongate. Apical mar- Moure and Hurd (1987) list 18 species. gins of S4 and S5 unmodified. Apical margin BIOLOGY: The nesting biology and mature of S6 not emarginate. Apical margin of S7 larva of C. cristata (Smith) were studied by with median process. Apical margin of S8 Claude-Joseph (1926, as a species of Halic- with median process; spiculum broad. Proc- tus). tiger unmodified. Gonobasal bridge broad; DISTRIBUTION: Species occur in Argentina and Chile. dorsal lobes weak. Basal process of gonostylus and parapenial lobe absent; dorsal process reduced to a ridge bearing setae. Ventral Genus Halictillus Moure surface of penis valve unmodified; dorsal Halictillus Moure, 1947: 7. Type species: Chlor- bridge near apices. alictus loureiroi Moure, 1941, monobasic and REVISIONS: There has been no attempt to original designation. revise the species of Halictillus. At present DIAGNOSIS: Species of Halictillus are the only two species, H. glabrescens (Cockerell) most Halictini-like among all augochlorines and H. loureiroi (Moure), are included in the and strongly resemble the genus Dialictus (in genus. fact, more often than not, they are misiden- BIOLOGY: The biology of H. glabrescens tified as species of Dialictus). The strong dis- was investigated by Claude-Joseph (1926, tal wing venation, medioapical cleft in the under the name Halictus glabriventris female T5, the absence of a pygidial plate in Friese). males, and the presence of a spiculum on S8 DISTRIBUTION: Halictillus occurs in north- in males all attest to the placement of this ern Argentina, southern Brazil, and Chile. 54 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

Genus Rhectomia Moure cies and presented a key for their identification. Rhectomia Moure, 1947: 9. Type species: Rhec- BIOLOGY: Nothing is known of Rhectomia tomia pumilla Moure, 1947, monobasic and biology, although the head size polymor- original designation. Corynurella Eickwort, 1969a: 398. Type species: phism present in at least one species (R. har- Corynurella mourei Eickwort, 1969a, monoba- risoni Engel) suggests the possibility of so- sic and original designation. cial behavior. DISTRIBUTION: Species of Rhectomia occur DIAGNOSIS: Rhectomia is most closely re- in northern Argentina, southern Brazil, Bo- lated to Rhinocorynura. The genus differs livia, Paraguay, and southern Peru. from Rhinocorynura by a broadly rounded mesoscutal anterior margin, a rounded me- Genus Rhinocorynura Schrottky soscutal lip, and the absence of clypeal ar- Figures 2, 18, 31, 33, 35, 74Ð75 mature and the anterior basitarsal brush. Corynura (Corynuropsis) Cockerell, 1901: 220. DESCRIPTION: Female. Mandible with Type species: Corynura (Corynuropsis) darwini weak subapical tooth. Labral distal process Cockerell, 1901 [ϭ Augochlora briseis Smith, narrowly triangular; basal elevation bilobed 1879], original designation. Nomen praeoccu- or orbicular; teeth absent. Prementum not patum (nec Corynuropsis Scott, 1894 [Crusta- greatly elongate. Galeal apex rounded; galeal cea: Entomostraca]). comb present; galeal base extending only Rhinocorynura Schrottky, 1909a: 147. Type spe- halfway to stipital base. Hypostomal ridge cies: Halictus (Corynura vel Corynuropsis) in- carinate; anterior angle rounded. Length of flaticeps Ducke, 1907, monobasic and original malar space less than basal mandibular designation. Ctenocorynura Schrottky, 1914: 628. Type spe- width. Epistomal sulcus obtuse. Ocelli not cies: Ctenocorynura vernoniae Schrottky, 1914 greatly enlarged; ocellar furrow absent. Ver- [ϭ Halictus (Corynura vel Corynuropsis) infla- tex expanded behind ocelli. Preoccipital ticeps Ducke, 1907], monobasic and original ridge rounded. Pronotal lateral angle pro- designation. duced, orthogonal to obtuse; dorsal ridge car- Corynuroides Sandhouse, 1943: 540. Nomen no- inate, usually with medial interruption sepa- vum pro Corynuropsis Cockerell, 1901. Type rating dorsal ridge into anterior and posterior species: autobasic with Corynuropsis Cockerell, halves on different planes; lateral ridge an- 1901. gled or carinate. Mesoscutal anterior border Rhynocorynura Sakagami and Moure, 1965: 303. rounded; mesoscutal lip rounded. Tegula Nomen emendatum (unjustified). oval. Anterior basitarsal brush absent. Basi- DIAGNOSIS: Refer to Diagnosis for Rhec- tibial plate bordered on all sides. Inner hind tomia. tibial spur pectinate. Apex of marginal cell DESCRIPTION: Female. Mandible with acute. Distal hamuli with irregular spacing strong subapical tooth. Labral distal process pattern. Basal area of propodeum smooth or broadly triangular; basal elevation bilobed; very finely striate; propodeal pit narrow. teeth absent. Prementum not greatly elon- Male. Mandible simple. Labrum without dis- gate. Galeal apex rounded; galeal comb pres- tal process; basal area not notched. Antenna ent; galeal base extending only halfway to extending back to scutellum; F2 roughly stipital base. Hypostomal ridge carinate or equal to F1. Inner hind tibial spur serrate. weakly lamellate; anterior angle rounded. Metasoma elongate. S4 unmodified. Apical Length of malar space less than basal man- margins of S5 and S6 emarginate. Apical dibular width. Clypeus frequently with ar- margin of S7 unmodified. Apical margin of mature of some design. Epistomal sulcus ob- S8 unmodified; spiculum narrow. Proctiger tuse. Ocelli not greatly enlarged; ocellar fur- with microtrichial fringe. Gonobasal bridge row absent. Vertex expanded behind ocelli. broad; dorsal lobes strong. Basal process of Preoccipital ridge rounded (except one spe- gonostylus present; parapenial lobe absent. cies is carinate). Pronotal lateral angle pro- Ventral surface of penis valve unmodified. duced, orthogonal to obtuse; dorsal ridge la- REVISIONS: The genus was recently treated mellate; lateral ridge angled. Mesoscutal an- by Engel (1995b) who recognized four spe- terior border narrowed; mesoscutal lip la- 2000 ENGEL: BEE TRIBE AUGOCHLORINI 55 mellate. Tegula oval. Anterior basitarsal posteriorly to posterior border of mesoso- brush present. Basitibial plate bordered on all ma or beyond ...... Ariphanarthra sides. Inner hind tibial spur pectinate. Apex 3. Pronotal dorsal surface not inflated; epistomal of marginal cell acute. Distal hamuli with ir- sulcus variable, but never forming a deep regular spacing pattern. Basal area of pro- projection extending into clypeus; basal area of propodeum variable ...... 4 podeum smooth. Male. Mandible simple. La- — Pronotal dorsal surface inflated; epistomal brum with distal process; basal area not sulcus forming a narrow, deep projection notched. Antenna extending back to scutel- extending into clypeus, nearly reaching lum; F2 approximately equal in length to F1. clypeal apex; basal area of propodeum Inner hind tibial spur serrate. Metasoma elongate, as long as scutellum and metan- elongate. Apical margins of S4 and S5 un- otum combined (fig. 39) ...... modified. Apical margin of S6 emarginate...... Chlerogelloides Apical margin of S7 with median projection. 4. Ocelli greatly enlarged (fig. 6); inner hind tib- Apical margin of S8 unmodified; spiculum ial spur serrate (fig. 48) ...... 5 narrow. Proctiger with microtrichial fringe. — Ocelli not enlarged (figs. 1Ð5); inner hind tibial spur variable ...... 6 Gonobasal bridge broad; dorsal lobes weak. 5. Compound eyes greatly enlarged, projecting Basal process of gonostylus present; para- above vertex in frontal aspect; clypeal apex penial lobe present or absent. Ventral surface relatively straight; marginal cell apex fee- of penis bly truncate and appendiculate ...... valve with keel...... Megaloptidia REVISIONS: There has been no revision of — Compound eyes not greatly enlarged, vertex the genus, which currently includes five de- projecting above upper tangent of com- scribed species. pound eyes; clypeal apex concave (figs. 5, 12); marginal cell apex acute ...... BIOLOGY: The nesting biology of one species, R. inflaticeps (Ducke), was investigated ...... Megommation (Megommation) 6. Basitibial plate with anterior border obsolete by Eickwort and Sakagami (1979). or all borders faint; inner hind tibial spur DISTRIBUTION: Species of Rhinocorynura variable (Megommation s.l. in part) . . . 7 occur in northern Argentina, southern Brazil, — Basitibial plate with all borders well devel- Bolivia, southern Ecuador, Paraguay, and oped; inner hind tibial spur serrate (fig. 48) Peru...... Micrommation 7. Scopa present (fig. 41); mandible normal, KEY TO GENERA AND SUBGENERA short with weak subapical tooth; inner hind OF AUGOCHLORINI tibial spur pectinate (figs. 46, 47) ..... 8 — Scopa absent (fig. 40); mandible long and The groups Electraugochlora, Glyptochlo- slender; inner hind tibial spur serrate (fig. ra, Oligochlora, Soliapis, and Xenochlora 48) .... Megommation (Cleptommation) 8. Basitibial plate short and rounded, borders are excluded from the male portion of this not defined ...... identification key as males are unknown for ...... Megommation (Megaloptina) these taxa. The key presented in Engel — Basitibial plate of normal length, narrowly (1998) mistakenly treated the marginal cell rounded, posterior border well developed apex of Oligochlora as acute while in all spe- ...... Megommation (Stilbochlora) cies the apex is, in truth, feebly truncate. 9. T5 with medioapical cleft (fig. 59); body sculpturing variable; mandible variably constructed; scopa frequently present; api- FEMALES cal margins of T1 and T2 not strongly de- 1. Prementum greatly narrowed and elongate, pressed or rimmed ...... 10 length seven or more times greater than — T5 lacking medioapical cleft; body coarsely width (fig. 13) ...... 2 punctured (figs. 7Ð9, 30, 56); mandible — Prementum not greatly narrowed or elongate, short, pointed, without subapical tooth; length less than seven times width (fig. 14) scopa absent (fig. 40); apical margins of T1 ...... 9 and T2 strongly depressed and rimmed 2. Maxillary palpus not elongate, extending pos- (figs. 56Ð57) ...... Temnosoma teriorly to base of prementum at most . . 3 10. Malar space elongate, as long as or longer — Maxillary palpi greatly elongate, extending than basal mandibular width (fig. 4) . . 11 56 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

— Malar space not elongate, length less than supplementary teeth; vertex without inter- basal mandibular width (figs. 1Ð3, 5, 8) . . ocellar furrow ...... 20 ...... 13 18. Ocelli enlarged, nearly touching compound 11. Flagellum with the normal complement of 10 eye (fig. 6); distal hamuli of hind wing flagellomeres; pronotal dorsal surface fre- closely packed and numerous (fig. 36); hairs quently inflated; basal area of propodeum of mesotibia, metatibia, and metabasitarsus as long as or longer than scutellum and me- variably colored, frequently amber, never tanotum combined (fig. 39), nonstriate . . . stiff and black (Megalopta s.l.) ...... 19 ...... 12 — Ocelli normal, not enlarged, more than one — Flagellum with only nine flagellomeres; pron- ocellar diameter from compound eye (figs. otal dorsal surface not inflated; basal area 1Ð5); distal hamuli of hind wing normal, of propodeum subequal to scutellum, short- with distinct spacing pattern (fig. 37); hairs er than scutellum and metanotum combined of mesotibia, metatibia, and metabasitarsus (fig. 38), frequently striate .... Chlerogas stiff and black ...... Xenochlora 12. Inner hind tibial spur densely pectinate, with 19. Scopa absent (fig. 40); mandible long and more than 10 long teeth; epistomal sulcus slender, without supplementary teeth; bas- forming orthogonal angle; pronotal dorsal itibial plate absent (fig.45)...... surface not inflated (Ischnomelissa in part) ...... Megalopta (Noctoraptor) ...... Ischnomelissa — Scopa present (fig. 41); mandible normal, — Inner hind tibial spur pectinate, fewer than 10 short with supplementary teeth on inner long teeth; epistomal sulcus forming acute margin (fig. 16); basitibial plate present, angle; pronotal dorsal surface frequently anterior border obsolescent (fig.44)..... inflated ...... Chlerogella ...... Megalopta (Megalopta) 13. Inner hind tibial spur pectinate (figs. 46, 47) 20. Pronotal dorsal ridge lamellate (fig.33)...... 14 ...... 21 — Inner hind tibial spur serrate (fig. 48) . . 36 — Pronotal dorsal ridge variable, but never la- 14. Inner posterior angle of tegula notched (fig. mellate ...... 22 34); pronotal dorsal ridge lamellate (fig. 21. Anterior surface of T1 modified into an acarinarium (basal glabrous area surrounded by 33); labral distal process quadrate, expand- dense hairs) (figs. 53Ð55); mesotrochanter ed at apex and base (fig. 20) (Augochlo- with dorsal hook (fig. 42); vertex short; ropsis s.l.)...... 15 marginal cell apex truncate; basitibial plate — Inner posterior angle of tegula rounded, not obsolescent on anterior border (fig. 44); notched (fig. 35); pronotal dorsal ridge var- basal area of propodeum covered with to- iable; labral distal process variable, but nev- mentum ...... Thectochlora er expanded at apex (figs. 17Ð19) .... 17 — Anterior surface of T1 unmodified (fig. 52); 15. Mesoscutal anterior border rounded (fig. 32); mesotrochanter lacking a dorsal hook; ver- mesoscutal lip rounded; vertex normal, lon- tex lengthened posteriorly; marginal cell ger than a single ocellar diameter; body apex acute; basitibial plate with strong bor- variously sculptured ...... 16 ders (fig. 43); basal area of propodeum var- — Mesoscutal anterior border narrow (fig. 33); iable, but never covered with tomentum . . mesoscutal lip lamellate; vertex short, bare- ...... Rhinocorynura ly a single ocellar diameter in length; body 22. Preoccipital ridge carinate (fig. 27); pronotal coarsely punctured ...... dorsal ridge carinate ...... 23 ...... Augochloropsis (Glyptochlora) — Preoccipital ridge rounded or angled, but dis- 16. Basal area of propodeum striate or pitted . . tinctly not carinate (fig. 26); pronotal dorsal ...... Augochloropsis (Augochloropsis) ridge variable ...... 27 — Basal area of propodeum smooth, granular, or 23. Basitibial plate with obsolete anterior border rugulose ...... (fig. 44); epistomal sulcus forming orthog- .....Augochloropsis (Paraugochloropsis) onal angle; mesoscutal anterior border 17. Labral keel of distal process with basal ex- broadly rounded (fig.32) ...... 24 pansion (fig. 19); if scopa present, then — Basitibial plate with well-developed borders mandible with supplementary teeth on in- (fig. 43); epistomal sulcus forming orthog- ner margin (fig. 16); vertex with interocel- onal or obtuse angle; mesoscutal anterior lar furrow (deep furrow between lateral border frequently narrowed (fig. 33) . . 25 ocelli, fig.11) ...... 18 24. Inner hind tibial spur pectinate, teeth well — Labral keel of distal process without basal ex- spaced, with fewer than 10 long teeth (figs. pansion (figs. 17, 18); mandible without 46, 47); pronotal dorsal ridge not produced 2000 ENGEL: BEE TRIBE AUGOCHLORINI 57

anteriorly; basal area of propodeum gran- ders; basal area of propodeum striate .... ular with fine basal striations ...... Neocorynurella ...... Andinaugochlora 31. Basitibial plate with all borders strongly de- — Inner hind tibial spur densely pectinate, with fined (fig. 43); anterior surface of T1 un- more than 10 long teeth; pronotal dorsal modified (fig.52)...... ridge produced anteriorly; basal area of pro- ...... Oligochlora (Soliapis) podeum with strong striae reaching to apex — Basitibial plate with anterior border obsolete ..... Caenaugochlora (Ctenaugochlora) (fig. 44); anterior surface of T1 with weak- 25. Angle of epistomal sulcus forming obtuse an- ly developed acarinarium (cf. fig.53).... gle; mesoscutal anterior border sometimes ...... Oligochlora (Oligochlora) narrowed (fig. 33); hairs of compound eyes 32. Basitibial plate with well-developed borders minute, shorter than, or scarcely longer (fig.43) ...... 33 than diameter of a single ommatidium (fig. — Basitibial plate with obsolete anterior border 29) ...... Neocorynura (fig.44) ...... 35 — Angle of epistomal sulcus forming orthogonal 33. Pronotal dorsal ridge not produced; anterior angle; mesoscutal anterior border broadly basitarsal brush present ...... 34 rounded (fig. 32); hairs of compound eyes — Pronotal dorsal ridge produced, frequently frequently longer than three or more om- with median interruption setting anterior maditial diameters (fig.28)...... and posterior halves on separate planes, ...... Caenaugochlora (Caenaugochlora) frequently with a small flange; anterior bas- 26. Preoccipital ridge strongly carinate; mesos- itarsal brush absent ...... Rhectomia cutul anterior border frequently narrowed 34. Basal area of propodeum with striae along bas- (fig. 33); basal labral elevation transverse al margin; labral distal process narrowly tri- ...... Neocorynura angular (fig. 17); galeal comb absent (figs. — Preoccipital ridge weakly carinate; mesoscu- 22, 23); propodeal pit not enclosed within a tal anterior border broadly rounded; basal V-shaped notch ...... Augochlorodes labral elevation orbicular ...... — Basal area of propodeum lacking any striae, ...... Neocorynurella part entirely granular or smooth; labral distal process broadly triangular (fig. 18); galeal 27. Inner hind tibial spur pectinate, teeth spaced comb present (figs. 24, 25); propodeal pit apart, with fewer than 10 long teeth (figs. enclosed within a V-shaped notch ..... 46, 47); basal area of propodeum variable, ...... Corynura (Corynura) part but shorter than scutellum and metanotum 35. Basal area of propodeum granular; hairs of combined (fig.38)...... 28 compound eye short (fig. 29); labral distal — Inner hind tibial spur densely pectinate, with process narrowly triangular (fig. 17); dull more than 10 long teeth, closely packed; and dark metallic blue-green or black . . . basal area of propodeum as long as or fre- ...... Halictillus quently longer than scutellum and metan- — Basal area of propodeum striate; hairs of otum combined (fig.39) ...... compound eye long (fig. 28); labral distal ...... Ischnomelissa part process broadly triangular (fig. 18); bril- 28. Marginal cell apex feebly truncate and appen- liant metallic green ...... diculate ...... 29 ...... Corynura (Callistochlora) — Marginal cell apex acute ...... 32 36. Epistomal sulcus forming obtuse angle . . . 37 29. Vertex without ridge posterior to ocelli; ep- — Epistomal sulcus forming orthogonal to acute istomal sulcus obtuse; galeal apex rounded angle ...... 39 (fig.24) ...... 30 37. Marginal cell apex acute; mesoscutal anterior — Vertex with transverse ridge posterior to ocel- border broadly rounded (fig. 32); preoccip- li (fig. 10); epistomal sulcus orthogonal; ital ridge variable ...... 38 galeal apex pointed (fig.23) ...... — Marginal cell apex truncate; mesoscutal an- ...... Pseudaugochlora terior border narrowed (fig. 33); preoccip- 30. Pronotal dorsal ridge carinate; labral teeth ital ridge carinate (fig.27) ...... strong; basitibial plate variable; basal area ...... Paroxystoglossa of propodeum granular or smooth, fre- 38. Preoccipital ridge rounded (fig. 26); basitibial quently without striae or with weak basal plate with well-developed borders (fig. 43); striae (fossil genus from Dominican amber: basal area of propodeum granular or Oligochlora s.l.) ...... 31 smooth, but never striate; galeal comb pre- — Pronotal dorsal ridge rounded; labral teeth absent (figs.24,25) ...... sent; basitibial plate with well-defined bor- ...... Corynura (Corynura) part 58 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

— Preoccipital ridge carinate (fig. 27); basitibial apex; mandible thin and twisted towards plate with obsolete anterior border (fig. 44); apex ...... Chlerogelloides basal area of propodeum striate; galeal 4. Ocelli greatly enlarged (fig.6)...... 5 comb absent (figs. 22, 23) . . Ceratalictus — Ocelli not enlarged (figs. 1Ð5)...... 6 39. Preocciptial ridge rounded (fig.26) .... 40 5. Marginal cell apex feebly truncate; S4 emar- — Preoccipital ridge carinate (fig.27) .... 41 ginate ...... Megaloptidia 40. Epistomal sulcus forming acute angle, very — Marginal cell apex acute; S4 not emarginate slightly protruding into clypeus (figs. 80, ...... Megommation (Megommation) 81); mesoscutal anterior border rounded; 6. Propodeal spiracle surrounded by a dense apex of marginal cell feebly truncate and patch of plumose setae ...... 7 appendiculate (Dominican amber fossil: — Propodeal spiracle not surrounded by a dense Augochlora s.l. in part, figs. 79Ð81)..... patch of plumose setae ...... 8 ...... Augochlora (Electraugochlora) 7. Apical margins of S2ÐS3 unmodified; head — Epistomal suclus forming orthogonal angle; and mesosoma brilliant metallic blue-green mesoscutal anterior border slightly nar- ...... Megommation (Megaloptina) rowed; marginal cell apex acute ...... — Apical margins of S2ÐS3 with short medial ...... Megaloptilla projections; head and mesosoma mostly am- 41. Epistomal sulcus forming acute angle, forming ber or brown ...... lobe which protrudes into basal margin of ...... Megommation (Cleptommation) clypeus; marginal cell apex truncate and ap- 8. F1 longer than F2; labral distal process absent pendiculate (Augochlora s.l. in part) . . 42 ...... Micrommation — Epistomal sulcus forming orthogonal angle; — F2 longer than F1; labral distal process present marginal cell apex acute ...... 43 ...... Megommation (Stilbochlora) 42. Mandible with large, strongly developed sub- 9. Malar space elongate, as long as, or usually apical tooth; basal labral elevation orbicular; much longer than basal mandibular width S1 frequently with a median ridge or tuber- (fig.4) ...... 10 cle (fig.58) ...... — Malar space not elongate, shorter than basal ...... Augochlora (Augochlora) mandibular width (figs. 1Ð3,5,8) .... 12 — Mandible normal, subapical tooth weak; basal 10. Flagellum with only 10 flagellomeres; inner labral elevation transverse; S1 rarely with hind tibial spur pectinate (figs. 46, 47); pron- median ridge or tubercle ...... otal dorsal surface not inflated; basal area of ...... Augochlora (Oxystoglossella) propodeum approximately as long as scutel- 43. Inner hind tibial spur with an enlarged, broad, lum (fig.38)...... Chlerogas basal tooth (fig.49) ...... Pereirapis — Flagellum with the normal complement of 11 — Inner hind tibial spur normally serrated, with- flagellomeres; inner hind tibial spur serrate out expanded basal tooth (fig.48) ..... (fig. 48); pronotal dorsal surface usually in- ...... Augochlorella flated; basal area of propodeum elongated, as long as scutellum and metanotum combined (fig.39)...... 11 MALES 11. Dorsal gonostylar process reduced without se- 1. Prementum greatly narrowed and elongate, tae or with extremely short setae ...... length seven or more times longer than wide ...... Ischnomelissa part (fig.13) ...... 2 — Dorsal gonostylar process reduced with long — Prementum not narrowed or elongate, length setae, frequently surpassing ventral gonos- less than seven times longer than wide (fig. tylar process ...... Chlerogella 14) ...... 9 12. Pronotal dorsal ridge lamellate ...... 13 2. Maxillary palpus not greatly elongate .... 3 — Pronotal dorsal ridge variable, but never la- — Maxillary palpus greatly elongate, extending mellate ...... 16 posterad beyond apex of mesosoma .... 13. Inner posterior angle of tegula with notch (fig...... Ariphanarthra 34); preoccipital ridge carinate or sharply 3. Pronotal dorsal surface not inflated; epistomal angled; S4 with medioapical lobe, laterally sulcus variable, but never forming a deep with apical processes (Augochloropsis s.l.) projection extending into clypeus; mandible ...... 14 unmodified ...... 4 — Inner posterior angle of tegula rounded, with- — Pronotal dorsal surface inflated; epistomal sul- out notch (fig. 35); preoccipital ridge round- cus forming a thin, deep projection extended; S4 without medioapical lobe, laterally ing into clypeus, almost reaching clypeal with or without apical processes ..... 15 2000 ENGEL: BEE TRIBE AUGOCHLORINI 59

14. S5 with medioapical emargination ...... dense setae; compound eyes frequently ...... Augochloropsis (Augochloropsis) with long hairs (fig.28) ...... — S5 without apical emargination ...... Caenaugochlora (Caenaugochlora) .....Augochloropsis (Paraugochloropsis) — S4 without medioapical tubercles, however, 15. S5 with apical margin emarginate; vertex nor- with a triangular medioapical setal patch; mal, not swollen posteriorly; antennae hairs of compound eyes short (fig. 29) . . . reaching beyond propodeum; F2 much lon- ...... Caenaugochlora (Ctenaugochlora) ger than F1 (fig.61) ..... Thectochlora 25. S5 with apical margin strongly depressed or — S5 without apical emargination; vertex swol- emarginate ...... 26 len posteriorly; antennae reaching at most — S5 with apical margin unmodified ..... 29 to propodeum; F2 and F1 equal in length 26. F2 much longer than F1 (fig. 61); marginal (fig.62) ...... Rhinocorynura cell apex acute or truncated; pronotal lat- 16. S4 with apical margin strongly concave or eral angle not produced ...... 27 emarginate ...... 17 — Length of F2 roughly equal to F1 (fig. 62); — S4 with apical margin unmodified or weakly marginal cell apex acute; pronotal lateral depressed ...... 25 angle produced ...... Rhectomia 17. Ocelli greatly enlarged (fig. 6); S4 with later- 27. Antennae extending to posterior border of pro- oapical notches on apical margin (fig. 66); podeum; mesoscutal anterior border rounded epistomal sulcus forming slightly acute an- (fig. 32); metasoma elongate ...... 28 gle; distal hamuli of hind wing dense and — Antennae extending to scutellum; mesoscutal numerous (fig. 36) (Megalopta s.l.) . . 18 anterior border slightly narrowed; metaso- — Ocelli not enlarged; S4 without lateroapical ma oval ...... Megaloptilla notches; epistomal sulcus variable; distal 28. Epistomal sulcus forming orthogonal angle; hamuli exhibiting distinct spacing pattern, venter of penis valve with prong ..... not densely packed together in a series (fig...... Andinaugochlora 37) ...... 19 — Epistomal sulcus forming obtuse angle; ven- 18. F2 about equal in length to F3; clypeus with ter of penis valve without prong ...... apical margin and scape anteriorly white; ...... Neocorynurella antennal socket diameter much greater than 29. Marginal cell apex feebly truncate ..... 30 ocular-antennal socket distance; S5 narrow- — Marginal cell apex acute ...... 32 ly emarginate . . . Megalopta (Megalopta) 30. Epistomal sulcus forming acute angle, form- — F2 about two-thirds length of F3; clypeus and ing a lobe extending into clypeus; metaso- scape completely black; antennal socket di- ma oval (Augochlora s.l.) ...... 31 ameter less than ocular-antennal distance; — Epistomal sulcus forming obtuse angle; me- S5 broadly depressed ...... tasoma frequently petiolate ...... Megalopta (Noctoraptor) ...... Neocorynura part 19. Marginal cell apex acute ...... 20 31. Outer ridge bordering gonostylus with short — Marginal cell apex feebly truncate ..... 22 setae ...... Augochlora (Augochlora) 20. S5 with apical margin unmodified ..... 21 — Outer ridge bordering gonostylus with long — S5 with apical margin strongly depressed . . . setae, surpassing ventral gonostylus length ...... Paroxystoglossa part ...... Augochlora (Oxystoglossella) 21. S5 with dense medial patch of setae; S6 with 32. Body variously sculptured, but not coarsely weak medio-apical notch, apical margin not punctate; T1 and T2 not depressed on basal produced ...... Augochlorodes halves; T7 not bilobed along apical margin — S5 without medial patch of setae; S6 with ...... 33 strong medio-apical notch, slightly pro- — Body coarsely punctate (fig. 30); T1 and T2 duced along apical margin . . . Pereirapis depressed on basal half (figs. 56, 57); T7 22. F2 approximately equal to F1 (fig.62) .... with apical margin bilobed (fig.67)...... 23 ...... Temnosoma — F2 much longer than F1 (fig.61)...... 33. Epistomal sulcus forming orthogonal angle ...... Paroxystoglossa part ...... 34 23. Vertex unmodified; terminal flagellomere un- — Epistomal sulcus forming obtuse angle .... modified (Caenaugochlora s.l.) ...... 24 ...... 35 — Vertex behind ocelli produced into a trans- 34. Preoccipital ridge rounded (fig. 26); F2 much verse ridge (fig. 10); terminal flagellomere longer than F1; basal area of propodeum (F11) produced into a hook (fig.63) .... elongated, as long as or longer than scutel- ...... Pseudaugochlora lum and metanotum combined (fig. 39) . . 24. S4 with medioapical tubercles covered in ...... Ischnomelissa part 60 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

— Preoccipital ridge carinate (fig. 27); F1 longer — Body brilliant metallic green; hairs of com- than F2; basal area of propodeum subequal pound eyes long (fig. 28); basal process of to scutellum (fig.38)..... Augochlorella gonostylus without setae ...... 35. Preoccipital ridge rounded (fig.26) .... 36 ...... Corynura (Callistochlora) — Preoccipital ridge carinate (fig.27) .... 38 38. F2 much longer than F1 (fig. 61); mesoscutal 36. Metasoma petiolate; apical margin of S6 anterior border narrowed (fig. 33); antennae emarginate (Corynura s.l.) ...... 37 — Metasoma elongate, but distinctly not petio- frequently just surpassing posterior border late; apical margin of S6 not emarginate of propodeum ...... Neocorynura part ...... Halictillus — F2 roughly equal in length to F1 (fig. 62); 37. Body brown, black, or dully metallic in col- mesoscutal anterior border rounded (fig. oration; hairs of compound eye short (fig. 32); antennae reaching back to mesoscutal 29); basal process of gonostylus with setae posterior border or to scutellum ...... Corynura (Corynura) ...... Ceratalictus

CLADISTICS DESCRIPTION OF CHARACTERS sotrochanteral hook, arrangement of the hamuli, the interocellar furrow). In total, 46 Eickwort’s (1969a) comparative morpho- new characters are used; 34 from adult mor- logical study of the Augochlorini served as phology and 12 from behavior. the primary resource for characters used be- Homology was determined using the prin- low. His work was largely supplemented and ciple of connections as it has been classically modified by the my above study of the gen- defined (Geoffroy Saint-Hilaire, 1818; era. A number of characters used by Eick- Owen, 1866; Remane, 1952). Identification wort (op. cit.) in his groundplans had to be of ethological homology follows the same excluded as the states could not be confi- principles as used for morphology (Baerands, dently differentiated. Examination of taxa for 1958; Wenzel, 1992; Greene, 1994), namely each group revealed overlapping variation topographic and phenetic similarity coupled for many of the features included in his with functional information. study. For example, Eickwort (op. cit.), as The following 84 characters were coded well as a recent cladistic reinterpretation of for all 39 genera and subgenera of Augo- his raw data (Danforth and Eickwort, 1997), chlorini as well as 10 genera of Halictini, 1 utilized characters that were treated as ‘‘unmodified’’ versus ‘‘modified’’. Although genus of Nomioidini, and two genera of these statements are not incorrect, the ho- Nomiinae (summarized in the data matrix mology assessments and codings can be fur- presented in appendix 2). As many species ther refined. For example, the apical margin as possible were examined from each genus of the fourth male metasomal sternum is one in order to survey variation in each group; in in which ‘‘unmodified’’ refers to a straight, several cases all known species for a given uninterrupted apical margin, whereas ‘‘mod- genus were examined (appendix 3 presents a ified’’ includes those taxa possessing a deep list of taxa examined for construction of the median cleft, a gently depressed or concave data matrix). In the character matrix (appen- margin, a strong median process, or deep lat- dix 2), interrogative marks represent un- eral notches. Thus, the state ‘‘modified’’ con- known information while dollar signs repre- tains little or no true grouping information. sent subset polymorphisms (elaborated on Of the 80 characters employed in the earlier below). The matrix was constructed in the studies (Eickwort, 1969a; Danforth and Eick- DADA (Nixon, 1995). Characters 0 through wort, 1997), 27 are used here and an addi- 33 are for the female sex, although some may tional 11 are reinterpreted based on direct equally apply to males, while 34 through 60 study of numerous augochlorine species (ap- are derived from the male. Characters 61 pendix 3). At the same time my own studies through 71 are autapomorphic, used to sup- revealed a number of new characters which port the monophyly of various genera, and are employed here for the first time (e.g., me- not listed in sequence by gender. The last 12 2000 ENGEL: BEE TRIBE AUGOCHLORINI 61 characters (characters 72 through 83) are long as an ommatidium diameter (fig. 29); based on ethological traits. State number (1) long, much longer than several ommatid- does not imply plesiomorphic versus apo- ium diameters (fig. 28). morphic polarity. The nomiine genera Dieu- 13. Ocelli: (0) normal (fig. 1Ð5); (1) great- nomia and Lipotriches were used as out- ly enlarged (fig. 6). groups to polarize the characters. For the def- 14. Interocellar furrow: (0) absent; (1) inition of particular morphological structures present (fig. 11). refer to the above section on General Mor- 15. Preoccipital area: (0) not carinate (fig. phology. 26); (1) carinate (fig. 27) or lamellate. 16. Pronotal dorsal surface between lateral FEMALE-DERIVED CHARACTERS angles: (0) short, mostly covered by mesoscutum; (1) convex, inflated, and greatly ex- 0. Apical margin of clypeus: (0) straight; panded anteriorly, not obscured by meso- (1) concave (fig. 12). scutum. 1. Distal process of labrum: (0) base 17. Mesoscutal anterior border: (0) broad- broadly joined to basal area, triangular (fig. ly rounded (fig. 32); (1) strongly narrowed 18); (1) base narrowly joined to basal area, and projecting forward (fig. 33). triangular (fig. 19); (2) quadrate (fig. 20). 18. Tegula: (0) oval (fig. 35); (1) inner 2. Keel of labral distal process: (0) thin posterior margin notched (fig. 34). ridge (figs. 17, 18); (1) expanded at base (fig. 19. Propodeal triangle length: (0) no lon- 19). ger than metanotum; (1) subequal to scutel- 3. Lateral teeth of labrum: (0) absent (fig. lum (fig. 38); (2) as long as or longer than 18, 19); (1) small, forming a serrated margin; scutellum and metanotum combined (fig. (2) strong, forming a weakly pectinate mar- 39). gin (fig. 17). 20. Propodeal posterior pit: (0) set into tri- 4. Length of malar space: (0) short, less angular recess; (1) not set into a triangular than basal width of mandible (figs. 1Ð3); (1) recess. elongate, as long as, or longer than, basal 21. Probasitarsal brush: (0) absent; (1) width of mandible (fig. 4). present. 5. Angle formed between dorsal and lat- 22. Basitibial plate: (0) absent (fig. 45); (1) eral clypeogenal sulci and opening toward present, anterior border obsolete (fig. 44); (2) compound eye: (0) obtuse or linear; (1) approximately orthogonal; (2) acute, forming present, rimmed on all borders (fig. 43). lobe projecting into clypeal base. 23. Metatibial spine: (0) absent; (1) pres- 6. Apex of galea: (0) lobed (fig. 24); (1) ent. distinctly acute (fig. 23). 24. Inner metatibial spur: (0) serrate (fig. 7. Galeal comb: (0) absent; (1) present 48); (1) pectinate, fewer than 10 teeth well (figs. 24, 25). spaced (figs. 46Ð47); (2) pectinate, more than 8. Base of galea: (0) extending posterad to 10 teeth, densely packed. middle of stipes (fig. 21); (1) extending pos- 25. Scopa: (0) absent (fig. 40); (1) present terad or near to base of stipes (fig. 22). (fig. 41). 9. Medioapical process of prementum: (0) 26. Basal area of propodeum: (0) normal, short, extending apicad to lateral processes; strongly angled from posterior surface; (1) (1) elongated, extending apicad beyond lat- declivitous. eral processes. 27. Apex of marginal cell: (0) acute; (1) 10. Prementum: (0) unmodified, less than truncate, frequently feebly appendiculate. 7ϫ longer than wide (fig. 14); (1) greatly 28. Distal wing venation: (0) strong; (1) narrowed and elongate, 7ϫ or more longer weakened. than wide (fig. 13). 29. Distal hamuli: (0) irregularly spaced 11. Anterior angle of hypostomal ridge: (fig. 37); (1) uniformly spaced and numerous (0) rounded; (1) angled, coming to a distinct (fig. 36). point. 30. Acarinarium of anterior surface of T1: 12. Compound eye hairs: (0) minute, as (0) absent (fig. 52); (1) present (figs. 53Ð55). 62 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

31. Median cleft in apical margin of T5: situated between apodemes of penis valves (0) absent; (1) present (fig. 59). in dorsal view; (1) broad, situated at or lat- 32. Gradulus of T6: (0) absent; (1) present. eral to apodemes of penis valves in dorsal 33. Gradulus of S4: (0) absent; (1) present. view. 55. Inner apical corner of volsella: (0) var- MALE-DERIVED CHARACTERS iously produced; (1) modified into a thin hook. 34. Distal process of labrum: (0) absent; 56. Ventral surface of penis valve: (0) un- (1) present (fig. 60). modified; (1) with broad prong; (2) with 35. Relative lengths of F1 versus length of keel. F2: (0) F1 equal to F2 (fig. 62); (1) F1 short- 57. Parapenial lobes: (0) absent (fig. 74); er than F2 (fig. 61); (2) F1 longer than F2. (1) present (fig. 73). 36. Dense, plumose setae surrounding pro- 58. Basal process of gonostylus: (0) ab- podeal spiracle: (0) absent; (1) present. sent; (1) present, without setae (fig. 77); (2) 37. Pygidial plate on T7: (0) absent; (1) present, with setae (figs. 74, 75). present. 59. Ventral process of gonostylus: (0) sin- 38. Anal filaments on apical margin of gle projection; (1) divided projection. proctiger: (0) absent; (1) present. 60. Dorsal process of gonostylus: (0) ab- 39. Microtrichiae on apical margin of sent; (1) setose ridge; (2) present, partially proctiger: (0) absent; (1) present. membranous; (3) present, sclerotized. 40. Medio-apical process on S3: (0) absent; (1) present. 41. Gradulus of S4: (0) continuous; (1) AUTAPOMORPHIES medially interrupted. 61. The female maxillary palpus: (0) un- 42. Gradulus of S4: (0) separated from an- modified, extending posterad to premental tecosta; (1) meeting antecosta medially. base at most; (1) greatly elongate, extending 43. Medio-apical margin of S4: (0) contin- posterad to metasoma. [Ariphanarthra-1] uous, straight; (1) gently concave; (2) cleft; (3) posterior process. 62. Hypostomal ridge posteriorly extended 44. Lateral notches on apical margin of into a lamellate flange [female: Megomma- S4: (0) absent; (1) present (fig. 66). tion proper—state 1]: (0) absent; (1) present. 45. Triangular setal patch medially on S4: 63. Medio-dorsal hook of mesotrochanter (0) absent; (1) present (fig. 65). [female: Thectochlora—state 1]: (0) absent; 46. Medio-apical margin of S5: (0) contin- (1) present (fig. 42). uous, straight (fig. 64); (1) gently concave; 64. Depressed and rimmed apical margin (2) cleft. of T1-2 [female: Temnosoma—state 1]: (0) 47. Gradulus of S6: (0) continuous; (1) absent; (1) present (figs. 56, 57). medially interrupted. 65. Number of units comprising antennal 48. Medio-apical margin of S6: (0) contin- flagellum (Nr. female/ Nr. male) [both sexes: uous, straight; (1) cleft (fig. 64). Chlerogas—state 1]: (0) 10/11; (1) 9/10. 49. Apical margin of S7: (0) continuous, 66. Dorsally extended transverse ridge be- straight; (1) with median process (figs. 68, hind ocelli on vertex [both sexes: Pseudau- 69); (2) with bilobed process (fig. 70); (3) gochlora—state 1]: (0) absent; (1) present with lateral processes (fig. 71). (fig. 10). 50. Apical margin of S8: (0) continuous, 67. F11 [male: Pseudaugochlora—state straight; (1) with median process. 1]: (0) unmodified, gently conical at apex; 51. Spiculum of S8: (0) absent; (1) pres- (1) hooked at apex (fig. 63). ent, broad (fig. 68); (2) present, narrow (fig. 68. Mesofemoral and mesotibial expansion 69). [male: Chlerogelloides—state 1]: (0) absent; 52. Point of fusion between S7 and S8: (0) (1) present. at apodeme apex; (1) before apodeme apex. 69. Inner hind tibial spur [male: Chlero- 53. Gonobase: (0) broad; (1) narrow. gas—state 1]: (0) serrate; (1) pectinate. 54. Gonobasal dorsal lobes: (0) narrow, 70. Broad medio-apical cleft on T7 [male: 2000 ENGEL: BEE TRIBE AUGOCHLORINI 63

Temnosoma—state 1]: (0) absent; (1) present (appendix 2). Subset polymorphisms (Nixon, (fig. 67). 1993) were coded for those taxa exhibiting 71. Long, anteriorly directed, dorsal pro- only two states of a multistate character. Sub- cess on penis valve [male: Temnosoma— set polymorphisms are indicated with the state 1]: (0) absent; (1) present (fig. 76). dollar sign ($) in the matrix and implemented with the Shift-$ command in DADA (Nixon, ETHOLOGICAL CHARACTERS 1995). The subgenus Augochlora was coded Behavioral information, mostly from nest as having subset polymorphism for the shape architecture, was derived from a number of of the labral distal process (character 1: states sources. Appendix 4 provides information as 0 and 1). Similarly, all nonparasitic taxa for to where pertinent ethological data were ob- which the social biology is unknown were tained for each genus. coded as subset polymorphisms for social be- 72. Social structure: (0) solitary; (1) clep- havior (character 72), thereby excluding the toparasitic; (2) communal; (3) semisocial; (4) parasitic state (the presence of a well-devel- eusocial. The classification of social catego- oped scopa, normal mandibles, and a basiti- ries as defined by Michener (1969a, 1974) is bial plate indicates that these taxa are not followed here. parasitic). This coding allowed these taxa to 73. Augochlorines are oligolectic, so spe- have any of the possible social states (soli- cific foraging preferences are not meaningful tary through eusocial; states 0, 2Ð4), but not for phylogenetic analysis. However, augo- to have the cleptoparasitic state (state 1). chlorine genera do vary in some aspects of their foraging behavior, most notably in the time of activity: diurnal (0); nocturnal (1). DATA ANALYSIS 74. Attachment of brood cell to burrow: All characters were run with equal weights (0) via long lateral; (1) sessile. and considered nonadditive. Analyses were 75. Brood cell distribution: (0) scattered; carried out using NONA (Goloboff, 1993) (1) clustered; (2) serial. and were run in two parts. The first analysis 76. Chamber containing cell cluster: (0) absent; (1) present, along main burrow; (2) consisted of 20 replications of random-taxon- present, offset from main burrow. addition with each replicate holding 500 77. Brood cell orientation: (0) horizontal trees in memory. This analysis was per- or slanting; (1) vertical. formed to identify a set of trees upon which 78. Nest plan: (0) branching before cells; more exhaustive branch-swapping could be (1) unbranched or branching only below first undertaken. The memory was then reallocat- set of cells. ed to hold as many topologies as possible 79. Cell location: (0) along a single bur- and the initial set of trees was submitted to row; (1) along multiple burrows. the program and analyzed using the max* 80. Short lateral tunnels bordering cells: command. Autapomorphic traits (characters (0) absent; (1) present. 61 through 71) were excluded from the cal- 81. Nesting substrate: (0) soil; (1) wood. culation of the consistency index (CI), reten- 82. Turret: (0) absent; (1) present. tion index (RI), and tree length. The two 83. Position of the pollen mass provisions nomiine genera (Dieunomia and Lipotriches) within the brood cell: pollen mass placed at were used to root the tree, thereby allowing cell apex (0); pollen mass occupying a lateral for the various halictine genera to potentially position within the cell (1). render the Augochlorini as nonmonophyletic. Trees were visualized and printed using SUBSET POLYMORPHISMS CLADOS (Nixon, 1993). 5488 trees exhib- Taxa which are polymorphic were coded ited length 283, CI 0.31, and RI 0.61 (strict with an interrogative mark in the data matrix consensus in fig. 78). 64 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

Fig. 78. Phylogeny of tribe Augochlorini (length 283, CI 0.31, RI 0.61, strict consensus). 2000 ENGEL: BEE TRIBE AUGOCHLORINI 65

DISCUSSION PHYLOGENETIC RELATIONSHIPS tribe, only the medio-apical cleft of the female fifth tergum and the pygidial plate of The phylogenetic analysis of augochlorine males were included. An approximate outline genera presented here is certainly not robust of augochlorine classification is presented in owing to the large number of trees, the low table 3. CI, and low RI; however, a strict consensus Because of the small sample of halictine of these topologies retains a remarkable genera examined, it was not possible to de- amount of resolution (fig. 78). On account of termine the sister-group of the Augochlorini, the high level of homoplasy in the analysis, but members of the caenohalictine genera I have conservatively avoided naming most (Caenohalictus, Habralictus, etc.) seem to be of the nodes in the cladogram. However, cer- the most likely candidates and should per- tain clades are well supported and are dis- haps be treated as a tribe separate from the cussed at length below. In the following dis- Halictini. In fact, a family-group name for cussion character and state numbers are pro- these genera was already proposed by Mich- vided in parentheses. ener (1954b) and could be reinstated to ac- commodate them as the tribe Caenohalictini. TRIBE AUGOCHLORINI A tribal classification of this subfamily rec- The Augochlorini is supported as mono- ognizing five monophyletic tribes seems phyletic among the Halictinae with the tribe more appropriate than the one in present use. Halictini apparently paraphyletic with re- The paraphyletic Halictini should be broken spect to the augochlorines. This conclusion into three entities: the Gastrohalictini (ϭ La- is not far removed from the one reached by sioglossum group), the Halictini (ϭ Halictus Pesenko (1983), although he considered the and Sphecodes groups), and the Caenohalic- Halictini to be monophyletic and his putative tini (ϭ Agapostemon and Caenohalictus synapomorphies of the Augochlorini are not groups) (table 4). all identical with those resulting from my studies (Engel, 1996a, 1998, herein). The SUBTRIBE CORYNURINA monophyly of the Augochlorini is supported by the medio-apical cleft of the fifth tergum The subtribes Augochlorina (discussed be- of females (31-1), absence of a pygidial plate low) and Corynurina are both recognized as in males (37-0), the medio-apical cleft of the monophyletic. The corynurine genera form sixth sternum in males (48-1), presence of a the basalmost clade of the tribe (fig. 78), spiculum (51-1,2), and sessile attachment of united by the presence of a strong glaeal brood cells to the main burrow in nests (74- comb (7-1) and the absence of a distal pro- 1). Additional possible synapomorphies, cess on the male labrum (34-0). All of these which are subsequently modified several genera are distributed in northern Chile and times within the tribe, are an unmodified Argentina, Paraguay, Bolivia, and the south- venter of the penis valve (56-0), the cluster- ernmost regions of Brazil and Peru. Moure ing of brood cells in the nest (75-1), and the and Hurd (1987) assigned the Ecuadorian presence of an excavated brood cell chamber species Halictus joannisi Vachal to Corynura (76-1). Although recent studies focusing on thereby extending the range of this genus the phylogeny of short-tongued bee families much further northward; however, this spe- (Alexander and Michener, 1995) failed to cies actually belongs to the high-altitude ge- support augochlorine monophyly, these anal- nus Andinaugochlora and is a member of the yses suffered from sparse taxon sampling at Augochlorina (Engel, 1996c). The genera lower ranks and therefore limited character comprising the Corynurina had been consid- choice. In the study of Alexander and Mich- ered paraphyletic with respect to the remain- ener (op. cit.) the augochlorines were repre- der of the tribe (Eickwort, 1969a; Danforth sented only by the genera Augochlora, Cor- and Eickwort, 1997) and this is somewhat ynura, and Pseudaugochlora, while of the supported by the absence of a cleft in the aforementioned synapomorphies for the male S6 of Halictillus, a character that could 66 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

potentially group the remaining corynurine presented by Alexander and Michener, 1995) genera with the Augochlorina. The paraphyly the a priori assumption that it is primitively of the subtribe, however, was primarily based present in the Augochlorini is not supported on the a priori assumption that the galeal by outgroup comparisons. A galeal comb is comb was a primitive trait and its loss con- not present among the Halictini and Nom- stituted a synapomorphy for the higher Au- ioidini and must therefore be considered in- gochlorini (here equivalent to the subtribe dependently derived in the common ancestor Augochlorina). Although this character is of these genera and not as a trait carried over certainly primitively present among many from a more distant ancestor with the Nom- higher-level bee groups (e.g., see distribution iinae or Rophitinae. The interpretation of the 2000 ENGEL: BEE TRIBE AUGOCHLORINI 67

posterad to meet the base of the stipes (8-1) as well as by the elongate median apical process of the prementum (9-1). Unlike the Cor- ynurina, the Augochlorina is widely distributed covering the range of the corynurine genera and extending northward to southern Canada as well as east into the West Indies. The subtribe can be further divided into two primary clades; the Megaloptomorpha and the Augochloromorpha (discussed below). The enigmatic genus Neocorynura is left un- assigned in the Augochlorina. A revision of the numerous species in Neocorynura is des- perately needed in order to conﬁrm its monophyly and to, it is hoped, conﬁdently place the genus among the Augochlorina.

MEGALOPTOMORPHA galeal comb as plesiomorphic for Augochlo- rini is also incongruent with the plesiom- The megaloptomorph augochlorines are orphic presence of a strongly distad galea rel- weakly united by the presence of a feebly ative to the stipes and the numerous charac- truncate marginal cell apex (27-1). This char- ters uniting Halictillus with Corynura (see acter, however, is homoplastic throughout the discussion below) if the Corynurina is con- tribe and is reversed in some species of the sidered paraphyletic. The analysis presented Ischnomelissiti and Megaloptiti clades (see here supports Corynurina monophyly and the below); it also appears independently in scat- absence of a cleft in the sixth sternum of tered species of the Augochloromorpha. Halictillus males is interpreteted as a second- Thus, no single character change serves to ary reversal. identify a megaloptomorph bee, instead the Within the Corynurina two main clades apomorphic presence of a truncate marginal are recognized (fig. 78: table 3); the Cory- cell apex and the absence of augochloro- nura group and the Rhinocorynura group. moph synapomorphies are together required The genera of the Corynura group share the to recognize this clade. Although megalop- presence of a triangular recess surrounding tomoph monophyly is weakly supported, the propodeal pit (20-0), an obsolescent an- they can be confidently excluded from the terior border to the basitibial plate (22-1), the Augochloromorpha and specific clades with- presence of a metatibial spine (23-1), a broad in the group can be confidently diagnosed spiculum (51-1), and the reduction of the (see below). dorsal gonostylar process to a setose ridge (60-1). In Corynura s.s. the basitibial plate ISCHNOMELISSITI structure is reversed to regain the anterior The Ischnomelissiti is a distinctive clade border. The Rhinocorynura group is united of genera that frequently have elongate by the combination of the male F1 length heads, perhaps an adaptation for visiting being equivalent to that of F2 (35-0), the flowers with deep corollas. The Ischnomelis- presence of microtrichiae on the proctiger siti are united by the elongate propodeum (39-1), a medially interrupted gradulus on the (19-2), the absence of distal process on the sixth sternum of males (47-1), and the pres- male labrum (34-0), and the reversal to an ence of a setose gonostylar basal process (58- acute marginal cell apex (27-0). The genera 2). Chlerogella and Chlerogelloides are further united by the presence of an acute epistomal SUBTRIBE AUGOCHLORINA sulcus (5-2), particularly so in Chlerogello- The subtribe Augochlorina is supported by ides (refer to description of this genus), and the elongation of the galea which extends an inflated pronotal dorsal surface (16-1). 68 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

The elongate head of Chlerogas is indepen- num in males (45-1), and the partially mem- dently derived, as this is a genus of the Au- branous dorsal gonostylar process (60-2). gochloromorpha (below). Augochloropsis s.l. monophyly is established by the quadrate distal labral process (1-2), THECTOCHLORITI ϩ MEGALOPTITI the tegular notch (18-1), the absence of a male labral process (34-0), the presence of a The two clades discussed below are united medioapical process on the fourth metasomal by the presence of a concave apical margin sternum of males (43-3), the broad spiculum on the fourth sternum (43-1) and the pres- (51-1), the inner apical hook of the volsella ence of a deep medioapical cleft on the fifth (55-1), the presence of a strong keel on the sternum of males (46-2). The structure of the venter of the penis valves (56-2), the vertical fourth sternum is further elaborated on in the orientation of brood cells in the nest (77-1), genus Megalopta (refer to description of ge- and the presence of short lateral tunnels bor- nus above). dering the cell clusters (80-1). The Megalopta group is a disinctive group THECTOCHLORITI of fairly robust bees and includes not only The modification of the anterior-facing the largest members of the tribe (i.e., some surface of the first metasomal tergum into an species of Megalopta s.s.) but also the most acarinarium is unique to the genera of Thec- diverse clade of nocturnal species within the tochloriti (30-1), although the charcter is re- tribe, inclusive of the only nocturnal parasitic versed in two apomorphic species of Oligo- bees of any family (Noctoraptor). The clade chlora (see discussion of the genus above). is supported by the broad distal labral pro- The acarinarium is described above under cess (1-0), the basal expansion of the distal General Morphology and is designed for the labral keel (2-1), the formation of an acute transport of phoretic instars of mites (see also angle by the epistomal sulcus that protrudes Fain et al., 1999). The modification of the into the basal margin of the clypeus (5-2), bee’s morphology for the transport of the the presence of the interocellar furrow (14- mites as well as the frequently species-spe- 1), the shortened and declivitous basal area cific association between the bees and mites of the propodeum (19-0 and 26-1), and the suggests that this relationship is a mutualistic utilization of a wood subtrate for nest con- one (at least within Augochlorini). No other struction (81-1). Augochlorini have acarinaria. AUGOCHLOROMORPHA MEGALOPTITI The unique character of broadly separated This clade is united by the fusion of the gonobasal lobes (54-1) unites this large clade gradulus with the antecosta on the fourth of genera. The genus Chlerogas is here con- sternum of males (42-1), although this is re- sidered the basalmost genus of the Augo- versed in Ctenaugochlora (this reversal be- chloromorpha based on the pectinate inner ing one of several synapomorphies for the metatibial spur. The serrate inner metatibial subgenus). spur unites the remaining genera, although it Among the Megaloptiti, the Augochlorop- is subsequently reversed in Pseudaugochlo- sis group is united by the apomorphic pres- ra, Stilbochlora, Megaloptina, and Augo- ence of a carinate preoccipital ridge (15-1), chlorodes. The next clade excludes Temno- presence of a gradulus on the fourth metasoma by the apomorphic presence of serrated somal sternum in females (33-1), and F1 and margins on the labral distal process (3-1). F2 being of equal lengths in males (35-0). This character is independently modified in Both of the included genera, Augochloropsis the genera Augochlora and Pereirapis and in and Caenaugochlora, are diverse. The mono- some species of the Megaloptidia group. phyly of Caenaugochlora s.l. is supported by Although the monophyly of the Paroxy- the apomorphic presence of an orthogonal stoglossa group was not supported by the epistomal sulcus (5-1), a triangular setal analysis presented here, the genera are diffi- patch medially on the fourth metasomal ster- cult to separate morphologically and there is 2000 ENGEL: BEE TRIBE AUGOCHLORINI 69 enough cohesion to the taxa that I have opted (57-1). The clade contains two distinctive to establish a group in the belief that it will groups. The first of these, the Pseudaugo- eventually be recognized as monophyletic. chlora group, possesses a pectinate inner me- The group can be recognized by the presence tatibial spur (24-1), a feebly truncate margin- of medioapical cleft on the fifth metasomal al cell apex (27-1), the presence of lateral sternum of males (46-2). If the monophyly processes on the apical margin on the sev- of the Paroxystoglossa group is confidently enth metasomal sternum of males (49-3), an established, then groups that I have herein unmodified venter on the penis valves (56- retained at the generic level should be con- 0), and strong ridge on the vertex (66-1). The sidered subgenera of a single genus. Megaloptidia group contains those augo- The genus Augochlorodes is placed in its chlorine genera sharing the absence of labral own monotypic genus group. The genus is teeth (3-0), the greatly narrowed prementum remarkably similar to species of the Augo- (10-1), and the presence of a labral distal chlora group (differentiated above in the di- process in males (34-1). Although relation- agnosis of the genus). It is difficult to con- ships within the clade are not well resolved, sider the following characters as synapomor- Megaloptidia is likely the most primitive ge- phies given that there is only one included nus of the group owing to the sclerotized species. However, Moure (in Schlindwein, dorsal gonostylar process; the remaining gen- 1998) has named a second species of the ge- era share the apomorphic presence of a par- nus that may potentially share some of these tially desclerotized dorsal process. Further features, thereby providing a good test for work on the internal phylogeny of the Me- the predictive power of the cladogram. How- galoptidiiti will need to explore this character ever, I have not seen seen material of this more fully. putative Augochlorodes, nor has Moure presented a description of the species (see sys- ANOMALIES tematic section on Augochlorodes above as well as appendix 1). Putatitive synapomor- Some anomalous features of the topology phies for the Augochlorodes group include (fig. 78) should be briefly noted. The posi- the combination of a pectinate inner metati- tions of the genera Andinaugochlora, Tem- bial spur (24-1), the meeting of the gradulus nosoma, and Ceratalictus are enigmatic and and antecosta on the fourth metasomal ster- should be investigated further. The genera num of males (42-1), the absence of apical Andinaugochlora and Neocorynurella are re- processes on the eigth metasomal sternum of markably similar and it is surprising that a males (50-0), and the broad spiculum (51-1). sister-group relationship, or at least closer af- The Augochlora-group of genera (Augo- finity, is not supported by these analyses. The chlora, Augochlorella, Ceratalictus, and Pe- placement of Temnosoma is also intuitively reirapis) are perhaps the most well-known displeasing. It has been long believed that group of augochlorine genera. The presence Temnosoma and Augochloropsis were sister of a carinate preoccipital ridge (15-1) and the taxa based on a unique labral morphology (1- presence of eusociality (72-4). One species 2) and the coarse sculpturing of Temnosoma of Augochlora s.l. (the fossil species, A. lep- (figs. 7Ð9, 30, 56, 57), Glyptochlora, and a toloba, described in appendix 1) has second- few primitive species of Paraugochloropsis. arily lost the preoccipital carina, while the Of these characters only the labral morphol- secondary loss of eusocial behavior is a syn- ogy is discrete and could be successfully apomorphy of species in Augochlora s.s. coded for cladistic analysis. The degree of sculpturing is quite variable, blending grad- MEGALOPTIDIITI ually from species of Temnosoma and Glyp- tochlora into the least sculptured species of The Megaloptidiiti includes thoe augo- Paraugochloropsis. Therefore, this feature, chlorines with a sharply pointed galeal apex while suggestive, provides little conclusive (6-1), the fusion of metasomal sterna seven information toward the grouping of these and eight prior to their lateral apices in males taxa. The quadrate labrum is incongruent (52-1), and the presence of a parapenial lobe with the pattern exhibited by other characters 70 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250 in the analysis and must therefore be inter- lower percentage reflectance in the nocturnal preted, for the time being, as convergent be- representative for wavelengths below 450 nm tween Augochloropsis and Temnosoma. (Engel, 1998, in prep.). Insect ocelli are po- Lastly, Ceratalictus also possesses a unique tentially involved in the detection of subtle character that seemingly allies it with Au- changes in light levels, especially at low in- gochlorella. Both of these genera have ex- tensities (Chapman, 1982). The enlargement ceedingly similar genitalia with a divided of the ocelli in each of these groups is pre- ventral gonostylar process (59-1). As it is, sumably an adaptation for navigating under Ceratalictus allies itself with Pereirapis in the extreme conditions of night where subtle the phylogeny, a particularly distressing po- changes in light levels might aid in their ori- sition. A more intuitive scenario would have entation abilities. Many crepuscular and noc- Augochlorella and Ceratalictus as sisters and turnal bees have proportionally larger ocelli Augochlora and Pereirapis as sisters. It will than diurnal species (Kerfoot, 1967). be important to determine whether these re- The nest architecture of the augochlorines lationships are stable to the addition of new is varied and has been the subject of phylo- character information. If a sister-group rela- genetic interpretation (e.g., Eickwort and tionship is ever supported for Ceratalictus Sakagami, 1979). When nest architecture and Augochlorella, then it would probably be traits are interpreted in light of the cladogram best to treat them as synonyms or as subgen- (fig. 78), the assumption that the primitive era of a single genus. nest architecture for the Augochlorini must have been something closely resembling the ETHOLOGICAL EVOLUTION general nest design seen in the tribe Halictini (Eickwort and Sakagami, op. cit.) is not sup- Details of behavioral evolution among the ported. A more congruent pattern of charac- augochlorines will be presented in a forth- ters entails a more complicated nest design coming paper specifically focusing on this with the following characteristics to be basal topic (Engel, in prep.). A preliminary outline among the augochlorines: (1) nests construct- of ethological evolution in Augochlorini is, ed in the soil, (2) a single turretless burrow, however, presented here. (3) burrow opening into a cell chamber, (4) The nocturnal genera Megalopta, Megom- cells oriented horizontally. Those nests re- mation s.s., and Megaloptidia do not form a sembling the simple Halictini-type (e.g., natural group, thereby suggesting indepen- lacking the cell chamber) are therefore re- dent derivations of this life-history strategy ductions. The pathways of change suggested and convergence on a presumably adaptive by the phylogeny are not as progressive as morphology. Each of these groups possess those of the previous studies (Sakagami and greatly enlarged ocelli, large body size, and Michener, 1962; Eickwort and Sakagami, pale integumental pigmentation, all possibly 1979). Instead of a series of changes, the to- specializations for activity at night. Interest- pology suggests a major change between the ingly, other nocturnal Hymenoptera, such as Halictini and Augochlorini with subsequent the nocturnal vespids Apoica and Provespa independent derivations or reductions in the and the nocturnal ichneumonids of the subtribe each occurring without evidence of pro- family Ophioninae, also exhibit strikingly ceeding through one of the ‘‘simpler’’ plans enlarged ocelli as well as generally pale body (Engel, 1998, in prep.). pigmentation (coloration is not universal, The genera Temnosoma, Megommation however, as some species of all of these (Cleptommation), and Megalopta (Noctorap- groups, including Megalopta and Megalop- tor) are not closely related, indicating three tidia, can be fairly dark). The general loss of derivations of cleptoparasitism in the tribe. body pigmentation may be the result of re- Each exhibits the typical suite of morpholog- laxed pressure from ultraviolet radiation. A ical characters associated with parasitic be- cursory examination of the UV-reflectance havior: absence of scopal hairs on the me- abilities of diurnal and nocturnal augochlor- tatibia, absence of a basitibial plate, and the ines as well as other brilliant metallic-green simple, bladelike or scythelike mandible. In- bees (e.g., Euglossini) reveals a consistently formation on the specific modes of parasit- 2000 ENGEL: BEE TRIBE AUGOCHLORINI 71 ism and biology of these species is desper- fied after the separation of the African and ately needed. Until the hosts of these taxa South American continents. Engel (1998) hy- and their specific biologies are discovered, pothesized that the primitive condition for little more can be determined regarding their augochlorine nests was a single, turretless evolution. burrow in the soil opening into a horizontally oriented cell chamber. Interestingly, trace FOSSIL HISTORY fossils of the Late Cretaceous from Argen- tina assigned to the ichnogenus Uruguay The augochlorines are the most speciose (Roselli, 1938; Genise and Bown, 1996) group in the Miocene fauna of Hispaniola as strongly resemble nests made by extant au- it is presently understood. There are six de- gochlorine species and generally fit the prim- scribed augochlorine species representing itive reconstruction hypothesized by Engel three genera; Augochlora, Neocorynura, and (1998). The tribe presumably originated in Oligochlora. The remainder of the Domini- southern South America and extended north- can amber bee fauna consists of the groups ward, reaching at least the area of the West Protandrenini (Rozen, 1996), Chilicolini Indies by the early Miocene as shown by the (Michener and Poinar, 1996; Engel, 1999e), presence of three genera in Dominican am- Euglossini (Engel, 1999c; Poinar, 1998), ber. The species from the West Indies, in- Halictini (Michener and Poinar, 1996), Me- cluding the fossils, all share affinities with gachilinae (Engel, 1999d), and Meliponini species occurring in northern South America (Wille and Chandler, 1964; Camargo et al., (Eickwort, 1988; Engel, 1995c), supporting 2000). Although the Miocene species are the the notion that the West Indian augochlorine oldest augochlorine fossils presently known, fauna is derived from South America and not the tribe is undoubtedly much older. None of North or Central America, thereby following the Miocene groups (Neocorynura, Augo- the South AmericanÐCaribbean track of Ro- chlora s.l., Oligochlora s.l.) are basal in au- sen (1975). Finer level studies on the internal gochlorine phylogeny as it is presently un- phylogenetic relationships of various genera derstood (fig. 78). The tribe as a whole may (particularly Augochlora and Augochlorop- be as old as the early Paleocene or, more sis) will greatly facilitate an understanding of likely, the Late Cretaceous, having diversi- augochlorine biogeography.

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bee, Augochlora pura (Hymenoptera: Wcislo, W. T. Halictidae). J. Kansas Entomol. Soc. 1993. Communal nesting in a North Ameri- 39: 157Ð192. can pearly-banded bee, Nomia tetra- Strand, E. zonata, with notes on nesting behavior 1921. Apidologisches, insbesondere u¬ber pa- of Dieunomia heteropoda (Hymenop- la¬arktische Andrena-Arten, auf Grund tera: Halictidae: Nomiinae). Ann. En- von Material des Deutschen Entomo- tomol. Soc. Am. 86: 813Ð821. logischen Museums. Arch. Naturg. Wcislo, W. T., and M. S. Engel Abt. A 87: 266Ð304. Taschenberg, E. 1996. Social behavior and nest architecture of 1883. Die Gattungen der Bienen (Anthophi- nomiine bees (Hymenoptera: Halicti- la). Berliner Entomol. Z. 27: 37Ð100. dae; Nomiinae). J. Kansas Entomol. Timberlake, P. H. Soc. Suppl. 69: 158Ð167. 1958. Temnosoma, a genus of bees new to the Wenzel, J. W. United States (Hymenoptera: Halicti- 1992. Behavioral homology and phylogeny. dae). Pan-Pac. Entomol. 34: 34. Annu. Rev. Ecol. Syst. 23: 361Ð381. Vachal, J. Wille, A., and L. C. Chandler « 1903. Etude sur les Halictus d’Ame«rique 1964. A new stingless bee from the Tertiary (Hym.). Misc. Entomol. 11: 89Ð104, amber of the Dominican Republic (Hy- 121Ð136. menoptera; Meliponini). Rev. Biol. 1904. E« tude sur les Halictus d’Ame«rique (Hym.). Ibid. 12: 9Ð24, 113Ð128, 137Ð Trop. 12: 187Ð195. 144. Wolda, H., and D. W. Roubik 1911. E« tude sur les Halictus d’Ame«rique 1986. Nocturnal bee abundance and seasonal (Hym.). Ibid. 19: 9Ð24, 41Ð56, 107Ð bee activity in a Panamanian forest. 112. Ecology 67: 426Ð433.

APPENDIX 1 Descriptions and Records of Bees Pertinent to this Study

The following species are treated in this work Augochlora (Augochlora) nitidior Moure, since they provide significant range extensions for nomen nudum their respective genera, are important taxonomic changes, are rare additions to the augochlorine Augochlora (Augochlora) nitidior Moure in Schlind- fauna, or provide valuable information for a mor- wein, 1998: 51. phological understanding of a particular genus. Augochlora (Electraugochlora) leptoloba, new Augochlora (Augochlora) nigrocyanea Cockerell species Augochlora nigrocyanea Cockerell, 1897a: 144. Halictus zophodes Vachal, 1911: 16. Figures 79Ð81 Augochlora smaragdina variety atrata Friese, 1916: 312. Augochlora smaragdina variety atra Friese, 1916: 313. DIAGNOSIS: As for the subgenus (refer to page Lapsus calami. 32). Augochlora (Odontochlora) essequibensis Cockerell, DESCRIPTION: Female. Total body length 8.5 1923: 445. NEW SYNONYMY. mm; forewing length 4.6 mm. Head length 2.1 Augochlora cyanaspis Cockerell, 1931: 552. mm. Lower third of clypeus below lower tangent of compound eyes. Basal vein distad cu-a by 3 COMMENTS: The possible synonymy of A. es- times vein width; 1r-m confluent with 1m-cu; 2r- sequibensis with A. nigrocyanea was first sug- m distad 2m-cu by 3 times vein width; first sub- gested to me by my late mentor George C. Eick- marginal cell about as long as combined lengths wort (personal commun.) and I have since been of second and third submarginal cells; second able to confirm his suspicions through examina- submarginal cell parallel-sided; distal hamuli ar- tion of the types. This synonymy is published here ranged 3-1-1-2. for the first time but should probably be attributed Clypeus and supraclypeal area with widely to Eickwort as he was the first person to correctly scattered faint, coarse punctures, integument be- recognize the conspecific nature of the taxa. tween faintly imbricate. Face and vertex minutely 80 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

Fig. 79. Photomicrograph of holotype female of Augochlora (Electraugochlora) leptoloba, new subgenus and new species, in Dominican amber (photograph by author).

Figs. 80Ð81. Augochlora (Electraugochlora) leptoloba, new subgenenus and new species, female. 80. Head, oblique anterior aspect. 81. Head in profile. 2000 ENGEL: BEE TRIBE AUGOCHLORINI 81 punctured, punctures separated by width or less, cell longer than combined lengths of second and integument otherwise faintly imbricate. Gena as third submarginal cells; second submarginal cell on face except punctures become separated by slightly narrowed anteriorly; anterior border of two times puncture width on lower third. Postgena third submarginal cell three-quarters that of pos- faintly imbricate and impunctate. Pronotum faint- terior border; distal hamuli arranged 2-1-2. ly imbricate. Mesoscutum and scutellum imbricate with very widely scattered, faint punctures. Clypeus with small, weak punctures separated Metanotum weakly rugulose. Preepisternum by 2Ð4 times puncture width, integument other- coarsely punctured, punctures contiguous on up- wise smooth. Supraclypeal area with minute per third, separated by width or less on lower two- punctures separated by 1Ð2 times puncture width, thirds. Mesepisternum coarsely punctured, punc- integument between smooth. Face at and below tures separated by puncture width or less, integ- level of antennae with punctures separated by less ument otherwise granular; hypoepimeron weakly than a puncture width, integument smooth; above rugulose with minute punctures along anterior level of antennae punctures become much smaller third separated by less than a puncture width. Me- and contiguous except at emargination of com- tepisternum weakly rugulose. Propodeal lateral pound eye where separated by less than width and surface faintly imbricate with widely scattered coarse punctures; posterior surface faintly imbri- forming impunctate spot bordering the crux of the cate; basal area imbricate with faint basal striae. eye. Punctures of vertex minute and separated by Terga and sterna imbricate. width, integument smooth. Gena as on vertex ex- Mandible and labrum brown. Head dull metal- cept punctures separated by 2Ð3 times puncture lic green with a few faint metallic copper high- width. Postgena imbricate and impunctate. Pron- lights except clypeal apex brown. Mesosoma as otum faintly imbricate. Mesoscutum with minute on head; legs brown; wings hyaline, venation punctures over smooth integument; medially brown. Terga dark brown with a few faint metallic punctures separated by width or slightly more on green highlights; sterna brown. anterior half and separated by 2Ð5 times width on Pubescence golden and generally sparse except posterior half; lateral thirds with punctures sepa- as indicated: on mesoscutum, scutellum, and metanotum fuscous; on outer surfaces of meso- and rated by width. Scutellum with minute punctures metatibiae, basitarsi, tarsomeres 2Ð4 long, simple, separated by 1Ð2 times width, integument be- fuscous, and stiff; on terga fuscous. tween smooth. Metanotum rugulose. Preepister- Male. Unknown. num strongly rugose. Punctures of mesepisternum HOLOTYPE: Female, Miocene amber of the Do- separated by puncture width on smooth integu- minican Republic (MACT), accession number M- ment. Metepisternum with punctures separated by 2521. less than puncture width, integument smooth. Pro- ETYMOLOGY: The specific epithet is a combi- podeal lateral and posterior surfaces imbricate; nation of the Greek words leptos (meaning basal area of propodeum with strong striae radi- ‘‘small’’) and lobos (meaning ‘‘lobe’’) and refers ating from basal margin, integument between stri- to the weakly developed epistomal lobe that protrudes into the basal margin of the clypeus. ae smooth and shining. Anterior surface of T1 with punctures very widely spaced, integument between smooth; remainder of T1 with minute Augochlora (Oxystoglossella) punctures separated 1Ð4 times puncture width, in- rightmyerae, new species tegument faintly imbricate; sculpturing of T2ÐT5 DIAGNOSIS: This species can be distinguished as on posterior half of T1; apical margins of terga from other Oxystoglossella by the sculpturing of imbricate and impunctate; sterna imbricate. the clypeus, face, mesoscutum, and pleura and the Mandible dark brown except middle third coloration of the face and mesoscutum (all de- which is amber and apex which is reddish brown. scribed below). Labrum dark brown. Clypeus and supraclypeal DESCRIPTION: Female. Total body length 8.8 area dark brown with strong metallic red or purple mm; forewing length 5.8 mm. Head as long as highlights except along upper border of supracly- wide (length, width 2.4 mm). Lower two-thirds of peal area which is brilliant metallic green. Anten- clypeus below lower tangent of compound eyes. na brown. Remainder of head brilliant metallic Intertegular distance 1.8 mm. Basal vein distad green with scattered gold or weakly coppery high- cu-a by three times vein width; 1r-m confluent lights. Mesosoma brilliant metallic green except with 1m-cu; 2r-m distad 2m-cu by seven times mesoscutum and scutellum which are dark brown, vein width, 2r-m gently curved; first submarginal nearly black; mesoscutum with a few faint metal- 82 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250 lic red or purple highlights on borders; scutellum NEW RECORD: One male; Costa Rica, Heredia with stronger metallic red or purple highlights, Province, ca. 12 km SW Horquetas (Rara Avis), particularly strong on axilla, and some metallic 550 m, 18-23 January 1989, D. A. Grimaldi green highlights along anterior border. Tegula (AMNH). COMMENTS: This species was, until now, known dark brown. Wing membrane hyaline; veins solely on the basis of the holotype female in the brown. Legs brown. Metasoma dark brown except collection of the LACM. A single male has re- metallic green highlights faint on central discs of cently been identified among unsorted Hymenop- T2ÐT5, stronger laterally on terga; sterna without tera at the AMNH and is reported upon here. This highlights. is the second species of the subgenus for which Pubescence golden or white except on scutel- males and females are both known. lum and T5ÐT6, which is slightly fuscous. Ante- Lasioglossum (Evylaeus) biciliatum rior surface of T1 with mat of appressed, short (Friese), new combination plumose hairs. Male. Unknown. Corynura biciliata Friese, 1916: 319. HOLOTYPE: Female; Bolivia, La Paz, Alto R«õo COMMENTS: This Costa Rican species was Beni, south of Rio Inicua, 1100 m, 15Ð18 January placed by Friese (1916) in the augochlorine genus 1976, Luis E. Penã (AMNH). Corynura, which, as presently understood, does ETYMOLOGY: The specific epithet honors Ms. not occur outside of southern South America. Ex- Molly G. Rightmyer, talented artist and entomol- amination of the type (ZMHB) reveals that this ogist, who has offered every assistance in my ef- species is, in fact, a member of the tribe Halictini forts to curate the AMNH bee collection. (or Gastrohalictini: see Table 4), genus Lasioglos- sum. Padre Moure had also examined the type, a Augochlorella eusticta Moure, nomen nudum year before me, as evidenced by the date on his determination label, and arrived at the same con- Augochlorella eusticta Moure in Schlindwein, 1998: 51. clusion. Augochlorodes clementis Moure, nomen nudum Megalopta (Megalopta) intermedia Augochlorodes clementis Moure in Schlindwein, 1998: Sakagami, nomen nudum 51. Megalopta intermedia Sakagami, 1979: 86.

Augochloropsis scabriceps Moure, nomen COMMENTS: This name appeared in a popular nudum article on the diversity and biology of halictine bees. Unfortunately no species has ever been pro- Augochloropsis scabriceps Moure in Schlindwein, 1995: posed with this epithet. 52. Neocorynurella cosmetor (Vachal), Caenaugochlora (Caenaugochlora) elisabethae new combination Engel Halictus cosmetor Vachal, 1911: 51. Caenaugochlora (Caenaugochlora) elisabethae Engel, Augochlora cosmetor (Vachal); Moure and Hurd, 1987: 1997c: 91. 276. Vachalius cosmetor (Vachal); Moure, 1999: 76. NEW RECORDS: One male; Costa Rica, Punta- renas Coto Brus, Las Alturas Biological Station, NEW RECORD: One female, Venezuela, Merida, near lecheria of Finca Las Alturas, near Coto«n, La Montana, cable car station, 2442 m, 23Ð25 1540 m, 13 June 1992, J.V. McHugh, lot #CR92- February 1968, P. and B. Wygodzinsky and M. 183 (CUIC). One female, four males; Costa Rica, Cormons (AMNH). Puntarenas, Las Alturas Field Station, 20 km N. San Vito de Hava, 1400 m, 20Ð24 May 1991, De Oligochlora (Soliapis) rozeni, Vries, malaise trap (AMNH). One female; Costa new species Rica, Puntarenas, Las Alturas Field Station, 20 Figure 82 km N. San Vito de Hava, 1540 m, 10Ð31 July 1992, C. Snyder, malaise trap (AMNH). DIAGNOSIS: As for the subgenus (refer to p. 47). Caenaugochlora (Ctenaugochlora) DESCRIPTION: Female. Total body length 8.2 donnae Engel mm; forewing length 5 mm. Head apparently about as long as wide (length, width 2 mm). Com- Caenaugochlora (Ctenaugochlora) donnae Engel, pound eyes strongly convergent below. Pronotal 1995a: 283. lateral angle orthogonal. Basal vein distad cu-a by 2000 ENGEL: BEE TRIBE AUGOCHLORINI 83

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

two times vein width; ScϩR pigmented as other ular; basal area granular, without basal striae. Ter- wing veins; 1r-m confluent with 1m-cu; 2r-m dis- ga faintly imbricate. tad 2m-cu by four times vein width, 2r-m rela- Mandible and labrum dark brown. Head bril- tively straight; first submarginal cell longer than liant metallic green-gold with scattered coppery combined lengths of second and third submarginal highlights. Antenna brown except with yellow cells; second submarginal cell parallel-sided; an- markings on inner surface of scape. Mesosoma as terior border of third submarginal cell approxi- on head except tegula brown; wings hyaline, ve- mately equal to that of second submarginal cell; nation dark brown; legs brown. Terga dark brown distal hamuli arranged 3-1-2. Inner hind tibial with metallic green highlights and apical margins spur with 4 teeth excluding apex. brown without highlights; sterna brown. Clypeus and supraclypeal area weakly granular, Pubescence golden except on mesoscutum and impunctate. Face below level of antennae weakly scutellum fuscous and on meso- and metatibiae granular with faint, crowded punctures appearing and tarsi which is fuscous, highly branched, and by margin of compound eye, clypeus, and near long. antennal socket; upper half of face apparently Male. Unknown. weakly granular with faint, nearly contiguous HOLOTYPE: Female; Miocene amber of the Do- punctures. Gena and postgena apparently faintly minican Republic (MACT), accession number M- imbricate. Pronotum faintly imbricate. Mesoscu- 2523. tum (and apparently the same on scutellum) ETYMOLOGY: The specific epithet is a patronym- weakly punctured, punctures separated by punc- ic honoring Dr. Jerome G. Rozen, Jr., of the ture width or less, integument between imbricate. AMNH. Jerry helped me enormously during the Metanotum imbricate. Preepisternum rugose. Me- undertaking of my graduate research and since sepisternum granular with faint, coarse punctures then in my postdoctoral studies at the AMNH. I separated by a puncture width or less. Metepis- am sincerely grateful for his support and friend- ternum imbricate. Propodeal lateral surface gran- ship. 84 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

Pereirapis semiaurata (Spinola) am confident of Ebmer’s comparisons. I therefore have considered H. caucasicus as identical to Halictus semi-auratus Spinola, 1853: 86. these latter two taxa and thereby as a synonym of Augochlora Titania Smith, 1853: 75. H. semiauratus. Augochlora semiaurata (Spinola): Smith, 1853: 79. I have also not had the opportunity to see Padre Halictus caucasicus Radoszkowsky, 1868: 77. NEW Moure’s types of P. rhizophila (located in DZUP), SYNONYMY. Augochlora titanea Dalla Torre, 1896: 96. Nomen emen- although I have seen specimens of this species datum (unjustified). identified by Moure. In 1944 and 1950 Moure Augochlora seminigra Cockerell, 1897a: 144. NEW considered his species as a synonym of A. semi- SYNONYMY. nigra. I am similarly confident with Moure’s de- Corynura titania (Smith): Cockerell, 1900: 375. termination which accords with the specimens Halictus hypixis Vachal, 1904: 127, 142. identified by him that I have examined. I therefore Halictus chrysaspis Vachal, 1911: 18. NEW SYNON- consider P. rhizophila as a synonym of H. sem- YMY. iauratus. Halictus myrrhites Vachal, 1911: 18. NEW SYNONY- MY. Halictus simotes Vachal, 1911: 22, 55. NEW SYNON- Pseudaugochlora pulchra, new species YMY. Halictus cerasis Vachal, 1911: 55. NEW SYNONYMY. DIAGNOSIS: This enigmatic species can be easily Augochlora cladopyga Cockerell, 1914: 324. NEW recognized from currently known Pseudaugochlo- SYNONYMY. ra species by the strongly protuberant clypeus, the Pereirapis rhizophila Moure, 1943a: 461. NEW SYN- ONYMY. ventrally bent mandible with its mediodorsal Augochlorella bidentata Michener, 1954b: 57. NEW flange, the long prementum which extends pos- SYNONYMY. terad to the procoxae, the serrate malus, the outer Augochlorella cerasis (Vachal): Moure in Ordway, hind tibial spur bent strongly at its apex, and the 1966b: 511. strong rugae of the propodeal basal area. Augochlorella chrysaspis (Vachal): Moure in Ordway, DESCRIPTION: Female. Total body length 11.34 1966b: 511. Augochlorella semiauratus (Spinola): Moure in Ordway, mm; forewing length 7.7 mm. Head slightly wider 1966b: 512. than long (length 2.7 mm, width 2.8 mm). Man- Pereirapis rhysophila Ordway, 1966b: 512. Lapsus cal- dible strongly bent ventrally near its midpoint ami. with dorsal flange or lamellae leading to strong, Augochlorella simotes (Vachal): Moure in Ordway, rounded, subapical tooth. Labral basal area 1966b: 512. strongly delimited from distal process by carina. Augochlorella cladopyga (Cockerell): Moure in Ord- way, 1966b: 512. Clypeus strongly protruding from face in profile. Pereirapis caucasica (Radoszkowsky): Ebmer, 1977: Hypostomal ridge weakly lamellate. Prementum 575. very long (although not narrowed; approximately Pereirapis cerasis (Vachal): Moure and Hurd, 1987: 5.5 times longer than wide), extending posteriorly 247. to procoxae. Glossa nearly as long as prementum. Pereirapis chrysaspis (Vachal): Moure and Hurd, 1987: Maxillary palpi relatively short. Compound eyes 247. Pereirapis simotes (Vachal): Moure and Hurd, 1987: only weakly converging below; strongly emargin- 248. ate above level of antennae. Intertegular distance Augochlora myrrhites (Vachal): Moure and Hurd, 1987: 1.96 mm. Strong anterior basitarsal brush; malus 277. of antenna cleaner serrate; mesotibial spine very Pereirapsis [sic] risofila Alves dos Santos, 1997: 6. Lap- short and strongly bent inward at apex; outer hind sus calami. tibial spur strongly bent inward towards apex; in- Pereirapis myrrhites (Vachal): Moure, 1999: 83. ner hind tibial spur with four long teeth (exclud- COMMENTS: Two synonymies are proposed ing apex). Basal vein distad cu-a by 4.5 times vein above even though I have not had the opportunity width; 1m-cu distad 1r-m by vein width; 2r-m dis- to examine the type material firsthand. The first is tad 2m-cu by five times vein width; pterostigma H. caucasicus the holotype for which is located narrow, barely wider than distance from C to in the Institute of Systematic Zoology, Polish ϩ Academy of Sciences, Kra«kow. According to the Sc R (including widths of these veins); first sub- curator the holotype is in poor condition and not marginal cell slightly longer than second and third suitable for shipment (W. Celary, personal com- combined; second cell not narrowed anteriorly; mun., 1996). Father Ebmer (1977), however, had anterior border of third cell longer than that of examined the type and considered it identical to second; distal hamuli arranged 3-1-1-2. A. seminigra and A. bidentata. I have seen the Clypeus with weak, coarse punctures separated types of both A. seminigra and A. bidentata and by less than a puncture width, integument be- 2000 ENGEL: BEE TRIBE AUGOCHLORINI 85 tween imbricate; supraclypeal area as on clypeus es, progressively becoming longer ventrally to- although faintly imbricate and shining. Face be- wards border of gena and postgena. Postgena with low antennae with coarse punctures separated by long, branched, white hairs. Pronotum with mi- less than a puncture width, punctures stronger nute, simple, appressed hairs on posterior border, than those of clypeus, integument between imbri- dorsally on anterior border and medially; laterally cate. Face above level of antennae with punctures such hairs nearly obscure the surface, thinning by smaller and contiguous; punctures becoming very pronotal lobe. Mesoscutum with scattered fuscous weak by ridged vertex. Gena smooth with minute hairs similar to those of face. Scutellum and me- punctures separated by 3Ð4 times puncture width. tanotum with pubescence similar to that of me- Postgena imbricate and impunctate. Pronotum soscutum although much longer. Pleura with a smooth with minute punctures separated by punc- few fuscous hairs as on mesoscutum, mostly with ture width or less; laterally punctures disappear- scattered golden hairs each with a few short ing. Mesoscutum with punctures separated medi- branches. Scopal hairs white; remaining pubes- ally by puncture width, integument between cence of legs gold except fuscous on outer sur- smooth; at borders punctures separated by less faces of mesotibia, metatibia, outer surface of pro- than width and much smaller, contiguous around tarsus, and entirety of meso- and metatarsus. Pro- parapsidal lines. Scutellum smooth with scattered podeal lateral surface with hairs like those of large punctures. Metanotum weakly roughened. pleura except intermixed wih short, simple, sub- Preepisternum rugulose on upper border, other- erect white hairs; posterior surface with short wise strongly and contiguously punctured. Mese- hairs like that of lateral surface plus long white pisternum smooth with punctures separated by hairs with short branches. Terga with scattered less than puncture width except on lower half punctures widely spaced. Metepisternum with suberect white hairs; anterior surface of T1 with weak transverse striae on upper third, remainder long, mostly simple, gold to white hairs; T2ÐT6 with scattered punctures and smooth integument with gradually more numerous fuscous hairs in- between. Propodeal lateral surface punctured, termixed on lateral and posterior borders, such punctures separated by less than puncture width, predominant by T5. Sterna with scattered, long, although on border with metepisternum and dorsal mostly simple, gold or fuscous hairs; fuscous surface faintly imbricate, a few weak transverse hairs most predominant on S5ÐS6. striae near metacoxal base; posterior surface Male. Unknown. smooth with widely spaced punctures; basal area HOLOTYPE: Female; Peru, Hua«nuco, Cueva de with strong rugae, integument between smooth. las Pavas, Tingo Maria, 12 July 1974, C. Porter Anterior surface of T1 smooth with widely spaced and L. Stange (CUIC). minute punctures, medially with larger punctures ETYMOLOGY: The specific epithet is derived separated by width or less, posterior border with from the Latin word pulcher meaning ‘‘beauti- microscopic punctures widely spaced; T2 with ful.’’ minute punctures separated by width, posterior COMMENTS: This species is quite unique among border as for that of T1; T3ÐT5 imbricate; sterna Pseudaugochlora and as more material is ac- imbricate with weak widely scattered punctures. quired it might be worthy of its own subgenus Mandible dark brown with reddish apex. Head (particularly if related species are discovered). For brilliant metallic green with a few coppery high- now, however, I hesitate to recognize subgenera lights except apex of clypeus and antennae dark in Pseudaugochlora and leave such a determina- brown. Mesosoma brilliant metallic green with tion to future students of the Augochlorini. scattered copper highlights, highlights strongest on mesoscutum and pleura; tegula brown except Rhectomia mourei (Eickwort) metallic green on inner border. Wing membrane hyaline; veins dark brown. Legs brown with metallic green highlights on all surfaces except tarsi. Corynurella mourei Eickwort, 1969a: 514. Metasoma brilliant metallic green except apical Rhectomia mourei (Eickwort): Engel, 1995b: 308. margins of T1ÐT3 brown; sterna brown with strong metallic green highlights on apical halves. NEW RECORD: One female; Bolivia, La Paz, Mandible with row of long, simple hairs on Alto R«õo Beni, south of R«õo Inicua, 1100m, 15Ð ventral margin. Face with scattered fuscous hairs, 18 January 1976, L.E. Penã (AMNH). such hairs with a few short branches. Hairs of COMMENTS: This is the first record of this genus gena suberect and golden with a few short branch- from Bolivia. 86 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250 2000 ENGEL: BEE TRIBE AUGOCHLORINI 87 88 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 250

APPENDIX 3 Species of Augochlorini Examined for Phylogenetic Studies Halictid bee species examined in the course of gochlora): C. costaricensis (Friese), C. cupriven- these studies. Daggers (†) indicate fossil species. tris (Vachal), C. elisabethae Engel, C. flagrans Not all characters were coded from each species (Vachal), C. gemmella (Cockerell), C. inermis (e.g., males are unknown for some species and (Vachal), C. jeffreyi Engel, Caenaugochlora others are represented solely by the holotype, (Caenaugochlora) spp. 1Ð2; Caenaugochlora thereby excluding them from some of the dissec- (Ctenaugochlora): C. algeri Engel, C. beethoveni tions necessary to examine certain traits). Many Engel, C. donnae Engel, C. perpectinata (Mich- additional species have been studied (particularly ener); Ceratalictus: C. ischnotes (Vachal), C. sti- in the genera Augochloropsis, Caenohalictus, gon (Vachal), C. clonius (Bre`thes) [ϭ C. theius Dialictus, Megalopta, and Neocorynura), but (Schrottky): see Moure, 1999]; Chlerogas: C. ar- these were not focused on in great detail or used aguaensis Brooks and Engel, C. boliviensis in constructing the data matrix presented in ap- Brooks and Engel, C. chlerogas (Vachal), C. col- pendix 2. Specimens are deposited in the institu- ombiensis Brooks and Engel, C. cyaneus Brooks tions listed under Materials and Methods. and Engel, C. hirsutipennis Cockerell, C. nephos Brooks and Engel, C. tiara Brooks and Engel, C. SUBFAMILY HALICTINAE townesi Brooks and Engel; Chlerogella: C. elon- AUGOCHLORINI: Andinaugochlora: A. joannisi (Vachal), A. micheneri Eickwort, Andi- gaticeps Michener, C. nasus (Enderlein), Chlero- naugochlora spp. 1Ð3; Ariphanarthra: A. palpalis gella spp. 1Ð15; Chlerogelloides: C. femoralis Moure; Augochlora (Augochlora): A. buscki Engel et al., C. simplex Engel and Brooks; Cor- Cockerell, A. decorata (Smith), A. esox (Vachal), ynura (Callistochlora): C. chloris (Spinola), C. A. essequibensis Cockerell, A. foxiana Cockerell, prothysteres (Vachal); Corynura (Corynura): C. A. hallinani Michener, A. ignifera Crawford, A. apicata (Spinola), C. chilensis (Spinola), C. cor- mulleri Cockerell, A. nigrocyanea Cockerell, A. inogaster (Spinola), C. cristata (Smith), C. herbsti pachytes (Vachal), A. praeclara Cresson, A. pura (Alfken), C. rubella (Haliday); Halictillus: Ha- (Say), A. regina Smith, A. repandirostris (Vach- lictillus spp. 1Ð3; Ischnomelissa: I. cyanea al), A. smaragdina Friese, Augochlora (Augochlo- Brooks and Engel, I. ecuadoriana Brooks and En- ra) spp. 1Ð4; Augochlora (Oxystoglossella): A. gel, I. lescheni Brooks and Engel, I. octogesima antonita Michener, A. aurifera Cockerell, A. cor- Brooks and Engel, I. rhina Brooks and Engel, I. diaefloris Cockerell, A. matucanensis Cockerell, zonata Engel; Megalopta (Megalopta): M. amo- A. nominata Michener, A. rightmyerae Engel (see ena (Spinola), M. centralis Friese, M. ecuadoria appendix 1), A. thalia Smith, Augochlora (Oxy- Friese, M. fornix (Vachal), M. genalis Meade- stoglossella) spp. 1Ð3; Augochlorella: A. aurata Waldo, M. ochrias (Vachal), M. purpurata (Smith), A. bracteata Ordway, A. comis (Vachal), (Smith), M. sodalis (Vachal), Megalopta spp. 1Ð A. edentata Michener, A. gratiosa (Smith), A. neg- 4; Megalopta (Noctoraptor): M. byroni Engel et lectula (Cockerell), A. persimilis (Viereck), A. po- al., M. noctifurax Engel et al.; Megaloptidia: M. moniella (Cockerell), A. striata (Provancher), A. contradicta (Cockerell), M. nocturna (Friese), M. urania (Smith), Augochlorella spp. 1Ð2; Augo- saulensis Engel and Brooks; Megaloptilla: M. chlorodes: A. turrifaciens Moure; Augochloropsis callopis (Vachal), M. byronella Engel and Brooks; (Augochloropsis): A. auriventris (Friese), A. bra- Megommation (Cleptommation): M. minutum chycephala Moure, A. callichroa (Cockerell), A. (Friese); Megommation (Megaloptina): M. ogil- flammea (Smith), A. ignita (Smith), A. multiplex viei (Cockerell), M. festivagum (Dalla Torre), Me- (Vachal), A. notophos (Vachal), A. wallacei gommation (Megaloptina) sp.; Megommation (Cockerell); Augochloropsis (Glyptochlora): A. (Megommation): M. insigne (Smith); Megom- atropos (Smith), A. ornata (Smith), A. refulgens mation (Stilbochlora): M. eickworti Engel et al., (Smith); Augochloropsis (Paraugochloropsis): A. Megommation (Stilbochlora) sp.; Neocorynura: anonyma (Cockerell), A. auriferina Michener, A. N. cercops (Vachal), N. colombiana Eickwort, N. aurifluens (Vachal), A. chloera (Moure), A. cras- discolor (Smith), †N. electra Engel, N. fumipennis siceps Moure, A. cupreola (Cockerell), A. electra (Friese), N. oiospermi (Schrottky), N. panamensis (Smith), A. hebescens (Smith), A. iris (Schrottky), Engel, N. papallactensis Engel, N. rhytis (Vach- A. metallica (Fabricius), A. smithiana (Cockerell) al), N. rufa Michener, N. sulfurea Engel, Neocor- [ϭ A. aphrodite (Schrottky)], A. sthena Schrottky, ynura spp. 1Ð7; Neocorynurella: N. cosmetor A. sumptuosa (Smith), A. vesta (Smith) [ϭ A. ni- (Vachal) (see appendix 1), N. seeleyi Engel and griscopis (Vachal)], Augochloropsis (Paraugo- Klein, N. virida Engel and Klein (nomen emen- chloropsis) sp. 1Ð6; Caenaugochlora (Caenau- datum); †Oligochlora: †O. eickworti Engel, †O. 2000 ENGEL: BEE TRIBE AUGOCHLORINI 89 grimaldii Engel, †O. micheneri Engel, †O. rozeni A. sericeus (Forster), A. texanus Cresson, A. vie- Engel (appendix 1), †Oligochlora sp. (Engel and quesensis Cockerell; Caenohalictus: C. eberhar- Rightmyer, in press); Paroxystoglossa: P. andro- dorum Michener, Caenohalictus spp. 1Ð5; Dia- mache (Schrottky), P. barbata Moure, P. brachy- lictus: D. breedi (Michener), D. busckiellus cera Moure, P. crossotos (Vachal), P. jocasta (Cockerell), D. mestrei (Baker), D. parvus (Cres- (Schrottky), P. mimetica Moure, P. seabrai Mou- son), D. proangularis (Ellis), Dialictus spp. 1Ð6; re, P. spiloptera Moure, P. transversa Moure; Habralictus: H. bimaculatus Michener, H. can- Pereirapis: P. semiaurata (Spinola) (see appendix alictulatus Moure, Habralictus spp. 1Ð2; Habra- 1); Pseudaugochlora: P. crawfordi (Vachal), P. lictellus: H. auratus (Ashmead), Habralictellus graminea (Fabricius), P. pandora (Smith), P. spp. 1Ð3; Halictus: H. farinosus Smith, H. ligatus praepotens (Vachal), P. sordicutis (Vachal) [ϭ P. Say; Homalictus: H. dampieri Cockerell, Hom- nigerrima (Friese)], P. pulchra Engel (see appen- alictus sp.; Lasioglossum: L. coriaceum (Smith), dix 1); Rhectomia: R. harrisoni Engel, R. lie- L. crocoturum (Vachal), L. leucozonium bherri Engel, R. mourei (Eickwort), R. pumilla (Schrank), L. trizonatum (Cresson); Mexalictus: Moure, Rhectomia spp. 1Ð2; Rhinocorynura: R. M. arizonensis Eickwort, M. mexicanus Eickwort, briseis (Smith), R. crotonis (Ducke), R. inflaticeps M. micheneri Eickwort, Mexalictus spp. 1Ð3; Rui- (Ducke); Temnosoma: T. metallicum Smith, T. zanthedella: R. mutabilis (Spinola), R. nigroca- smaragdinum Smith; Thectochlora: T. alaris erulea (Spinola); Zonalictus: Z. rufobasalis (Alf- (Vachal); Xenochlora: X. chalkeos Engel et al., X. ken), Z. viridifilosus (Cockerell). ianthina (Smith), X. nigrofemorata (Smith), X. ochrosterna Engel et al. SUBFAMILY NOMIINAE NOMIOIDINI: Nomioides: N. variegatus Dieunomia: D. heteropoda (Smith), D. trian- (Olivier). gulifera (Vachal); Lipotriches: Lipotriches sp. HALICTINI: Agapostemon: A. poeyi (Lucas),

APPENDIX 4 Sources of Nest Architectural Characters Information on the nesting biology of halictine Rau (1929: as Epinomia), Wcislo (1993), Wcislo genera was extracted from the sources indicated and Engel (1996); Habralictus: Michener and below. These data were used in constructing the Lange (1958b), Michener et al. (1979); Halictil- matrix presented in appendix 2. lus: Claude-Joseph (1926: as Halictus); Halictus: Eickwort (1985); Lasioglossum: Knerer and At- -Rhopalomelissa sen ؍) Agapostemon: Abrams and Eickwort (1980), wood (1962); Lipotriches Eickwort (1981), Eickwort and Eickwort (1969), su Pauly, 1990): Hirashima (1961), Michener Roberts (1969); Augochlora: Engel (personal (1969b), Wcislo and Engel (1996); Megalopta: obs.), Eickwort and Eickwort (1973b), Sakagami Janzen (1968), Sakagami (1964), Sakagami and and Moure (1967), Stockhammer (1966); Augo- Moure (1967); Megommation:Jo¬rgensen (1912: chlorella: Engel (personal obs.), Eickwort and as Megalopta), Michener and Lange (1958b), Eickwort (1973a), Knerer and Atwood (1962), Sakagami and Moure (1967); Neocorynura: Ordway (1961, 1966a), Sakagami and Moure Michener et al. (1966), Sakagami and Moure (1967); Augochlorodes: Michener and Seabra (1967), Schremmer (1979); Nomioides: Pesenko (1959); Augochloropsis: Michener and Lange (1983); Oxystoglossella: Eickwort and Eickwort (1972), Sakagami and Moure (1967); Paraugo- (1959), Michener and Seabra (1959); Caenaugo- chloropsis: Eickwort and Sakagami (1979), Mich- chlora: Michener and Kerfoot (1967: as Pseudau- ener and Lange (1959); Paroxystoglossa: Miche- gochloropsis); Caenohalictus: Michener and ner and Lange (1958a, b), Michener and Seabra Lange (1958b: as Caenaugochlora), Michener et (1959); Pereirapis: Eickwort and Sakagami al. (1979), Sakagami and Moure (1967); Callis- (1979), Oliveira Campos (1980); Pseudaugochlo- tochlora: Claude-Joseph (1926: as Halictus); Cor- ra: Michener and Lange (1958b), Sakagami and ynura: Claude-Jospeh (1926: as Halictus); Dia- Moure (1967: as Pseudaugochloropsis); Rhino- lictus: Engel (personal obs.), Michener and Sea- corynura: Eickwort and Sakagami (1979); Rui- bra (1959), Michener et al. (1979), Sakagami and zanthedella: Claude-Joseph (1926: as Halictus); Moure (1967); Dieunomia: Cross and Bohart Xenochlora: Engel et al. (1997), Smith (1861: as (1960: as Nomia), Parker et al. (1986: as Nomia), Megalopta).