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A Revision of the Bee Genus Nomada in Argentina (Hymenoptera, Apidae, Nomadinae)

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A Revision of the Bee Genus Nomada in Argentina (Hymenoptera, Apidae, Nomadinae) Roig Alsina: Revision of the bee genusRev. Mus. Nomada Argentino Cienc. Nat., n.s.221 11(2): 221-241, 2009 Buenos Aires, ISSN 1514-5158 A revision of the bee genus Nomada in Argentina (Hymenoptera, Apidae, Nomadinae) Arturo ROIG ALSINA Museo Argentino de Ciencias Naturales «Bernardino Rivadavia,» Av. Angel Gallardo 470, 1405 Buenos Aires, Argentina. Abstract: A revision of the bee genus Nomada Scopoli in Argentina is presented. Nine species are recognized in this region, five of which are described as new: N. mesopotamica, N. longula, N. chacoana, N. missionica, and N. turrigera. Lectotypes are designated for N. pampicola Holmberg, 1886, and N. costalis Brèthes, 1909. A new name, N. holmbergiana, is proposed for Hypochrotaenia parvula Holmberg, 1886, preoccupied in Nomada, and a neotype is designated for H. parvula Holmberg. A key to the species, descriptions, distributional data, and illustrations are provided. Key words: Cleptoparasitic bees, Nomadini, Argentina, new species. Resumen: Revisión de las abejas del género Nomada en la Argentina (Hymenoptera, Apidae, Nomadinae). Se presenta una revisión de las abejas del género Nomada Scopoli en la Argentina. Se reconocen nueve especies en esta región, de las cuales cinco se describen como nuevas: N. mesopotamica, N. longula, N. chacoana, N. missionica y N. turrigera. Se designan lectotipos para N. pampicola Holmberg, 1886, y N. costalis Brèthes, 1909. Se propone un nuevo nombre, N. holmbergiana, para Hypochrotaenia parvula Holmberg, 1886, preocupado en Nomada, y se designa un neotipo para H. parvula Holmberg. Se presenta una clave para las especies, descipciones, datos de distribución e ilustraciones. Palabras clave: Abejas cleptoparásitas, Nomadini, Argentina, nuevas especies. ____________ INTRODUCTION Species of Nomada have been associated with hosts in several families of bees, but the majority The species of Nomada Scopoli belong in the of them belong to the genus Andrena Fabricius, an subfamily Nomadinae, a large and diverse group of association that may be ancestral for the genus bees all of which are cleptoparasites (Michener, (Alexander, 1991). The information on host asso- 2007). The genus Nomada is by far the most speciose ciation for the Neotropical species is extremely within the subfamily, with over 800 described spe- scanty, since hosts are only known for three spe- cies (Alexander & Schwarz, 1994). These bees occur cies in the Caribbean, which parasitize species of in all the continents, but are particularly abundant Exomalopsis Spinola and Agapostemon Guérin in the northern hemisphere. The South American (Alexander, 1991), and for one unidentified species fauna is rather poor in species, and a single group is of Nomada from Brazil, which parasitizes represented of the seventeen species-groups that Exomalopsis auropilosa Spinola (Rozen, 1997). The Alexander (1994) recognized for the world fauna. life histories of Argentinean species are yet to be This group has been variously treated in the past, as studied. the subgenus or genus Hypochrotaenia Holmberg Species of Nomada may be confused with spe- (Michener, 1954; Snelling, 1986), as the modesta cies of the related tribe Brachynomadini and allies, species-group (Alexander, 1991), or as the vegana which have a similar habitus and size. They can be species-group (Alexander, 1994). There is evidence easily distinguished because species of Nomada in that it is a monophyletic group (Alexander, 1994). South America always have yellow integumental The systematics of Nomada in South America has markings, while brachynomadines, also patterned received little attention and it is in a poor state of with red and black, completely lack yellow mark- knowledge, in spite of the reduced number of spe- ings, having instead bands and patches of white to cies present in the region. A recent contribution of yellowish pubescence. Schwarz & Gusenleitner (2004) has begun to rem- edy this situation with a detailed redescription of MATERIAL AND METHODS critical specimens. The present contribution pre- sents a revision of the nine species that occur in Specimens studied are deposited in the fol- Argentina, five of which are described as new. lowing institutions: Instituto y Fundación Miguel 222 Revista del Museo Argentino de Ciencias Naturales, n. s. 11 (2), 2009 Lillo, Tucumán (IFML); Museo Argentino de Species groups Ciencias Naturales «Bernardino Rivadavia,» There are four distinctive species-groups Buenos Aires (MACN); Museo de La Plata, La present in the Argentinean fauna. Two of them Plata (MLP); United States National Museum of agree with the delimitation proposed by Snelling Natural History, Washington D.C. (USNM); (1986) respectively for Hypochrotaenia s. str. and Zoologische Staatssammlung, München (ZSM). Micronomada Cockerell & Atkins. The other two Acronyms are used to indicate depositories of the are very distinctive small groups, and may prove to specimens. be derivatives from wihtin the former two. A phy- The following abbreviations are used: AOD, logenetic analysis of the species of Hypochrotaenia antennocular distance; IAD, interantennal dis- (the vegana group of Alexander, 1994) is badly tance; POD, post-ocellar distance; OOD, ocellocular needed, but is beyond the scope of this contribu- distance; CA, distance from anterior margin of tion. clypeus to lower tangent of antennal sockets; AMO, Nomada bonaerensis Holmberg, N. mesopo- distance from lower tangent of antennal sockets to tamica n. sp., N. costalis Brèthes, and N. holmber- lower tangent of median ocellus. The maximum giana n. n. belong in the group that Snelling (1986) diameter of the median ocellus (MOD) is used as characterized as Hypocrotaenia in a strict sense. It a reference to express the length of the pubes- is distinguished by the dorsal surface of the hind cence and other structures. The metasomal terga tibia with coarse, dark spiniform setae arising from (T) and sterna (S) are identified with Arabic nu- the spicules, and by the hind coxa laterally ex- merals. The sex of the specimens is indicated by panded, with the dorsal outer margin keeled or not. F, female, and M, male. The forecoxa bears an apical spine or at least a tooth-like projection. The forewing is darker along SYSTEMATICS the costal margin, or it is entirely infuscate with a clear preapical spot. Usually the males have the All the Neotropical species of Nomada are clas- sculpture of the integument stronger than that of sified in the subgenus Hypochrotaenia Holmberg, the females (Figs. 34-35). This group includes the the vegana species-group of Alexander (1994). largest number of Neotropical species of Nomada. Snelling (1986) defined Hypochrotaenia, which he Nomada pampicola Holmberg was included by treated at the genus level, by the following charac- Snelling (1986) in Micronomada, due to the ters, not taking into account those that he men- rounded hind coxa, and the feeble setae present on tioned as variable: gonocoxite with a digitiform in- the dorsal margin of the hind tibia. The pronotal ner dorsal lobe, forecoxa with an apical tooth or collar has rounded lateral angles, and the forecoxa spine, paraocular carina absent, mandible without bears an apical spine. It is the only species present preapical tooth, first flagelar segment at least as in Argentina of this group, which has its maximal long as second, lack of specialized subgenital brush, diversity in North America. gonostylus without specialized groups of setae, and Nomada longula n. sp. belongs in a group char- anterior margin of the pronotum subangulate to acterized by their wings with an amber membrane angulate laterally. The first three characters are and yellowish veins and pterostigma. All included synapomorphies of the monophyletic vegana spe- species have two submarginal cells due to the loss cies-group in the cladistic analysis of Alexander of the second abscissa of vein Rs. The face is broad, (1994), while the last character unites the vegana squarish, with the antennal sockets well above the with the roberjeotiana species-group (the last one middle of the face, the swollen area between the equivalent to the subgenus Nomadita Mocsáry). antennal sockets nearly reaching the middle ocel- The problems of recognizing subgenera within lus, and the upper paraocular areas also swollen. Nomada were stated by Alexander (1994), who re- The body is elongate and the color patterns remind frained from using them and divided the genus in those of some social vespids. The epithets coined 17 species-groups, two of them remaining as by Ducke (1908, Nomada polybioides) and by paraphyletic. It would be desirable to have a Cockerell (1916, Polybiapis mimus) refer to this subgeneric classification of Nomada, but this fact. According to Snelling (1986) N. abnormis seems not feasible until the taxonomy of its spe- Ducke is related to polybioides and mimus. Schwarz cies and the phylogenetic relationships are better & Gusenleitner (2004) have carefully redescribed understood. In any case, I have found useful to mimus. In this group, like in bonaerensis and al- recognize species-groups among the South Ameri- lies, the dorsal surface of the hind tibia has coarse, can species, which are not equivalent to dark spiniform setae arising from the spicules and Alexanders species-groups, but all fall within his the hind coxa is laterally expanded, usually with vegana group. the dorsal outer margin keeled. Roig Alsina: Revision of the bee genus Nomada 223 Nomada missionica n. sp. and N. turrigera n. pointed on the wing margin, missionica and sp. belong,
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