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Classification of the Bee Tribe Augochlorini (Hymenoptera: Halictidae)

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Classification of the Bee Tribe Augochlorini (Hymenoptera: Halictidae) View metadata, citation and similar papers at core.ac.uk brought to you by CORE 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 (285–247 B.C.) by 70–72 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 sub- jective 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 augo- chlorine 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 sum- marized and infratribal groups defined. Most notably, the tribe is divided into two monophy- letic 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. nigro- cyanea 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
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