Zootaxa 3683 (5): 501–537 www.mapress.com / zootaxa / Copyright © 2013 Magnolia Press ISSN 1175-5326 (print edition) Article ZOOTAXA ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3683.5.1 http://zoobank.org/urn:lsid:zoobank.org:pub:E841BEF3-A5A9-4FD6-9262-15ABFB7FFF40 Revision of the bee subgenus Centris (Wagenknechtia) Moure, 1950 (Hymenoptera: Apidae: Centridini) FELIPE VIVALLO Laboratório de Hymenoptera, Departamento de Entomologia, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, São Cristóvão 20940-040 Rio de Janeiro, RJ, Brazil. E-mail: fvivallo@yahoo.com Table of contents Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 501 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 501 Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 502 Taxonomic history of the species of Centris (Wagenknechtia) Moure, 1950 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 503 Taxonomy of the species of Centris (Wagenkechtia) Moure, 1950 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 505 Centris (Wagenknechtia) Moure, 1950 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 505 Centris (Wagenknechtia) cineraria Smith, 1854 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 505 Centris (Wagenknechtia) escomeli Cockerell, 1926 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 509 Centris (Wagenknechtia) moldenkei Toro & Chiappa, 1989 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 511 Centris (Wagenknechtia) muralis Burmeister, 1876 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 512 Centris (Wagenknechtia) orellanai Ruiz, 1940 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 515 Centris (Wagenknechtia) rhodophthalma Pérez, 1911 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 517 Centris (Wagenknechtia) vardyorum Roig-Alsina, 2000 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 520 Key to the species of the subgenus Centris (Wagenknechtia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 521 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 529 Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 532 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 533 Abstract In this paper, a complete taxonomic revision of the species of Centris (Wagenknechtia) Moure is presented for the first time. The following species are recognized: Centris cineraria Smith, C. escomeli Cockerell, C. moldenkei Toro & Chiappa, C. muralis Burmeister, C. orellanai Ruiz, C. rhodophthalma Pérez and C. vardyorum Roig-Alsina. Floral associations, distribution records, and diagnoses of both sexes based on type specimens, are given. An identification key, illustrations, along with an updated catalogue of all species of the subgenus, are also provided. In addition, a neotype for Centris orellanai is designated. Key words: Oil-bees, taxonomy, Neotropical Region Introduction Centris Fabricius, 1804 is one of the most important and diverse genera of solitary bees of the Neotropical Region, occurring from southern United States to southern South America (Michener, 1979, 2000). The more than 200 described species are organized into 12 subgenera (Moure et al., 2007) widely distributed in the Neotropics, although some of the latter have more restricted distributions, especially in xeric areas of South America. Among the currently recognized subgenera, Centris (Wagenknechtia) Moure, 1950 is, along with C. (Penthemisia) Moure, 1950 (sensu Zanella, 2002) one with the lowest number of species described. This subgenus occurs mainly in dry areas of southern South America, from southern Peru, central Chile and Argentina to Accepted by C. Rasmussen: 6 Jun. 2013; published: 8 Jul. 2013 501 Patagonia. Centris cineraria Smith, 1854 is the species with the southernmost distribution records not only of the subgenus, but also of the entire tribe Centridini. In general, species of Centris (Wagenknechtia) are well known and their identification is relatively easy, due to the regional treatments of Roig-Alsina (2000) and Vivallo et al. (2002), who studied the species occurring in Argentina and Chile, respectively. Despite this, there has been neither a comprehensive taxonomic revision nor an identification key available for all known species of the subgenus. Although Centris (Wagenknechtia) contains only seven described species (Vivallo et al., 2002; Moure et al., 2007) a number of papers have been published since the beginning of the last century containing different aspects of the biology of various species of the subgenus. Especially numerous are papers dealing with floral and distribution records. However, just as occurs with the taxonomy of the group, there are no publications that encompass all available, updated and standardized information for all described species. This lack of organization has created some confusion, both in the taxonomy of some species, as well as in their floral and distribution records, such as for Centris cineraria, C. escomeli Cockerell, 1926 and C. vardyorum Roig-Alsina, 2000 (see for example Jörgensen, 1909, 1912a, b; Brèthes, 1910; Herbst, 1921; Wagenknecht, 1971). The misidentifications of these species have led to the erroneous association of various bionomic data that actually belong to other species. Certain biological aspects are relatively well known for some of the species of Centris (Wagenknechtia), such as flight period, plants visited, nest architecture, immature stages and natural enemies. These data are available for Centris cineraria mainly through the publications of Claude-Joseph (1926), Ruiz (1940), Rozen (1965) and Wagenknecht (1971), and for C. rhodophthalma Pérez, 1911 through the contributions made by Ruiz (1940), Wagenknecht (1971) and mainly by E. Chiappa, whose work has greatly increased our knowledge of the bionomics of this species (Chiappa, 1998, 2000; Chiappa & Rodríguez, 2001; Chiappa et al., 2000). As result of these contributions, C. rhodophthalma has become the best and most studied species of the entire subgenus. On the other hand, this contrasts with the situation in other species, such as C. escomeli, C. moldenkei Toro & Chiappa, 1989 and C. orellanai Ruiz, 1940, where many relevant aspects of biology remain unknown. In this paper, a complete taxonomic revision of all species of Centris (Wagenknechtia) is presented for the first time; diagnoses, floral associations and distribution records are provided for all species of the group. An identification key, illustrations of both sexes and an updated catalogue for all species of the subgenus are also provided. In addition, a neotype for Centris orellanai is designated. Material and methods The material studied is deposited in the following collections: American Museum of Natural History, New York, USA (AMNH), British Museum of Natural History, London, England (BMNH), California Academy of Sciences, California, USA (CAS), Central Texas Melittological Institute, Texas, USA (CTMI), Coleção Entomológica Pe. Jesus Santiago Moure, Departamento de Zoologia, Universidade Federal do Paraná, Curitiba, Paraná, Brazil (DZUP), Colección Entomológica de la Universidad de Tarapacá, Arica, Chile (IDEA), Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Buenos Aires, Argentina (MACN), Museo de Historia Natural de Valparaíso, Valparaíso, Chile (MHNV), Museo Nacional de Historia Natural, Santiago, Chile (MNHN), Muséum National d’Histoire Naturelle, Paris, France (MNHP), Museu Nacional, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil (MNRJ), Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (MZUSP), Laurence Packer Collection, York University, Toronto, Ontario, Canada (PCYU), Pontificia Universidad Católica de Valparaíso, Valparaíso, Chile (PUCV), Coleção Camargo, Departamento de Biologia, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de São Paulo, Ribeirão Preto, São Paulo, Brazil (RPSP), United States National Museum, Washington, USA (USNM) and Zoological Museum of the University of Copenhagen, Copenhagen, Denmark (ZMUC). General morphological terminology follows Michener (2000), mandible morphology is according to Michener & Fraser (1978) and the oil-collecting apparatus (elaiospathe) according to Neff & Simpson (1981) and Snelling (1984). Antennal flagellomeres are indicated as F1, F2, etc.; metasomal terga and sterna as T1, T2, S1, S2, etc., respectively. All the measurements are given in millimeters (mm). The position of vertex in relation to eyes was considered in frontal view. The upper interocular distance (UID) was measured considering the shortest distance between the eyes, in frontal view. The lower interocular distance (LID) was measured at the same level of the 502 · Zootaxa 3683 (5) © 2013 Magnolia Press VIVALLO maximum clypeal width. The length of the antennal flagellomeres was measured along their mid line. Mandibular length was measured from the acetabulum to the apex of the apical tooth. The maxillary palpomeres were numbered from the base to the apex of the palpus. The measurements of Centris muralis correspond to the average of four specimens of each sex randomly selected from a sample of 50 specimens. The labels were transcribed under the section “Material examined”. The backward slash (\) indicates different labels on the pin of the same specimen. Misspelled names were given correctly between square brackets as “[=...]”. The distribution maps were based on locality records taken from specimen labels and from records available in the literature listed in the section “References”. The erroneous records from misidentifications of species were not considered. This also was done for the compilation of floral records (Table 1). All new distribution and/ or floral records were marked with an asterisk (*). In the sections “Distribution” and “Material examined” the countries, their main geopolitical divisions (Regions in Chile, Departments in Peru and Provinces in Argentina) and the localities within them were ordered alphabetically, when possible. Chile is divided into 15 Regions designated by a name and a Roman numeral, assigned from north to south. Nevertheless there are three exceptions: the Metropolitan Region (designated RM and informally recognized with the number XIII), and the recently created Regions of Los Ríos in the south (XIV), and Arica y Parinacota in the north (XV). In this paper were used both nomenclatures (name and Roman numeral), because it is possible to find in the literature or in the labels of the specimens the name of the Region, its Roman numeral or both simultaneously. The distribution maps were created using ArcGis software (10.1 version). Photographs were taken using a Leica DFC 500 camera attached to a Leica MZ16 stereomicroscope. All images were first treated using Auto-Montage Pro (Syncroscopy) software of the Projeto Taxonline, Rede Paranaense de Coleções Biológicas of the Universidade Federal do Paraná (UFPR), Curitiba, Paraná, Brazil. Later, they were enhanced with Adobe Photoshop® (ver. 7.0) without distorting the morphological characters of the specimens. The images of the oil-collecting structures were taken using a scanning electron microscope ZEISS DSM 940 in the Laboratório de Microscopia Eletrônica of the Instituto de Biociências of the Universidade de São Paulo (USP), São Paulo, Brazil. In the discussion about biogeography the hierarchization of biogeographic units proposed by Morrone (2001) was used, where the Notropical Region is divided into subregions and these in turn into biogeographic provinces recognized by characteristic groups of organisms. For a complete list of taxonomical groups see references therein. The literature cited below each species corresponds to an update of the information presented by Moure et al. (2007) in the Catalogue of Bees in the Neotropical Region. Taxonomic history of the species of Centris (Wagenknechtia) Moure, 1950 The taxonomic history of the species of Centris (Wagenknechtia) began in the mid-nineteenth century with the publication of Smith (1854) on the Hymenoptera of the British Museum. This author described several species of Neotropical bees, one of them being Centris cineraria, based on a single female collected in Chile. Burmeister (1876) studied a series of one female and three males collected in Argentina, describing them as Centris muralis and also publishing notes on some other species of the genus that occur in that country. Gribodo (1893) described Anthophora plumigera and A. virgo from a female and a male, respectively, collected in Patagones, Argentina. Three years later, Dalla Torre (1896) published a catalogue of Hymenoptera, citing Centris cineraria, C. muralis and transferring the species described by Gribodo (1893) to the genus Podalirius Latreille, 1802, citing them as P. plumiger and P. virgo. Friese (1900) published a monograph of the genus Centris, proposing the subgenera C. (Cyanocentris), C. (Rhodocentris), C. (Poecilocentris) and C. (Melacentris), including C. cineraria in the first and C. muralis in the last. Schrottky (1902a) published a list of species deposited in the National Museum of Buenos Aires listing Centris (Melanocentris) muralis for the central region of Argentina and C. cineraria, as C. (Cyanocentris) chilensis (Spinola, 1851), for the south of that country. Schrottky (1903) reported on both species with new distribution records in Argentina. Friese (1908) described Centris muralis melanopus, based on a number of males and females collected in Mendoza, Argentina. That year, Jensen-Haarup (1908) reported for the first time C. nigerrima (Spinola, 1851) in Argentina, information that was subsequently referred to by Friese (1908, 1910). Brèthes (1910) published a short list of Hymenoptera of Chile, proposing Centris smithii Friese, 1899 (= C. chilensis) as a junior synonym of C. cineraria, due to the confusion in the identity of both species. That year Friese REVISION OF THE BEE SUBGENUS CENTRIS (WAGENKNECHTIA) Zootaxa 3683 (5) © 2013 Magnolia Press · 503 (1910) also published a list of bees from southern Argentina, listing Centris chilensis in Neuquén and C. nigerrima in Mendoza. This author also reported this latter species in Concepción, Chile. In the next year Centris rhodophthalma Pérez, 1911, was described from a single female collected in Chañarcillo, northern Chile. Friese (1912) published a list of bees from southern Argentina citing among them C. muralis melanopus. In the same year Jörgensen (1912b) published a revision of the bees from Mendoza (Argentina), transferring the species of Centris to Hemisia Klug, 1807 citing Hemisia nigerrima, H. muralis and H. muralis melanopus. Schrottky (1913a) published the geographical distribution of Argentinian Hymenoptera, and as proposed by Jörgensen (1912b), reported Hemisia muralis and H. nigerrima in central Argentina, and H. chilensis in Patagonia. In 1919 Cockerell published a list of the species deposited in the United States National Museum, listing Centris cineraria as C. chilensis, apparently following the interpretation given by Brèthes (1910) for this species. Along with this species Cockerell (1919) also listed C. muralis and C. muralis melanopus. In 1921 Herbst discussed the real identity of Centris cineraria, indicating that the interpretation of Brèthes (1910) on this species was erroneous. Subsequently, Friese (1924) described several new species and subspecies of Centridini, including Centris chilensis neoqueenensis, based on a number of males and females collected in Neuquén, Argentina, all apparently in poor condition. Cockerell (1926) published a small work on Peruvian bees, describing Centris escomeli, from a single male collected in Arequipa, Peru. In 1940 Ruiz published one of the first revisions of the bees of Chile, giving a brief overview of taxonomy, biology, floral association and distribution. Ruiz reported on Centris cineraria and C. rhodophthalma and described C. nigerrima orellanai, based on a female collected in northern Chile. In 1950 Moure proposed some new groupings of Neotropical bees, describing the subgenus Centris (Wagenknechtia), with C. cineraria as the type species, C. rhodophthalma, and C. orellanai, recognized as valid species and not as subspecies of C. nigerrima, and, tentatively, C. muralis. In that paper Moure (1950) indicated that the interpretation given by Friese (1924) for C. chilensis corresponds to C. cineraria, following Herbst (1921). Michener (1951) recognized Hemisia as the valid name for Centris and published a key to the subgenera described thus far, including in the subgenus Hemisia (Wagenknechtia) the species H. cineraria, H. muralis, H. orellanai and H. rhodophthalma. Moure (1960) re-recognized Centris as the valid name of the genus, proposing the synonymy of Anthophora plumigera and A. virgo with C. muralis, based on type material of all species, choosing a male of the type series of the latter species as lectotype. In 1971 Wagenknecht published the fifth part of his contribution to the biology of Chilean bees, providing bionomics, distribution records and floral associations of several species of Centris, citing in C. (Wagenknechtia) the species C. cineraria, C. muralis, C. orellanai and C. rhodophthalma. Snelling (1974) dealt with the taxonomy and distribution of North American species of Centris, listing in C. (Wagenknechtia) the species C. cineraria, C. muralis, C. orellanai and C. rhodophthalma. Toro (1986b) published a preliminary list of Chilean bees, including in C. (Wagenknechtia) the species C. cineraria, C. escomeli, C. orellanae [sic], C. rhodophthalma, and C. cineraria neoqueenensis, which was reported for the first time from Chile, this time as a subspecies of C. cineraria and not of C. chilensis as originally described by Friese (1924). Toro & Chiappa, 1989 described Centris moldenkei, a species endemic to northern Chile. In 1998 Chiappa redescribed the female and described the male of Centris rodophthalma [sic], based on material collected in the IV Coquimbo Region, Chile. This author also provided a key to the species of C. (Wagenknechtia) known at that time to occur sympatrically with C. rhodophthalma: C. cineraria and C. orellanai. Roig-Alsina (2000) reviewed the Argentinian species of Centris, describing C. vardyorum and providing distribution records for C. muralis and C. cineraria. He also indicated that the data published by Friese (1908) and Jörgensen (1909) on C. nigerrima, and by Jörgensen (1912a, b) and Schrottky (1913a) on Hemisia nigerrima actually refer to C. vardyorum. Additionally, Roig-Alsina (2000) proposed C. muralis melanopus and C. chilensis neoqueenensis as junior synonyms of C. muralis and C. cineraria, respectively. In that paper, Roig-Alsina (2000) also described C. hyptidoides, which he tentatively included in C. (Wagenknechtia). However, this species belongs to a different lineage, as indicated by Vivallo & Melo (2009). Chiappa et al. (2000) published some notes on the biology of Centris rodophthalma [sic] comparing them with the published data on other species of the genus that occur in Chile. These authors listed in C. (Wagenknechtia) the species C. rodophthalma [sic] and C. cineraria. Vivallo et al. (2002) partially reviewed the species of C. (Wagenknechtia), providing diagnoses and biological data for the Chilean species of the subgenus, along with an identification key. These authors recognized in this subgenus Centris cineraria cineraria, C. cineraria neoqueenensis, C. escomeli, C. moldenkei, C. muralis, C. orellanai, C. rhodophthalma and C. vardyorum. This paper reported for the first time that the data published by Wagenknecht (1971) and Toro (1986b) for Centris garleppi (Schrottky, 1913b) actually pertain to C. escomeli. The last taxonomic work published about 504 · Zootaxa 3683 (5) © 2013 Magnolia Press VIVALLO the species of Centris (Wagenknechtia) was the Catalogue of Bees in the Neotropical Region (Moure et al., 2007). In that paper C. cineraria, C. escomeli, C. moldenkei, C. muralis, C. orellanai, C. rhodophthalma and C. vardyorum were recognized. Moure et al. (2007) also indicated that distribution records and floral association published by Jensen-Haarup (1908) and Friese (1910) for C. nigerrima refer to C. vardyorum. Taxonomy of the species of Centris (Wagenkechtia) Moure, 1950 Centris (Wagenknechtia) Moure, 1950 Centris (Wagenknechtia); Snelling, 1956. 32: 5 (Systematic note). Anderson, 1979. 11: 222 (Bionomics). Neff & Simpson, 1981. 54: 106, 107, 118 (Morphology, distribution, note on phylogeny, comparison with Centris (Paracentris) Cameron, 1903 and C. (Melacentris) Moure, 1996). Coville et al., 1983. 56: 116, 117, 121 (Cited, systematic note, nesting behaviour). Snelling & Brooks, 1985. 369: 23 (Cited). Coville et al., 1986. 59: 325 (Cited). Buchmann, 1987. 18: 355 (Descriptive note, floral records, bionomics). Vogel, 1988. 26: 348, 350 (Figure about phylogeny within Centridini, cited). Cocucci, 1991. 174: 31 (Cited). Chiappa, 1998. 22: 90 (Key). Chiappa et al., 2000. 24: 24 (Systematic note). Michener, 2000: 732−735 (Cited, key, distribution, comparison with Centris (Paracentris)). Fernández, 2001. 2: 109 (List, distribution). Ayala, 2002. 25: 2 (Cited). Fernández, 2002. 2: 129 (List, distribution). Machado et al., 2002. 4: 357 (Descriptive note). Silveira et al., 2002: 93 (Cited). Vivallo et al., 2003: 78 (Cited, systematic note). Machado, 2004: 258 (Descriptive note, floral record). Rocha-Filho et al., 2009. 26: 302 (Cleptoparasites). Cappellari et al., 2011. 107: 1331 (Morphology, behaviour). Cilla & Rolón, 2012b. 67: 573 (Bionomics). Giannini et al., 2013. 258: 76, 79 (Floral relationships, phylogenetic relationships within Centris). Diagnosis. The species of this subgenus are identified by a combination of the following characters (both sexes): head and mesosoma with integument dark brown to black, never with yellow areas (Figs. 1, 3, 5, 7, 9, 11, 13, 15, 17, 19, 21, 23, 25, 27); metasoma with metallic reflections (in some species) (Figs. 2, 4, 18, 20, 22, 24, 26, 28, 35 and 36), wing membranes translucent (Figs. 2, 4, 6, 8, 10, 12, 14, 16, 18, 20, 22, 24, 26, 28) and maxillary palpus with five palpomeres. Females: basitibial plate with a concavity in the lower half of the primary plate (Fig. 35) (except in C. muralis Fig. 37), with secondary plate fused with the primary plate, evidenced by a convexity on the upper half (lateral view). Males: preapical surface of gonocoxite with a papillate projection (Fig. 38). Centris (Wagenknechtia) cineraria Smith, 1854 (Figures 1−4, 29, 30, 38 and 39) Centris cineraria Smith, 1854. 2: 378 (Original description). Herbst, 1917. 1917: 265 (Cited). Etcheverry & Valenzuela, 1960. 1: 51 and plates N°11 and N°13 (Morphology, distribution, floral records, nesting behaviour, nest architecture, bionomics, figures of nest and larvae). Herrera & Etcheverry, 1960. 1: 64 (List). Vogel, 1974. 7: 124, 152, 155, 211, 230, 245−247 (Distribution, floral records, figure of an adult, bionomics). Batra & Schuster, 1977. 9: 136, 137 (Distribution, nesting behaviour, bionomics). Kalin et al., 1982. 69: 93−96 (Distribution, floral records). Snelling & Brooks, 1985. 369: 19 (Cleptoparasites). Toro, 1986a. 57: 83 (Distribution, bionomics). McGinley, 1989. 494: 12 (List). Toro et al., 1991. 16: 98 (Cited). Rozzi et al., 1997. 132: 174, 176 (Floral record, cited). Camousseight & Barrera, 1998. 25: 78−80 (Distribution, list, cited, floral records). Chiappa, 1998. 22: 90 (Key). Chiappa, 2000. 46: 19, 23 (Cited, descriptive note of cell and morphology of larva). Chiappa et al., 2000. 24: 20−24 (Cited, nesting behaviour, distribution, bionomics, nest architecture, floral records, cleptoparasite, comparison with Centris rhodophthalma). Alonso-Amelot, 2008. 34: 888 (Floral record, bionomics). Montalva & Ruz, 2010. 35: 45 (List, distribution). Cilla & Rolón, 2012b. 67: 580 (Nest architecture, distribution). Cosacov et al., 2012. 47: 465, 367 (Floral record, bionomics, cited). Espinoza et al., 2012. 85: 506 (Floral record). Rolón & Cilla, 2012. 66: 37 (Nesting biology). Centris (Wagenknechtia) cineraria; Vogel, 1974. 7: 151 (Cited). Magunacelaya, 1981. 14: 226 (Cited, systematic note). Neff & Simpson, 1981. 54: 105−107 (Descriptive note, figure of fore elaiospathe). Coville et al., 1983. 56: 117, 118, 120 (Nesting behaviour, nest architecture). Toro, 1986b. 13: 127 (Distribution). Rocha-Filho et al., 2009. 26: 301 (Cleptoparasite). Centris chilensis neoqueenensis Friese, 1924. 3: 21. Centris (Wagenknechtia) cineraria neoqueenensis; Toro, 1986b. 13: 127 (Distribution). Chiappa, 1998. 22: 91 (Distribution). Vivallo et al., 2003: 81 (Distribution). Hemisia cineraria; Michener, 1953. 35: 1072 (Descriptive note of larva). Comments. A complete redescription of this species can be found in Vivallo et al. (2002), so here are indicated only diagnostic characters for the identification of both sexes of this species. REVISION OF THE BEE SUBGENUS CENTRIS (WAGENKNECHTIA) Zootaxa 3683 (5) © 2013 Magnolia Press · 505 Diagnosis. Both sexes with integument dark brown on head and mesosoma. Metasoma with bluish metallic reflections. Mesoscutum and scutellum with whitish or light yellow hairs. The rest of the body with dark brown to black pilosity. Female: fore elaiospathe located in the distal half of forebasitarsus, with the secondary comb formed by three giant spatulate hairs (Fig. 29). Hairs of hind elaiospathe not overlapping (Fig. 30). Middle and hind claws simple, fore claws with small preapical tooth. The specimens with southernmost distribution records have slightly yellow pubescence on mesoscutum and scutellum. This species is similar to Centris orellanai, differing (both sexes) in the coloration of the pubescence on mesoscutum and scutellum (relatively whitish in C. cineraria (Figs. 1−4); light brown to orange in C. orellanai (Figs. 17−20)) and by the metallic reflections on metasoma (bluish in C. cineraria (Figs. 2 and 4); relatively greenish in C. orellanai (Figs. 18 and 20)). Both females also differ in the hair number of the secondary comb of the fore elaiospathe (three in C. cineraria; four in C. orellanai), and in the preapical tooth of the claws of middlegs (absent in C. cineraria; present in C. orellanai). Type material. Centris cineraria: Holotype female (BMNH, examined). Centris chilensis neoqueenensis: Syntypes male and female (Museum für Naturkunde, Humboldt-Universität zu Berlin, Germany (ZMB), not examined). Type locality. Centris cineraria: Chile. Centris chilensis neoqueenensis: Argentina: Neuquén. Taxonomic decision for synonymy. Roig-Alsina (2000). Floral records. See table 1. FIGURES 1–4. Centris cineraria Smith, 1854. Fig. 1. Female (Viña del Mar, Chile): Head, frontal view. Fig. 2. Habitus, lateral view. Fig. 3. Male (Viña del Mar, Chile): Head, frontal view. Fig. 4. Habitus, lateral view. Scale bars 1 mm. 506 · Zootaxa 3683 (5) © 2013 Magnolia Press VIVALLO Distribution. This species ranges in Chile from IV Coquimbo Region as far south as XII Magallanes y la Antártica chilena Region in Chilean Patagonia, and in Argentina from Neuquén Province to Santa Cruz Province. ARGENTINA: (Herbst, 1921). CHUBUT: (Cockerell, 1919; Ruiz, 1940; Wagenknecht, 1971; Montalva & Ruz, 2010). Parque Nacional Los Alerces (Roig-Alsina, 2000). *Esquel. NEUQUÉN: (Friese, 1910; Ruiz, 1940; Wagenknecht, 1971; Vogel, 1974; Vivallo et al., 2002, 2003; Montalva & Ruz, 2010). Fortín Chacabuco, Ñirihuau (Vogel, 1974), San Martín de Los Andes (Roig-Alsina, 2000). Nahuel Huapi (Vogel, 1974). *Cabecera E. lago Huenchulafquén. RÍO NEGRO: (Montalva & Ruz, 2010). Alto Ñiriguao, Bariloche (Roig-Alsina, 2000). SANTA CRUZ: (Holmberg, 1884, 1903; Schrottky, 1902a, 1903; Montalva & Ruz, 2010). Perito Moreno (Roig-Alsina, 2000). *Lago Buenos Aires. CHILE: (Dalla Torre, 1896; Friese, 1910; Moure, 1950; Batra & Schuster, 1977; Toro, 1986a; Chiappa & Toro, 1994; Rozzi et al., 1997; Montalva & Ruz, 2010). IX ARAUCANÍA: Angol (Cockerell, 1937), Lonquimay (Ruiz, 1940; Wagenknecht, 1971; Vivallo et al., 2002). XI AYSÉN DEL GENERAL CARLOS IBÁÑEZ DEL CAMPO: Chile Chico (Toro, 1986b; Chiappa, 1998; Vivallo et al., 2002). VIII BIO-BÍO: Mulchén (Ruiz, 1940; Wagenknecht, 1971). *Las Trancas, *Quenco, *Termas de Chillán. IV COQUIMBO: (Chiappa et al., 2000a). Illapel (Wagenknecht, 1939, 1971; Vivallo et al., 2002). Valle del Choapa, Vicuña (Ruiz, 1940). *Cuncumén, *Las Breas, *Puerto Oscuro. VI LIBERTADOR GENERAL BERNARDO O’HIGGINS: Termas del Flaco. XII MAGALLANES Y LA ANTÁRTICA CHILENA: *Laguna Amarga. VII MAULE: Fundo Bellavista (Ruiz, 1940). Talca (Hacienda Las Mercedes) (Gazulla, & Ruiz, 1928; Ruiz, 1940; Wagenknecht, 1971). Reserva Nacional Los Ruiles (Espinoza et al., 2012), *Curicó, *La Jaula, *Los Queñes. REGIÓN METROPOLITANA: La Obra (Wagenknecht, 1971). La Parva (Kalin et al., 1982; Camousseight & Barrera, 1998). Las Condes (Wagenknecht, 1971). Lo Águila (Ruiz, 1940). Maipú (Cockerell, 1919, probable record). Santiago (Brèthes, 1910; Ruiz, 1923, 1940; Etcheverry & Valenzuela, 1960; Vogel, 1974; Cilla & Rolón, 2012b). *Cajón del Maipo, *Caleu, *El Canelo, *El Manzano, *Farellones, *Lagunillas, *Lo Marín, *Lo Prado, *Los Pinos, *Manquehue, *Quebrada de Macul, *Quebrada del Peumo, *Tiltil, *Vega de Las Vacas. V VALPARAÍSO: Aconcagua (Ruiz, 1940). Cerro La Campana (Vivallo, 2000). Limache (Ruiz, 1940), Casablanca, Ocoa, Rocas de Santo Domingo, Viña del Mar (Wagenknecht, 1971). *Belloto, *Chacabuco, *Colliguay, *Concón, *Cuesta La Dormida, *El Salto, *Horcón, *Los Andes, *Montemar, *Papudo, *Peñuelas, *Piedras Blancas, *Quilpué, *Recreo, *Reñaca, *Río Blanco, *Río Colorado, *Valparaíso, *Villa Alemana, *Zapallar. Material examined. ARGENTINA: CHUBUT: 1 female: Argentina Chubut 24 km N. of Esquel 42°51’209” 071°07’507” 2700ft. 20.XII.06 A.-I. Gravel (PCYU). NEUQUÉN: 1 female and 1 male: 1627\ Argentina: Neuquén: Cabecera E. lago Huechulafquén, sobre Adesmia sp. leg. G. Germán Roitman & Norberto H. Montaldo 11/12/1997\ Centris cineraria Smith, 1854 Roig Alsina det. 2000 (MACN). 1 female: 10266\ Neuquén\ Centris cineraria Smith, 1854 Roig Alsina det. 2000 (MACN). RÍO NEGRO: 2 females: R. A. Río Negro Bariloche 12/ 1964 A. Giai\ Centris cineraria Smith, 1854 Roig Alsina det. 2000 (MACN). 1 male: Río Negro Alto Ñirihuao 03/ 01/1964\ Centris cineraria Smith, 1854 Roig Alsina det. 2000 (MACN). SANTA CRUZ: 1 female: Santa Cruz P. Moreno lago Bs. Aires 12/1982 J. Carreras\ Centris cineraria Smith, 1854 Roig Alsina det. 2000 (MACN). CHILE: 1 female: Holotype\ B.M.Type Hym 17B.925\ Chili\ cineraria Type Sm.\ Centris cineraria Type Smith (BMNH). IX ARAUCANÍA: 1 female: VIII R. Angol 05/11/1985 Chiappa (PUCV). 1 female: VIII R. Angol 05/11/ 1985 De la Hoz (PUCV). 1 male: VIII R. Angol 05/11/1985 C. Chaparro (PUCV). 1 female: Malleco Angol 26/11/ 1980 De la Hoz (PUCV). 1 female: Angol, 23/11/1944 J. Daúdelh (MNHN). XI AYSÉN DEL GENERAL CARLOS IBÁÑEZ DEL CAMPO: 1 male and 1 female: Chile Chico Aysén Chile 27/12/1960 L. E. Peña (DZUP). VIII BIOBÍO: 1 female: Termas de Chillán 27/01/1977 M. Cerda, col\ Coll. Cerda/ MNHN Chile (MNHN). 1 male and 2 females: Chillán Las Trancas 15/12/1977 O. Martínez (PUCV). 1 male: VIII R. Quenco 07/11/1985 O. Martínez (PUCV). 1 male: VIII R. Quenco 07/11/1985 De la Hoz (PUCV). IV COQUIMBO: 1 male: IV R. Cuncumén 13/ 09/1985 C. Chaparro (PUCV). 1 female: Coquimbo Las Breas 17/10/1979 Magunacelaya (PUCV). 1 male: IV R. Pto. Oscuro 09/1986 C. Tobar (PUCV). 2 females: Illapel 19/10/1933 (MNHN). XII MAGALLANES Y LA ANTÁRTICA CHILENA: 3 females: Laguna Amarga Magallanes Chile diciembre 1960. L. Peña (MNHN). VII MAULE: 1 male: Curicó La Jaula 11/1991 H. Toro (PUCV). 3 males: Curicó Los Queñes 31/10/1983 F. Rodríguez (PUCV). 1 male: Curicó Los Queñes 11/1991 Toro (PUCV). VI LIBERTADOR GENERAL BERNARDO O’HIGGINS: 1 female: VI R. Termas del Flaco 05/12/1982 H. Flores (PUCV). 2 males and 1 female: VI R. Termas del Flaco 05/12/1982 B. Dyer (PUCV). 4 males and 1 female: VI R. Termas del Flaco 05/12/1982 Magunacelaya (PUCV). REGIÓN METROPOLITANA: 1 female: 17.462\ Centris chilensis 1904 ♀ Friese det\ Chile Santiago 1890 Philippi\ 97292\ Chile Santiago 1890 Philippi col. 3 #MZ 17 962\ Centris (Wagenknechtia) cineraria Smith, REVISION OF THE BEE SUBGENUS CENTRIS (WAGENKNECHTIA) Zootaxa 3683 (5) © 2013 Magnolia Press · 507 1854 F. Zanella det. 1999 (DZUSP). 1 female: Santiago Chile\ Centris cineraria Sm ♀ det D. B. Baker 1980 (BMNH). 1 male: Santiago Los Pinos 04/10/1970 M. Cerda, col\ Coll. Cerda/ MNHN Chile (MNHN). 1 male: Santiago Quebrada del Peumo 21/10/1966 T. Ramírez, col (MNHN). 1 male: Santiago Manquehue 05/10/1970 M. D. E., col (MNHN). 1 male: Santiago Lo Prado 07/01/1996 M. Cerda, col\ Coll. Cerda/ MNHN Chile (MNHN). 5 females: Santiago C° San Cristóbal 01/11/1952 (MNHN). 2 males and 1 female: Santiago Quebrada de Macul 12/ 10/1988 M. Cerda, col\ Coll. Cerda/ MNHN Chile (MNHN). 1 male: Santiago Apoquindo 09/10/1966 R. Pérez, col (MNHN). 2 males: Santiago V. Paulina 04/11/1981 O. Martínez (PUCV). 1 male: Santiago Cajón del Maipo 11/ 1970 L. Alfaro, col (MNHN). 1 male: Santiago La Obra 19/09/1991 M. Cerda, col\ Coll. Cerda/ MNHN Chile (MNHN). 1 male: Santiago La Obra 11/1968 T. Ramírez, col (MNHN). 2 males: Santiago La Obra 09/1968 T. Ramírez, col (MNHN). 1 male: Santiago El Manzano 21/10/1970 M. Cerda, col\ Coll. Cerda/ MNHN Chile (MNHN). 1 male: Santiago El Canelo 27/10/1970 M. Cerda, col\ Coll. Cerda/ MNHN Chile (MNHN). 1 female: Santiago El Canelo 15/10/1933 Ureta, col (MNHN). 1 female: Santiago Lagunillas 09/10/1969 Moroni, col (MNHN). 1 male: Santiago La Parva Vega de las Vacas 3000 msnm. 5-7/01/1979 A. Camousseight, leg (MNHN). 1 female: Farellones 25/09/1979 P. Uslar\ en Valenzuelia trinervis (PUCV). 2 females: Farellones 15/12/1979 M. Arroyo\ en Amarthophillum cumingii (PUCV). 1 female: Farellones 18/12/1979 M. Arroyo\ en Sisyrinchium philippii (PUCV). 1 female: Farellones 19/12/1979 M. Arroyo\ en Adesmia radicifolia (PUCV). 1 female: Farellones 21/12/1979 M. Arroyo\ en Sisyrinchium philippii (PUCV). 1 female: Farellones 27/12/1979 M. Arroyo\ en Adesmia conferta (PUCV). 1 female: Farellones 27/12/1979 M. Arroyo\ en Astragalus curvicaulis (PUCV). 1 female: Farellones 05/01/1980 M. Arroyo\ en Adesmia conferta (PUCV). 1 female: Farellones 05/01/1980 M. Arroyo\ en Astragalus curvicaulis (PUCV). 3 males: Farellones 06/01/1980 M. Arroyo\ en Astragalus curvicaulis 2750 (PUCV). 2 females: Farellones 14/01/1980 M. Arroyo\ en Adesmia conferta (PUCV). 2 males: Farellones 24/ 01/1980 M. Arroyo\ en Astragalus curvicaulis 2750 (PUCV). 2 males and 1 female: Farellones (La Parva) 1970-72 Moldenke (PUCV). 1 female: 27646\ Chile, Santiago La Parva site. Int. Biol. Program 1970-72 leg. A. R. Moldenke #refers to host & date (PUCV). 1 male: Caleu Lo Marín 30/09/1984 M. Cerda, col\ Coll. Cerda/ MNHN Chile (MNHN). 1 female: Tiltil Caleu 21/10/1986 M. Cerda, col\ Coll. Cerda/ MNHN Chile (MNHN). 2 females: Caleu El Roble 07/11/1971 Sielfeld, col\ Coll: Sielfeld/ MNHN Chile (MNHN). 2 males: Chile Region Metropolitano Cerro Roble nr. Til-Til 17.xi.01 L. Packer (PCYU). V VALPARAÍSO: 1 female: Papudo- Zapallar Moldenke, leg (MNHN). 1 male: Papudo 17/10/1957 G. Kuschel, leg (MNHN). 1 male: Papudo Zapallar Moldenke (PUCV). 1 female: Horcones 25/10/1964 De la Hoz (PUCV). 6 males: Reñaca 27/09/1964 H. Toro (PUCV). 1 male: Reñaca 11/1964 Dazarola (PUCV). 1 male and 1 female: Reñaca 11/1964 J. Rocca (PUCV). 1 female: Montemar 10/1957 (MHNV). 1 male: Montemar 09/1957 (MHNV). 1 female: Recreo 24/10/1965 Solervicens (PUCV). 4 males and 2 females: Viña del Mar Salinas 21/10/1951 M. Cerda, col\ Coll. Cerda/ MNHN Chile (MNHN). 1 male: Viña del Mar 10/1965 (MHNV). 5 males: El Salto 18/10/1962 N. Hiching (PUCV). 2 males: El Salto 09/1957 (MHNV). 1 male: Valparaíso 07/11/1958 H. Toro G. (PUCV). 1 male: Valparaíso 09/11/1958 H. Toro G. (PUCV). 1 male: Valparaíso Reed (MNHN). 2 females: Peñuelas 01/11/1963 E. Cruzat P. (PUCV). 1 male and 3 females: Cº Roble Moldenke (PUCV). 1 female: 35189\ Chile Pr. Valparaíso & Santiago Cerro Roble Int. Biol. Program site leg. A. R. Moldenke #refers to host & date (PUCV). 1 male: Concón 05/10/1958 H. Toro G. (PUCV). 1 male: Piedras Blancas 12/10/1965 Solervicens (PUCV). 1 male and 1 female: Colliguay 10/1964 J. Rocca (PUCV). 3 males: Colliguay 18-20/09/1970 F. Rojas, col (MNHN). 1 male: Colliguay 15/09/1970 (MNHN). 4 males: Colliguay 10/1964 De la Hoz (PUCV). 1 male: Colliguay 18/10/1964 Chiappa (PUCV). 1 male: Colliguay 18/10/1964 V. Cabezas (PUCV). 3 males: Colliguay 18/10/1964 Rojas (PUCV). 6 males: Colliguay 18/10/1964 H. Toro (PUCV). 1 male: Colliguay 25/10/1964 L. Ruz (PUCV). 2 males: Cc\ Chile Region V Colliguay Los Yuyos 6.X.00 L. Packer (PCYU). 1 male: Quilpué 09/1960 M-Klug D. (PUCV). 1 Male: Quilpué 10/1960 M. Guzmán (PUCV). 1 male: Quilpué 20/09/1980 A. Schwalm (PUCV). 1 male: Quilpué 08/10/1967 L. Ruz (PUCV). 2 males: Quilpué 08/10/1967 Dazarola (PUCV). 1 male: Quilpué 09/1957 (MHNV). 1 male: Villa Alemana 01/10/1961 Solervicens (PUCV). 1 male: Villa Alemana 17/10/1963 J. Solervicens (PUCV). 1 male: Villa Alemana 21/10/ 1963 J. Solervicens (PUCV). 1 male: Belloto 10/1963 Dazarola (PUCV). 1 male: Belloto 25/09/1964 M. Aravena (PUCV). 7 males and 3 females: Cuesta La Dormida Moldenke (PUCV). 1 male: Cuesta La dormida Moldenke, leg (MNHN). 1 male: Limache 14/12/1925 F. Ruiz, col (MNHN). 2 females: Limache 02/01/1935 (MNHN). 1 male: Los Andes Río Colorado 2300 msnm. 01/11/1971 Sielfeld, col\ Coll: Sielfeld/ MNHN Chile (MNHN). 1 male: Los Andes 18/08/1958 (PUCV). 1 male: Los Andes 16/09/1958 (PUCV). 4 males: Los Andes 17/09/1958 (PUCV). 3 males: Los Andes 18/09/1958 H. Toro G. (PUCV). 1 male: Los Andes 19/09/1958 R. de Toro (PUCV). 1 male: Los 508 · Zootaxa 3683 (5) © 2013 Magnolia Press VIVALLO Andes 17/11/1958 R. de Toro (PUCV). 1 male: Los Andes Chacabuco 09/1959 (MHNV). 1 male: Río Blanco 12/ 1969 Montenegro (PUCV). 2 males: Río Blanco 03/10/1975 E. De la Hoz (PUCV). Centris (Wagenknechtia) escomeli Cockerell, 1926 (Figures 5−8 and 40) Centris escomeli Cockerell, 1926. 59: 28, 29 (Original description). Simpson, 1989. 14: 409, 410 (Cited). Toro et al., 1996. 23: 68, 69, 74 (Distribution, floral record). Chiappa, 1998. 22: 91 (Distribution). Montalva & Ruz, 2010. 35: 45 (List, distribution). Centris (Wagenknechtia) escomeli; Toro, 1986b. 13: 127 (Distribution). Cocucci et al., 1996. 34: 191 (Probable floral record). Vivallo et al., 2003: 78 (Distribution). Centris (Penthemisia) garleppi; Wagenknecht, 1971. 4: 279, 280 (Morphology, comparison with Centris nigerrima, distribution, floral record (misidentification)). Toro, 1986b. 13: 127 (List, distribution (misidentification)). Comments. A complete redescription of the male and the description of the female of this species can be found in Vivallo et al. (2002), so here are indicated only diagnostic characters for the identification of both sexes of this species. Diagnosis. Both sexes with blackish integument and brown pilosity (Figs. 5–8). Female: fore elaiospathe located in the distal half of forebasitarsus. Hairs of hind elaiospathe almost overlapping. All claws with preapical tooth. Male: mandible long and slender, with the posterior margin of second and anterior edge of the third tooth broadly curved (Fig. 7). This is the only species of Centris (Wagenknechtia) with dark brown pilosity and without metallic reflections on the metasoma (Figs. 6 and 8). The only two species with dark pilosity on the entire body are Centris rhodophthalma and C. vardyorum, but both species have intense bluish violet metallic reflections on the metasoma (Figs. 22, 24, 26 and 28). Type material. Holotype male (USNM, examined). Type locality. Peru: Arequipa. Floral records. See table 1. Distribution. This species ranges from southern Peru (Arequipa) to the III Atacama Region (Quebrada de Paipote) in northern Chile. CHILE: (Montalva & Ruz, 2010). II ANTOFAGASTA: Antofagasta (Wagenknecht, 1971; Toro, 1986b; Chiappa, 1998; Vivallo et al., 2002). Quebrada de Cifuncho, Quebrada de Paposo, Quebrada de Taltal (Toro et al., 1996; Vivallo et al., 2002). *Aguas Verdes, *Calama, *Chiuchiu, *Encanche, *La Negra, *Pampa Blanca, *Tumbre (Laskar). XV ARICA Y PARINACOTA: Chusmisa, Codpaquilla, Guatacondo, Parca, (Wagenknecht, 1971; Vivallo et al., 2002). *Ariquilda, *Mamiña, *Quebrada de Guatacondo, *Socoroma, *Termas de Mamiña. *ATACAMA: Chañaral, Copiapó, Potrerillos, Quebrada de Paipote. III TARAPACÁ: (Wagenknecht, 1971). *Iquique. PERÚ: (Soukup, 1943; Vivallo et al., 2002, 2003; Montalva & Ruz, 2010). AREQUIPA: Arequipa (Cockerell, 1926). Material examined. CHILE: II ANTOFAGASTA: 1 male: Atacama Aguas Verdes 22/10/1968 P. Millas, col (PUCV). 1 male and 1 female: Encanche 10/11/1974 Sielfeld, col\ Coll: Sielfeld/ MNHN Chile (MNHN). 1 male: La Negra 12/92 H. Toro, col (PUCV). 2 females: Antofagasta Chiuchiu 24/01/1972 H. Toro, col\ en Medicago sativa (PUCV). 2 females: Antofagasta Chiuchiu 24/01/1972 M. Rojas, col (PUCV). 2 females: Antofagasta Chiuchiu 24/01/1972 Montenegro, col\ en Medicago sativa (PUCV). 4 males and 8 females: II R. 30 Km. E. Calama 01/1992 S. Rodríguez, col (PUCV). 1 female: Calama Antofagasta Recarrhen 02/01/0972 R. Mendoza, col\ en Medicago sativa (IDEA). 1 female: Tumbre N. W. Laskar Vol. Antofagasta Prov. 3500 mts. 11/1965\ Centris (W.) cf escomeli Cockerell, 1920 F. Zanella det., 1998 (AMNH). XV ARICA Y PARINACOTA: 1 male: Quebrada Guatacondo, 2200 msnm. 15/03/1967 Loayza, col\ Coll. Cerda/ MNHN Chile (MNHN). 1 male: Ariquilda N. E. Iquique 28-29/04/1969, L. Peña, col (MNHN). 1 male: Tarapacá Guatacondo 09/1971 H. Toro, col\ en Medicago sativa (CTMI). 1 female: Tarapacá Guatacondo 09/1971 L. Ruz, col\ en Medicago sativa (CTMI). 1 male: Socoroma 16/04/1997 H. Vargas C., col (IDEA). 1 male: Mamiña 19/03/1968 (DZUP). 1 male: Guatacondo Tarapacá 19/09/1971 (PUCV). 10 males and 2 females: Tarapacá Guatacondo 19/09/1971 L. Ruz, col\ en Medicago sativa (PUCV). 6 males and 2 females: Tarapacá Guatacondo 09/1971 H. Toro, col\ en Medicago sativa (PUCV). 4 males: Tarapacá Guatacondo 18/04/1971 Montenegro, col\ en Medicago sativa (PUCV). 1 male: REVISION OF THE BEE SUBGENUS CENTRIS (WAGENKNECHTIA) Zootaxa 3683 (5) © 2013 Magnolia Press · 509 Tarapacá Guatacondo 19/04/1971 Montenegro, col\ en Medicago sativa (PUCV). 4 males: Tarapacá Chusmisa 19/ 04/1971 H. Toro, col\ en Medicago sativa (PUCV). 1 male: Tarapacá Chusmisa 19/04/1971 Montenegro col\ en Medicago sativa (PUCV). 2 males: Tarapacá Codpaquilla 22/04/1971 W. Sielfeld, col (PUCV). 1 male: Chile Region I 20km W Chuzmisa 11.iv.00 L. Packer\ Centris (Wagenknechtia) cf. escomeli det. J. S. Ascher 2004 (PCYU). 1 male: Tarapacá Codpaquilla 22/04/1971 Zuñiga, col (PUCV). 1 male: Tarapacá Codpaquilla 22/04/ 1971 L. Ruz, col\ en Medicago sativa (PUCV). III ATACAMA: 1 female: Chañaral Pampa Blanca 04/12/1990 R. Mendoza, col (IDEA). 1 male: On Loasa\ Chile, Region II R5 km1006, Chañaral S26°14.97’ W70°28.83’ 19.xi.2002 J. Grixti & A. Zayed\ Centris (Wagenknechtia) cf. escomeli det. J. Ascher 2004 (PCYU). 1 female: Potrerillos 15/01/1970 Sielfeld, col\ Coll: Sielfeld/ MNHN Chile (MNHN). 1 male: Copiapó Quebrada de Paipote 17/10/1988 H. Vargas C. y D. Bobadilla G., col\ en Prosopis chilensis (IDEA). 8 males and 2 females: Copiapó 15/ 01/1970 Jorquera, col\ Coll: Sielfeld/ MNHN Chile (MNHN). I TARAPACÁ: 2 males: Iquique Termas de Mamiña 14/07/1977 H. Vargas, col (IDEA). PERÚ: AREQUIPA: 1 male: Arequipa Peru (Escomel)\ Type No. 40493 U.S.N.M.\ Centris escomeli Ckll. Type\ USNM ENT 00534193 (USNM). 1 male and 3 females: Arequipa 12/1926 Dr. Escomel, col. Hemisia clypeata Fr. (MACN). 1 male: Perú Arequipa 2200 m. 19/02/1955 A. Meza, coll (CAS). 1 male: Arequipa 2200 m. 05/1955 A. Meza, col (DZUP). FIGURES 5–8. Centris escomeli Cockerell, 1926. Fig. 5. Female (Guatacondo, Chile): Head, frontal view. Fig. 6. Habitus, lateral view. Fig. 7. Male (Guatacondo, Chile): Head, frontal view. Fig. 8. Habitus, lateral view. Scale bars 1 mm. 510 · Zootaxa 3683 (5) © 2013 Magnolia Press VIVALLO Centris (Wagenknechtia) moldenkei Toro & Chiappa, 1989 (Figures 9−12, 31, 32 and 41) Centris moldenkei Toro & Chiappa, 1989. 15: 246, 247 (Original description). Toro et al., 1993. 18: 23, 27 (Floral record, distribution). Toro, 2002: 268, 270 (Floral record, bionomics). Montalva & Ruz, 2010. 35: 45 (List, distribution). Centris (Wagenknechtia) moldenkei; Vivallo et al., 2003: 82 (Descriptive note). Comments. Toro & Chiappa (1989) made a relatively complete description of this species, making its separation from other species of the subgenus easy. Some additional morphological characters of both sexes can be found in Vivallo et al. (2002). FIGURES 9–12. Centris moldenkei Toro & Chiappa, 1989. Fig. 9. Female (Guatacondo, Chile): Head, frontal view. Fig. 10. Habitus, lateral view. Fig. 11. Male (Guatacondo, Chile): Head, frontal view. Fig. 12. Habitus, lateral view. Scale bars 1 mm. Diagnosis. Head and mesosoma blackish with light brown legs (Figs. 9−12). Metasoma dark brown with translucent light brown distal margins of terga and sterna (Figs. 10 and 12). Pilosity yellow, darker on legs. Female: fore and hind elaiospathes vestigial, with the combs formed by separate hairs (Figs. 31 and 32). Male: mandible long and slender, with the second tooth closer to the apical tooth than to the third tooth (Fig. 11). Type material. Holotype male (AMNH/ Toro Collection, not examined). Type locality. Chile: I Tarapacá Region: La Tirana. Floral records. See table 1. Distribution. This species occurs exclusively in northern Chile, from XV Arica y Parinacota Region REVISION OF THE BEE SUBGENUS CENTRIS (WAGENKNECHTIA) Zootaxa 3683 (5) © 2013 Magnolia Press · 511 (Quebrada Vitor) to II Antofagasta Region (San Pedro de Atacama). CHILE: (Montalva & Ruz, 2010). II ANTOFAGASTA: San Pedro de Atacama (Vivallo et al., 2002). *Quillagua. XI ARICA Y PARINACOTA: La Tirana (Toro et al., 1993; Vivallo et al., 2002), Pica (Toro & Chiappa, 1989). *Guatacondo, *Quebrada de Tarapacá, *Quebrada Tana, *Quebrada Vítor. Material examined. CHILE: II ANTOFAGASTA: 1 male: Quillagua 10/1969 Montenegro, col\ Coll. Cerda/ MNHN Chile (MNHN). XV ARICA Y PARINACOTA: 1 female: Quebrada Vítor 21/10/1969 Montenegro, col\ Coll. Cerda/ MNHN Chile (MNHN). 52 males and 21 females: Tarapacá Qda. Tana 10/1969\ col. L. Ruz, Chile (PUCV). 52 males and 23 females: Tarapacá Qda. Tana 10/1969\ col. Montenegro, Chile (PUCV). 27 males and 16 females: Tarapacá Qda. Tana 101969\ col. Dazarola, Chile (PUCV). 64 males and 27 females: Tarapacá, Qda. Tana 10/1969\ col. Toro, Chile (PUCV). 1 female: Tarapacá Qda. Tana 10/1969 (PUCV). 1 male and 1 female: Tarapacá Qda. Tana 10/1969/col. F. Ramírez, Chile (PUCV). 1 male: Tarapacá Qda. Tana 10/1969\ col. De la Hoz, Chile (PUCV). 1 male: I Región La Tirana 29/09/1983\ B. Dyer, col. Chile\ PARATYPE (PUCV). 1 male: I Región La Tirana 29/ 09/1983\ C. Massad col. Chile (PUCV). 4 females and 20 males: Guatacondo 19/09/1971 Montenegro, col (PUCV). 8 males: Guatacondo 19/09/1971 Pizarro, col (PUCV). 5 females and 25 males: Guatacondo 19/09/1971 Toro, col (PUCV). 9 males and 2 fermales: Guatacondo 19/09/1971 L. Ruz, col\ en Medicago sativa (PUCV). 1 male: Km. 1680 04/10/1997 H. Toro, col\ en Prosopis tamarugo (PUCV). 1 male: Guatacondo 19/09/1971 Zúñiga, col (PUCV). 1 male: Quebrada de Tarapacá 20/11/1968 Sugaeta, col (MNHN). 1 female: Guatacondo 19/11/1971 L. Ruz, col (PUCV). Centris (Wagenknechtia) muralis Burmeister, 1876 (Figures 13−16, 37 and 39) Centris muralis Burmeister, 1876. 37: 162 (Original description). Vogel, 1974. 7: 216−218, 221, 245 (Floral records, distribution, cited). Neff & Simpson, 1981. 54: 107, 109 (Morphology, figure of fore elaiospathe). Chiappa, 1998. 22: 91 (Distribution). Medan et al., 2002. 34: 237 (Distribution, list). Sarzetti & Genise, 2011. 84: 179−180, 182 (Nesting biology, predators, figures of adult, larvae, nest and cells, nest architecture, cited). Cilla & Rolón, 2012a. 6: 5−8, 11, 12 (Distribution, nesting biology, nest architecture, figure of nest and cells). Cilla & Rolón, 2012b. 67: 575−581 (Cited, nest architecture, figures of adult, nest and cells, distribution, floral records, comparison with Centris nigerrima and C. pallida Fox, 1899). Rolón & Cilla, 2012. 66: 35−38 (Nest architecture, figures of nest and cells, floral records, bionomics). Centris (Wagenknechtia) muralis; Neff & Simpson, 1981. 54: 107, 109 (Morphology, distribution). Michelette & Camargo, 2000. 17: 655, 658, 663 (Distribution, floral records, cited). Rocha-Filho et al., 2009. 26: 301 (Cleptoparasite). Cilla & Rolón, 2012b. 67: 574 (Morphology, distribution, floral records). Vélez & Vivallo, 2012. 3357: 52 (Morphology). Anthophora plumigera Gribodo, 1893. 25: 391. Anthophora virgo Gribodo, 1893. 25: 390. Centris muralis melanopus Friese, 1908. 10: 58. Comments. The male of this species has a great variation in size and coloration of the pilosity with specimens almost completely covered by whitish pubescence (known as metander or beta males, see Snelling, 1984, Michener, 2000 and Vivallo & Zanella, 2012), and others with brown hairs on legs and metasoma. This variation is correlated with mating behaviour, as observed in other species of the genus, as Centris pallida Fox, 1899 and C. aenea Lepeletier, 1841 (see Michener, 2000 and references therein). Some diagnostic characters of this species can be found in Moure (1960) and Roig-Alsina (2000), but complete diagnoses of both sexes have not been published. Here both sexes are redescribed to help the identification of this species. Diagnosis. Both sexes with the body covered mainly with whitish pilosity, at least on head and mesosoma (Figs. 13−16). Female: primary basitibial plate with one concavity near the anterior and one near the posterior margin, without secondary plate (Fig. 37). Male: apex of the third mandibular third tooth concave (Fig. 15). Female. Measurements. Approximate body length: 18.6; head width: 6.2; forewing length: 13.6; intertegular span: 5.9; F1 length: 1.0; F2 length: 0.2; F3 length: 0.2; UID: 3.4; LID: 3.1; mandible length: 2.2; mandible basal width: 0.8; labrum length: 1.0; labrum width: 1.3. Coloration. Integument dark brown, mandible reddish brown, legs light brown (Fig. 13). Metasoma without metallic reflections (Fig. 14). Wing membranes slightly yellowish with veins light brown. Surface sculpture. Clypeus with coarse and relatively fine and sparse punctation, with smooth surface in the center of the disc. Labrum with fine and very dense punctation, without smooth areas, except on the basal margin. Paraocular area with fine, dense and uniform punctation, sparser near the lateral ocellus. 512 · Zootaxa 3683 (5) © 2013 Magnolia Press VIVALLO Mesoscutum and scutellum not areolate, with dense and relatively uniform punctation. Terga with fine and dense punctation, slightly sparser and coarser on T4 than on other terga. Pubescence. Whitish on head, mesosoma and T1 (Figs. 13 and 14). The rest of the body with brown hairs. Prepygidial fimbria light brown with plumose and simple hairs intermixed. Pygidial fimbria with brown, long and simple hairs. Structure. Flabellum circular with the anterior margin slightly acute and projected forward. Labial palpus with the 4th palpomere apically truncate. Maxillary palpus with five palpomeres (2nd>3rd>1st>4th≈5th). Mandible with four teeth with acute apex. Trimmal angle relatively straight, well developed. Acetabular carina reaching base of 4th tooth. Clypeal disc convex. Labrum relatively semicircular. Inner orbits of compound eyes subparallel (Fig. 13). Vertex in frontal view above the upper ocular tangent (Fig. 13). Distance between the lateral ocellus and compound eye more than three times the diameter of median ocellus (Fig. 13). Scutellum without longitudinal depression. Outer surface of the fore tibia with a short and shallow concavity. Fore elaiospathe with the anterior primary comb forming a compact structure, positioned along the basitarsus. Secondary anterior comb formed by three giant spatulate hairs. Hind elaiospathe with overlapping hairs. Primary basitibial plate elliptical with one concavity near the anterior and one near the posterior margin; without secondary plate (Fig. 37). Pygidial plate with rounded apex, secondary plate with open apex. Claws with preapical tooth very reduced on hind tarsus. Male. Measurements. Approximate body length: 16.6; head width: 5.3; forewing length: 14.1; intertegular span: 5.2; F1 length: 1.0; F2 length: 0.2; F3 length: 0.3; UID: 2.6; LID: 3.0; mandible length: 1.9; mandible basal width: 0.8; labrum length: 0.8; labrum width: 1.1. Coloration. Integument dark brown, reddish brown on mandible (Fig. 15). Metasoma without metallic reflections. Wing membranes slightly yellowish with veins light brown (Fig. 16). Surface sculpture. Clypeus with coarse and relatively fine and disperse punctation, with smooth and shiny surface in the center of the disc. Labrum with fine and very dense punctation, without smooth areas, except on the basal margin. Paraocular area with fine, dense and uniform punctation, sparser near the lateral ocellus. Mesoscutum and scutellum not areolate, with relatively uniform and not very dense punctation. Metasomal terga with fine and dense punctation, slightly sparser and coarser from T3 to T5. Pubescence. Whitish on head, mesosoma and T1 (Figs. 15 and 16). Legs with whitish and brown pubescence. Terga with white and brown pilosity (in some specimens the white or brown pilosity can be widely extended on metasoma) (Fig. 16). Structure. Mandible with three teeth of acute apex, except the third concave (Fig. 15). Trimmal angle acute, well developed. Acetabular carina reaching the base of the 3rd tooth. Clypeal disc convex. Labrum relatively triangular. Inner orbits of compound eyes converging upward (Fig. 15). Vertex at the same height of the upper ocular tangent. Distance between the lateral ocellus and the eye almost twice the diameter of median ocellus. Scutellum without longitudinal depression. Outer surface of the fore tibia with a short and shallow concavity. Claws with preapical tooth. T7 without pygidial plate. Type material. Centris muralis: Lectotype male (MACN, examined). Anthophora plumigera: Holotype female (Museu Cívico de Storia Naturale “Giacomo Doria” Genoa, Italy (MCG), not examined). Anthophora virgo: Holotype male (MCG, not examined). Centris muralis melanopus: Syntypes male and female (ZMB, not examined). Taxonomic decision for synonymy. Moure (1960) and Roig-Alsina (2000). Type locality. Centris muralis: Argentina: Buenos Aires: Carmen de Patagones. Anthophora plumigera: Argentina: Buenos Aires: Patagones. Anthophora virgo: Argentina: Buenos Aires: Patagones. Centris muralis melanopus: Argentina: Mendoza. Floral records. See table 1. Distribution. This species occurs exclusively in Argentina, from Salta Province (Cafayate and Chuscha) to Río Negro Province (El Carmen). ARGENTINA: (Chiappa, 1998). BUENOS AIRES: Buenos Aires (Jörgensen, 1912a, b; Schrottky, 1913a). Carmen de Patagones (Schrottky, 1902a, 1903), Delta (Roig-Alsina, 2000). CATAMARCA: San Fernando, Santa María (Roig-Alsina, 2000). *Andalgala, *Chumbicha, *Colpes, *Hualfia, *Joyango, *Pogoncillos, *Tinogasta. CÓRDOBA: Chancaní (Roig-Alsina, 2000). LA PAMPA: Cuchillocó (RoigAlsina, 2000). LA RIOJA: Anillaco (Cilla & Rolón, 2012a, Rolón & Cilla, 2012). Capayán (Cilla & Rolón, 2012a, b). Illiar, Patquía (Roig-Alsina, 2000). Pituil (Cilla & Rolón, 2012a). Udpidango (Cilla & Rolón, 2012a, b). Valle Los Colorados (Rolón & Cilla, 2012). Villa Casteló (Cilla & Rolón, 2012b). Vinchina (Sarzetti & Genise, 2011). *Santa Rosa. MENDOZA: (Vogel, 1974). Chacras de Coria (Jörgensen, 1909), Challao (Roig-Alsina, 2000). Desierto del Monte (Neff & Simpson, 1981). Mendoza (Friese, 1900, 1906, 1908; Jensen-Haarup, 1908; Jörgensen, 1909, 1912a, 1912b; Schrottky, 1902a, 1903, 1913a; Cockerell, 1919; Vivallo et al., 2002; Cilla & Rolón, 2012b). REVISION OF THE BEE SUBGENUS CENTRIS (WAGENKNECHTIA) Zootaxa 3683 (5) © 2013 Magnolia Press · 513 Papagallos (Roig-Alsina, 2000). Río Blanco (Medan et al., 2002). Tupungato, Uspallata (Roig-Alsina, 2000). *Chalalián. NEUQUÉN (Roig-Alsina, 2000; Vivallo et al., 2002). RÍO NEGRO: (Friese, 1900; Schrottky, 1903, 1913a; Vogel, 1974). Carmen de Patagones (Schrottky, 1902a, 1903; Wagenknecht, 1971). General Roca, Río Colorado (Roig-Alsina, 2000). *Lamarque. SALTA: *Cafayate, *Chuscha. SAN JUAN: (Cockerell, 1919; Schrottky, 1902b, 1903, 1913a). Pismanta, Rodeo (Roig-Alsina, 2000). Valle de Zonda (Michelette & Camargo, 2000). *Iglesia. SAN LUIS (Roig-Alsina, 2000). TUCUMÁN (Roig-Alsina, 2000). FIGURES 13–16. Centris muralis Burmeister, 1876. Fig. 13. Female (Mendoza, Argentina): Head, frontal view. Fig. 14. Habitus, lateral view. Fig. 15. Male (Mendoza, Argentina): Head, frontal view. Fig. 16. Habitus, lateral view. Scale bars 1 mm. Material examined. ARGENTINA: BUENOS AIRES: 1 male: Carm. Patag.\ 223\ 100\ Centris muralis holotypus Burm.\ Lectotipo desig. Moure, 1960 (MACN). 1 male: Carm. Patag. Argentina Dr. Burmeister 1881/ 114\ Co/type (BMNH). CATAMARCA: 2 males: Pr. Catamarca (Hualfia) 28/10/1973 J. L. Neff, coll\ en Larrea divaricata (CTMI). 4 males: Pr. Catamarca (Hualfia) 28/10/1973 J. L. Neff, coll. in Larrea divaricata (CTMI). 1 male: Pr. Catamarca (Hualfia) 04/11/1973 J. L. Neff, coll. C. muralis metandric male! Reared from nest Centris muralis ♂ J. L. Neff 02 (CTMI). 2 males: Pr. Catamarca Joyango/ Colpes 19/10/1973 J. L. Neff, coll.\ in Larrea divaricata (CTMI). 2 males and 4 females: Pr. Catamarca Joyango/ Colpes 19/10/1973 J. L. Neff, coll\ In Larrea divaricata (CTMI). 1 female: Catamarca Joyango/ Colpes 24/10/1972\ Larrea divaricata (CTMI). 4 females and 7 males: Chumbicha Catamarca Arg. Out-2957 M. Fritz, leg Dpto. Zool. U. F. Paraná (DZUP). 1 male: Chumbicha Catamarca-Arg. Out-1957 M. Fritz-leg. Dpto. Zool. U. F. Paraná (DZUP). 1 male: Chumbicha La Rioja R. Argentina (DZUP). 2 females: Pr. Catamarca (Tinogusta) 04/11/1973 J. L. Neff, coll\ In Bulnesia retama (CTMI). 514 · Zootaxa 3683 (5) © 2013 Magnolia Press VIVALLO 1 female: Pr. Catamarca (Poguncillos) 17/11/1973 J. L. Neff, coll\ In Larrea divaricata (CTMI). 1 female: Pr. Catamarca (Andalgala Desert site) 15/10/1973 J. L. Neff, coll\ in Bulnesia retama (CTMI). 1 male: Pr. Catamarca 28/10/1973 J. L. Neff, coll\ in Larrea divaricata (CTMI). LA RIOJA: 1 female and 1 male: Patquia Arg. Out-1975 A. Martínez (col) (DZUP). 2 males: Santa Rosa La Rioja Octubre de 1948-R. Arg. (DZUP). 1 male: Prov. La Rioja Patquia 10/1932 K. J. Hayward B. M. 1933-58 (DZUP). 1 male: Argentina Prov. La Rioja Patquia 10/1932 K. J. Hayward B. M. 1933/58 (BMNH). 1 male and 1 female: Argentina Prov. La Rioja Patquia 10/1932 K. J. Hayward B. M. 1933/58 (BMNH). MENDOZA: 1 female: 223\ Mendoza Centris muralis Burm. Allotypus\ muralis Burm. Mendozae (MACN). 1 female: 17/965\ Hemisia muralis (Burm.) forma melanopus Friese C. Schrottky det. 1910\ Mendoza Rep. Argentina\ 97298\ 17/965 Centris muralis Burm. F. melanopus Fr. Argentina Schrottky det (DZUSP). 1 female: Pr. Mendoza Chalalián 21/11/73, J. L Neff, coll\ in Larrea divaricata (CTMI). 1 male: Cord. de Mendoza 16/11/1907 P. Jörgensen (ZMUC). 1 male: 16/11/1907 P. Jörgensen (ZMUC). 1 male: Mendoza 19/10/ 1908 P. Jörgensen (ZMUC). 1 male: Mendoza 23/10/1908 P. Jörgensen (ZMUC). 1 male: Mendoza 05/11/1908 P. Jörgensen (ZMUC). 2 males: Mendoza 06/11/1908 P. Jörgensen (ZMUC). 2 males: Mendoza 08/11/1908 P. Jörgensen (ZMUC). 1 male: Mendoza 09/11/1908 P. Jörgensen (ZMUC). 2 males and 4 females: Arg. Mendoza above Uspallata 1,940 m. 06/02/1979 C. & M. Vardy B. M. 1980/67 (BMNH). 1 male: Arg. Mendoza above Uspallata 1,940 m. 06/02/1979 C. & M. Vardy B. M. 1980/67 all bees of this sp. nesting in holes in vertical sandy cliff face (BMNH). RÍO NEGRO: 1 female and 2 males: Lamarque R. Negro R. A. 10/1975 M. Fritz\ coleção Campos Seabra (DZUP). SALTA: 2 males: Salta Cafaite-RA R. Chuscha 11/1960 R. J. Llano-leg (DZUP). SAN JUAN: 1 female: Zonda San José Argentina 10-X-1992 10:30h Michelette leg. 949725 (RPSP). 1 male: Zonda San José Argentina 10-X-1992 10:15h Michelette leg. 940726 (RPSP). 1 male: Iglesia San José Argentina 31-X-1992 10:40h Michelette leg. 940751 (RPSP). 1 male: Iglesia San José Argentina 31-X-1992 19:00h Michelette leg. 940753 (RPSP). 1 male: Iglesia San José Argentina 31-X-1992 09:00h Michelette leg. 940752 (RPSP). Centris (Wagenknechtia) orellanai Ruiz, 1940 (Figures 17−20, 35, 36 and 40) Centris nigerrima orellanai Ruiz, 1940. 44: 335 (Original description). Centris orellanai; Snelling & Brooks, 1985. 369: 19 (Cleptoparasite). Chiappa, 1998. 22: 90, 91 (Distribution, key). Montalva & Ruz, 2010. 35: 45 (List, distribution). Centris (Wagenknechtia) orellanai; Rocha-Filho et al., 2009. 26: 301 (Cleptoparasite). Giannini et al., 2013. 258: 78 (Floral record). Centris (Wagenknechtia) orellanae [sic]; Toro, 1986b. 13: 127 (Distribution). Comments. During the preparation of this publication, I visited the collection of the Colegio San Pedro Nolasco, Santiago, Chile (CSPN) to photograph the holotype of Centris orellanai. According to J. Salamanca, the curator of the collection, the holotype had unfortunately been destroyed along with other specimens of that collection. In this paper a neotype for this species is designated and, described in detail to facilitate the differentiation of this species from others in the subgenus. A complete description of the male of this species can be found in Vivallo et al. (2002). This species is similar to Centris cineraria, differing (both sexes) in the coloration of the pubescence on mesoscutum and scutellum (light brown to orange in C. orellanai (Figs. 17–20); relatively whitish in C. cineraria (Figs. 1–4)), and by the metallic reflections on metasoma (relatively greenish in C. orellanai (Figs. 18 and 20); bluish in C. cineraria (Figs. 2 and 4)). Both females also differ in the hair number of the secondary comb of the fore elaiospathe (four in C. orellanai; three in C. cineraria) and in the preapical tooth of the claw of midleg (present in C. orellanai; absent in C. cineraria). Diagnosis. Both sexes with integument dark brown on head and mesosoma (Figs. 17 and 19). Metasoma with greenish metallic reflections. Mesoscutum and scutellum with light brown or orange hairs. The rest of the body with dark brown pilosity (Figs. 18 and 20). Female: fore elaiospathe located in the distal half of foretibia with the secondary comb formed by four giant spatulate hairs. Hairs of hind elaiospathe not overlapping. Claws with preapical tooth, except in hind legs. REVISION OF THE BEE SUBGENUS CENTRIS (WAGENKNECHTIA) Zootaxa 3683 (5) © 2013 Magnolia Press · 515 FIGURES 17–20. Centris orellanai Ruiz, 1940. Fig. 17. Female (Neotype female: Los Andes, Chile): Head, frontal view. Fig. 18. Habitus, lateral view. Fig. 19. Male (Baños del Toro, Chile): Head, frontal view. Fig. 20. Habitus, lateral view. Scale bars 1 mm. Neotype female. Measurements. Approximate body length: 14.5; head width: 5.3; forewing length: 8.9; intertegular span: 5.9; F1 length: 1.3; F2 length: 0.2; F3 length: 0.2; UID: 3.0; LID: 3.3; mandible length: 3.2; mandible basal width: 1.1; labrum length: 1.1; labrum width: 1.5. Coloration. Integument dark brown to black, malar area light brown, legs slightly reddish brown (Fig. 17). Metasomal terga and sterna dark brown with greenish metallic reflections except at the distal margins (Fig. 18). Wing membranes slightly yellowish with veins light brown (Fig. 18). Surface sculpture. Clypeus with coarse and relatively uniform punctation throughout, without defined smooth or elevated areas. Labrum with similar punctation but denser, mainly toward the distal edge; basal margin with a smooth area, slightly shorter at sides. Paraocular area with fine, dense and uniform punctation, sparser near the lateral ocellus. Mesoscutum and scutellum not areolate, with dense and relatively uniform punctation. T1 with fine and dense punctation at sides, sparser on disc. T2 with similar punctation, but the punctures of the disc are denser than on disc of T1. T3 and T4 with fine and dense punctation throughout, including the discs. T5 with similar punctation, but much denser than on previous terga. Pubescence. Dark brown, except mesoscutum, scutellum and tegula with light brown to orange hairs (Fig. 17 and 18). T1 with dark brown slightly blackish, long and plumose hairs laterally, sparser, shorter and simple on disc. All other terga with dark brown to black, short and simple pilosity, mixed with some longer hairs but not covering the entire surface. Prepygidial fimbria with plumose and simple hairs intermixed. Pygidial fimbria with brown, long and simple hairs. Structure. Flabellum circular with the anterior margin slightly acute and projected forward. Labial palpus with the 4th 516 · Zootaxa 3683 (5) © 2013 Magnolia Press VIVALLO palpomere apically truncate. Maxillary palpus with five palpomeres (2nd>3rd>1st>4th≈5th). Mandible with four teeth of acute apex (Fig. 17). Apical tooth slightly widened at the base. Gap between third and fourth teeth longer than between others. Trimmal angle obtuse, poorly developed, almost absent. Acetabular carina reaching the base of the 4th tooth. Clypeal disc convex. Labrum semicircular. Inner orbits of compound eyes diverging below. Vertex above the upper ocular tangent. Distance between the lateral ocellus and the eye more than twice the diameter of the median ocellus. Scutellum slightly depressed longitudinally. Outer surface of the anterior tibia with a short and shallow concavity. Fore elaiospathe with the anterior primary comb forming a compact structure only in the distal half. Secondary anterior comb formed by four giant spatulate hairs. Hind elaiospathe with the hairs separated by more than twice their own diameter. Primary basitibial plate elliptical with rounded distal end and a deep cavity in the distal half. Secondary basitibial plate indicated only by an elevation from the anterobasal margin of the primary plate towards the posterior margin, bordering the cavity (Fig. 35). Pygidial plate with rounded apex and without secondary plate (Fig. 36). Claws of foreleg with preapical tooth very reduced in midleg and absent in hind leg. Type material. Holotype female (CSPN) destroyed. Neotype female here designated with the following information: Chile: V Región: Los Andes 05/02/2002 Vivallo, col\ Neotype Centris (Wagenknechtia) orellanai Ruiz, 1940. The neotype will be deposited at MNHN. Type locality. Chile: V Valparaíso Region: Los Andes. Floral records. See table 1. Distribution. This species occurs exclusively in Chile, from IV Coquimbo Region (Baños del Toro) to V Valparaíso Region (Los Andes). CHILE: (Moure, 1950; Montalva & Ruz, 2010). IV COQUIMBO: (Toro, 1986b). Los Pelambres (Ruiz, 1940), Baños del Toro, Cuncumén, Río La Laguna (Vivallo et al., 2002), Río Seco, Talahuén (Wagenknecht, 1971). *Illapel, *Lagunitas. VI LIBERTADOR GENERAL BERNARDO O’HIGGINS: *Termas del Flaco. V VALPARAÍSO: Los Andes (Toro, 1986b). *Los Maitenes. Material examined. CHILE: IV COQUIMBO: 2 males: Coquimbo entre Vicuña y frontera Int. 1970-72. leg. A. R. Moldenke (CTMI). 4 males: Baños del Toro 06/01/1966 L. Peña, col\ Coll. Cerda/ MNHN Chile (MNHN). 1 female: Baños del Toro 24/09/1946 (MNHN). 2 males: Baños del Toro 07/01/1950 (MNHN). 1 female: Baños del Toro 07/01/1950 (MNHN). 1 male: Baños del Toro 4 En.48\ orellanai\ ♂ Centris orellanae-Ruiz (DZUP). 2 females: Río Laguna 22/09/1946 Sielfeld, col\ Coll: Sielfeld/ MNHN Chile (MNHN). 1 female: Rio Laguna Chile 22-XI-1946 Wagenknecht (DZUP). 1 female: R. Laguna 2700 mts 22-XI-1946 (DZUP). 2 females: Río Seco Lagunitas 16/12/1940 Sielfeld, col\ Coll: Sielfeld/ MNHN Chile (MNHN). 1 female: Lagunitas 3900 mts. 16-XII1940\ orellanai (DZUP). 1 female: Hda. Illapel Chile 11-XI-54 2800 m.alt. L.E. Pena\ Centris orellanai (Ruiz) Det. J. S. Moure 1956 (DZUP). VI LIBERTADOR GENERAL BERNARDO O’HIGGINS: VI Región T. del Flaco\ 5-XII1982\ Magunacelaya col. Chile (PUCV). V VALPARAÍSO: 1 male and 3 females: Los Andes: Los Maitenes 3500 msnm. 03/02/1973 Sielfeld, col\ Coll. Cerda/ MNHN Chile (MNHN). Centris (Wagenknechtia) rhodophthalma Pérez, 1911 (Figures 21−24 and 41) Centris rhodophthalma Pérez, 1911. 15: 55, 56 (Original description). Snelling & Brooks, 1985. 369: 19 (Cleptoparasites). Toro, 1986a. 57: 83 (Distribution, bionomics). Montalva & Ruz, 2010. 35: 45 (List, distribution). Sérsic & Cocucci, 1999. 194: 717 (Floral record, male behaviour (misidentification of Centris vardyorum)). Tadey, 2011. 59: 399, 402 (Floral record, male behaviour (misidentification of C. vardyorum)). Centris (Wagenknechtia) rhodophthalma; Simpson et al., 1977. 247: 151 (Floral record, distribution). Toro, 1986b. 13: 127 (Distribution). Cocucci et al., 1996. 34: 191 (Floral record). Vivallo et al., 2003: 79 (Cited). Rocha-Filho et al., 2009. 26: 301 (Cleptoparasite). Centris rodophthalma [sic]; Simpson, 1989. 14: 409−411 (Distribution, floral record, descriptive note, cited, figure of a female on plant host). Chiappa, 1998. 22: 87−91 (Cited, taxonomy, redescription, distribution, floral records, key, note, figures of morphology of adults). Chiappa, 2000. 46: 19, 21−25 (Cited, figures of immature stages and cells, bionomics, descriptive note of cell and larva, morphology, comparison with Centris caesalpiniae Cockerell, 1897, C. tamarugalis Toro & Chiappa, 1989 and C. violacea Lepeletier, 1841). Chiappa et al., 2000. 24: 20−25, 27 (Distribution, floral records, figures of nest, cells and flying male behaviour, nest architecture, bionomics, parasite, nesting and adult behaviour, cited, comparison with C. nigerrima and C. cineraria). Chiappa et al., 2000. 64: 134−137 (Cited, bionomics, floral records, distribution, morphology, figures of collecting behaviour). REVISION OF THE BEE SUBGENUS CENTRIS (WAGENKNECHTIA) Zootaxa 3683 (5) © 2013 Magnolia Press · 517 Comments. Both sexes of this species were redescribed by Chiappa (1998) and Vivallo et al. (2002). Here only diagnostic characters that permit identification of both sexes are given. Diagnosis. Both sexes with dark brown integument on head, legs and mesosoma (Figs. 21–24). Metasoma with bluish violet metallic reflections (Figs. 22 and 24). Body completely covered with brownish pilosity. Compound eyes red in live specimens. Female: fore elaiospathe with the anterior comb starting below the strigilis and extending to the apex of the forebasitarsus. Both elaiospathes with overlapping hairs. This species is similar to Centris vardyorum. Both species share the dark pilosity on the body and the bluish violet metallic reflections on metasoma, along with the red eyes of the females in living specimens. They differ in the shape of the labrum (semicircular in C. rhodophthalma (Figs. 21 and 23); triangular in C. vardyorum (Figs. 25 and 27)). The females differ mainly in the morphology of the fore elaiospathe (with compact primary and secondary combs and disposed along the basitarsus in Centris rhodophthalma; highly modified, forming a short comb disposed on the upper margin with short, dense and branched hairs on the ventral surface of the basitarsus in C. vardyorum (Fig. 33)). The males differ in the color of the eyes in living specimens (red in C. rhodophthalma; green in C. vardyorum). FIGURES 21–24. Centris rhodophthalma Pérez, 1911. Fig. 21. Female (El Pangue, Chile): Head, frontal view. Fig. 22. Habitus, lateral view. Fig. 23. Male (El Pangue, Chile): Head, frontal view. Fig. 24. Habitus, lateral view. Scale bars 1 mm. Type material. Holotype female (MNHP, examined). Type locality. Chile: III Atacama Region: Chañarcillo. Floral records. See table 1. 518 · Zootaxa 3683 (5) © 2013 Magnolia Press VIVALLO Distribution. This species occurs exclusively in Chile, from III Atacama Region (Parque Nacional Llanos de Challe) to V Valparaíso Region (Putaendo). CHILE: (Moure, 1950; Toro, 1986a; Simpson et al., 1977; Montalva & Ruz, 2010). III ATACAMA: Chañarcillo (Pérez, 1911; Wagenknecht, 1971; Simpson, 1989; Chiappa, 1998). Cuesta Pajonales, La Travesía, Las Juntas, Los Loros, Totoral (Chiappa, 1998; Vivallo et al., 2002). *Cachiyuyo, *Chañaral de Aceituno, *Copiapó, *Domeyko, *Parque Nacional Llanos de Challe, *Quebrada Cóndores, *Vallenar. IV COQUIMBO: Andacollito (Chiappa et al., 2000; Chiappa & Rodríguez, 2001). El Pangue (Chiappa, 2000). Hurtado (Wagenknecht, 1939; Chiappa et al., 2000; Chiappa & Rodríguez, 2001). Illapel (Wagenknecht, 1971), Paihuano, Pisco Elqui (Wagenknecht, 1939; Ruiz, 1940), Alcohuás, Balala, Cochiguás, Combarbalá, El Tofo, La Higuera, Las Breas, Los Choros, Los Molles, Pajaritos (Chiappa, 1998; Vivallo et al., 2002). Cuesta del Almendro, Monte Grande (Wagenknecht, 1939). Puntilla de Diaguitas (Wagenknecht, 1939; Ruiz, 1940; Chiappa, 1998). Río Claro, Río Cochiguaz (Monte Grande) (Ruiz, 1940). Varillar (Chiappa, 1998, 2000; Chiappa et al., 2000; Chiappa & Rodríguez, 2001; Vivallo et al., 2002). Vicuña (Wagenknecht, 1939; Toro, 1986b; Chiappa, 1998). *Algarrobal, *Chañares, *Condoriaco, *El Chacay, *El Molle, *Guampulla, *Incahuasi, *La Serena, *Las Cardas, *Los Hornos, *Río Turbio. *V VALPARAÍSO: Coimas, Putaendo. Material examined. CHILE: III ATACAMA: 1 female: Type\ Chañarcillo\ Centris rhodophthalma JP (MNHP). 11 females: On Dinemagonum gayanum\ Chile, Region III P. N. Llanos de Challe 6.X.2002 J. Grixti & A. Zayed (PCYU). 2 males and 1 female: Camino Domeyko 24/09/1977 O. León, leg (MNHN). 1 male: Cachiyuyo 24/09/1972 Sielfeld, col\ Coll: Sielfeld/ MNHN Chile (MNHN). 1 male: Vallenar 13/10/1972 M. Pino, col\ Coll: Sielfeld/ MNHN Chile (MNHN). 1 male: Atacama Las Juntas 09/1968 F. Ramírez, col (PUCV). 1 female: Vallenar 20/10/1988 H. Vargas C. y D. Bobadilla G., col (IDEA). 1 male: III R. Las Juntas 05/10/1982 V. Farías, col (PUCV). 1 male: III R. Las Juntas 05/10/1982 H. Toro, col (PUCV). 1 male: III R. Las Juntas 05/10/1982 Montenegro, col (PUCV). 1 female: Atacama Copiapó 03/10/1972, J. Neff, col (PUCV). 1 female: Atacama Qda. Cóndores 09/10/1977 Balart, col (PUCV). 1 female: Atacama Qda. Cóndores 09/10/1977 L. Ruz, col (PUCV). 1 female: Atacama Qda. Cóndores 09/10/1977 V. Cabezas, col (PUCV). 1 female: Totoral 06/10/1982 B. Dyer, col (PUCV). 1 male: Las Juntas 05/10/82 Montenegro, col (PUCV). 1 male: Travesía 11/10/1987 E. Chiappa, col (PUCV). 2 females: Las Juntas 09/1968 De la Hoz, col (PUCV). 1 male: Atacama Las Juntas 09/1968 F. Ramírez, col (PUCV). 1 female: Chañaral de Aceituno 10/1858. L. Peña, col (DZUP). 1 female: Coquimbo Los Loros 22/09/ 1984 O. Martínez, col (PUCV). 6 females: Pajonales 16/10/1947 (MNHN). 1 female: Pajonales 24/10/1947/ Coquimbo Chile Wagenknecht (PUCV). IV COQUIMBO: 1 male: Las Cardas 17/08/1946 (MNHN). 1 male: Las Cardas 23/09/1947 G. Kuschel, col (MNHN). 1 male: Vicuña 14/09/1938 (DZUP). 1 female: Vicuña 12/09/1938 (DZUP). 1 female: Coquimbo El Molle 350 msnm. 21/09/1973 H. Vásquez C., col (IDEA). 2 males: Coquimbo El Molle 350 msnm. 20/09/1973 A. Aguilera, col (IDEA). 1 female: Paihuano 01/10/1935 (MNHN). 1 male: Vicuña 13/09/1939 (MNHN). 1 female: Coquimbo 10/12/1946 Wagenknecht, leg (MNHN). 1 male: La Serena 11/10/1947, G. Kuschel, col (MNHN). 1 female: El Pangue 18/09/1948 (MNHN). 1 female: El Tofo 21/10/1967 B. W. H., col (MNHN). 2 females: Incahuasi 10/1970 Sielfeld, col\ Coll: Sielfeld/ MNHN Chile (MNHN). 1 male: Coquimbo Incahuasi 09/1968 De la Hoz, col (PUCV). 1 male: Coquimbo Incahuasi 09/1968 L. Ruz, col (PUCV). 1 male: Coquimbo Incahuasi 09/1968 Montenegro, col (PUCV). 1 male: Coquimbo Vicuña 09/1968 De la Hoz col (PUCV). 2 males: Coquimbo Vicuña 09/1968 H. Toro, col (PUCV). 4 males: Coquimbo Incahuasi 09/1968 H. Toro, col (PUCV). 5 males: Coquimbo Vicuña 09/1968 L. Ruz, col (PUCV). 2 males and 1 female: Coquimbo Vicuña 09/1968 H. Toro, col (PUCV). 1 male: Coquimbo Vicuña 09/1968 De la Hoz, col (PUCV). 5 males: Coquimbo Vicuña 09/1968 L. Ruz, col (PUCV). 5 females: Coquimbo Vicuña 09/1968 L. Ruz, col (PUCV). 1 female and 2 males: Coquimbo Vicuña 09/1968 Montenegro, col (PUCV). 1 female: Coquimbo Vicuña 09/1968 Ramírez, col (PUCV). 1 male and 1 female: Coquimbo Incahuasi 09/1968 L. Ruz, col (PUCV). 4 males and 1 female: Coquimbo Incahuasi 09/1968 H. Toro, col (PUCV). 1 male and 1 female: Coquimbo Incahuasi 09/1968 De la Hoz, col (PUCV). 2 males and 4 females: Coquimbo Vicuña 09/1968 H. Toro, col (PUCV). 5 females and 1 male: Coquimbo Vicuña 09/1968 De la Hoz, col (PUCV). 2 females: Coquimbo Incahuasi 22/09/1970 De la Hoz, col (PUCV). 1 female: Coquimbo Incahuasi 22/09/1970 H. Toro, col (PUCV). 1 female: Coquimbo Incahuasi 22/ 09/1970 V. Cabezas, col (PUCV). 1 male: Coquimbo El Pangue 10/1972 H. Toro, col (PUCV). 4 males: Coquimbo El Pangue 10/1972 O. Martínez, col (PUCV). 4 males and 1 female: Coquimbo El Pangue 10/1972 V. Cabezas, col\ en Stachys serrata (PUCV). 7 males: Coquimbo El Pangue 10/1972 V. Cabezas, col\ en Adesmia melanthes (PUCV). 4 males: Coquimbo El Pangue 10/1972 L. Ruz, col\ en Stachys serrata (PUCV). 5 males: Coquimbo El Pangue 10/1972 L. Ruz, col\ en Adesmia melanthes (PUCV). 2 females: Algarrobal 800 msnm. 10/09/1973 H. REVISION OF THE BEE SUBGENUS CENTRIS (WAGENKNECHTIA) Zootaxa 3683 (5) © 2013 Magnolia Press · 519 Vásquez C., col (IDEA). 1 female: El Molle 350 msnm. 19/09/1973 H. Vásquez C., col (IDEA). 1 female and 1 male: El Molle 350 msnm. 21/09/1973 H. Vásquez C., col (IDEA). 1 female: Pisco Elqui 1300 msnm. 23/09/1973 H. Vázquez C., col (IDEA). 1 male: Río Turbio 11/11/1939 (MNHN). 1 female: Coquimbo Qda. Los Choros 12/10/ 1977 V. Cabezas, col (PUCV). 1 male: El Pangue Coquimbo 13/10/1977 De la Hoz, col (PUCV). 1 male: El Pangue Coquimbo 13/10/1977 Magunacelaya, col (PUCV). 2 males: El Pangue Coquimbo 13/10/1977 Balart, col (PUCV). 1 male: Coquimbo Las Breas 17/10/1979 L. Ruz, col (PUCV). 13 males: Coquimbo Las Breas 17/10/ 1979 Magunacelaya, col (PUCV). 1 female: Coquimbo Las Breas 17/10/1979/ Chile leg. H. Toro (PUCV). 1 female: Coquimbo Las Breas 17/10/1979 H. Toro, col (PUCV). 1 male: Coquimbo Las Breas 23/09/1980 V. Cabezas, col (PUCV). 1 male: Coquimbo Las Breas 26/09/1980 H. Donoso, col (PUCV). 1 male: Incahuasi 10/10/ 1981 B. Dyer, col (PUCV). 1 male: Incahuasi 10/10/1981 H. Burgos, col (PUCV). 1 female: Coquimbo Incahuasi 10/10/1981 H. Burgos, col (PUCV). 2 females: Coquimbo Incahuasi 10/10/1981 O. Martínez, col (PUCV). 1 female: Coquimbo Chañares 12/09/1984, H. Toro, col (PUCV). 1 male: El Pangue 10/1987 De la Hoz, col (PUCV). 1 male: Hurtado 10/1987 O. Martínez, col (PUCV). 1 female: El Tofo 10/1987 E. Chiappa, col (PUCV). 1 female: Incahuasi 22/09/1980 Toro (col) (PUCV). 1 male: Incahuasi 14/10/1988 E. Chiappa, col (PUCV). 1 male: Incahuasi 14/10/1988 V. Cabezas, col (PUCV). 13 females: Coquimbo Incahuasi 14/10/1988 V. Cabezas, col (PUCV). 2 females: Coquimbo Incahuasi 14/10/1988 L. Ruz, col (PUCV). 5 females: Coquimbo Incahuasi 14/10/ 1988 E. Chiappa, col (PUCV). 1 female: Coquimbo Incahuasi 14/10/1988 H. Toro, col (PUCV). 1 female: IV Región Incahuasi 14/10/1988\ M. Rojas col. Chile (PUCV). 1 female: Chile Region IV Guampulla SW. of Samo Alto 19.x.01 Packer & Fraser (PCYU). 2 females: Vicuña 11/1994. D. Bobadilla, col (IDEA). 1 female: Chile Region IV 2–4km S Vicuna HWY 41 9.X.01 Packer & Fraser (PCYU). 1 female: Codoriaco 20/11/1968 (MNHN). 1 male: Condoriaco 17/08/1946 (MNHN). 1 female: On Loasa tricolor\ Chile, Region IV El Chacay S29°40.05 W71°09.88’ 24.x.2002 J. Grixti & A. Zayed (PCYU). 1 female: On Loasa tricolor\ Chile Region IV PanAm N. Los Hornos S29°34.89’ W 71°14.50’ 9.X.2002 J. Grixti & A. Zayed (PCYU). V VALPARAÍSO: 2 males: Coimas 15/09/1974\ Coll: Sielfeld/ MNHN Chile (MNHN). 2 males and 1 female: Putaendo 17/09/2001 F. Vivallo, col (MNRJ). Centris (Wagenknechtia) vardyorum Roig-Alsina, 2000 (Figures 25−28, 33, 34 and 41) Centris (Wagenknechtia) vardyorum Roig-Alsina, 2000. 2: 190 (Original description). Tadey, 2011. 59: 715–717 (Floral records, cited, bionomics, collecting behaviour). Centris (Wagenknechtia) sp.; Neff & Simpson, 1981. 54: 106, 107 (Distribution, morphology, figure of fore elaiospathe). Centris (Wagenknechtia) sp. nova; Simpson et al., 1990. 173: 210, 213, 220, 221 (Morphology, floral record). Centris sp. nova; Simpson et al., 1990. 173: 216 (Morphology). Centris nigerrima; Vogel, 1974. 7: 217 (Distribution, floral record (misidentification)). Montalva & Ruz, 2010. 35: 45 (Partim, Mendoza record (misidentification)). Centris rhodophthalma; Sérsic & Cocucci, 1999. 194: 717 (Floral record, male behaviour (misidentification)). Tadey, 2011. 59: 399, 402 (Floral record, male behaviour (misidentification)). Comments. This species was described recently (Roig-Alsina, 2000), so its redescription seems redundant. Below are given some diagnostic characters to help the identification of both sexes of this species. Diagnosis. Both sexes with integument and pilosity dark brown (Figs. 25–28). Metasoma with relatively weak metallic bluish violet reflections (Figs. 26 and 28). Female: fore elaiospathe strongly modified, with short, plumose and very dense pilosity on its ventral surface (Fig. 33). These hairs extend throughout the ventral surface of the fore tarsi. Hind elaiospathe with the hairs of the primary comb slightly separated, less than their diameters (Fig. 34). Eyes red in females and green in males (in live specimens). Type material. Holotype female (MACN, examined). Type locality. Argentina: Río Negro: 8 km SW General Roca, Paso Córdoba. Floral records. See table 1. Distribution. This species occurs exclusively in Argentina, from Mendoza Province (Chacras de Coria) to Río Negro Province (San Antonio). ARGENTINA: (Vivallo et al., 2002). CATAMARCA: Punta de Balasto/ Minas Capillitas (Simpson et al., 1990). MENDOZA: Chacras de Coria (Jörgensen, 1912a, b; Roig-Alsina, 2000). Mendoza (Friese, 1908, 1910; Jensen-Haarup, 1908; Jörgensen, 1909, 1912a, b; Schrottky, 1913a; Wagenknecht, 520 · Zootaxa 3683 (5) © 2013 Magnolia Press VIVALLO 1971). Pedregal (Jörgensen, 1909, 1912a). NEUQUÉN: Arroyito, El Chocón (Tadey, 2011). RÍO NEGRO: San Antonio (Simpson et al., 1990), Paso Córdoba (8 Km. SW General Roca) (Roig-Alsina, 2000). Material examined. ARGENTINA: RÍO NEGRO: 1 female: Arg: Río Negro Paso Córdoba; river 8k sw General Roca. 6-7.xi.1997 C. & M. Vardy\ Centris vardyorum ♀ sp. n. Holotypus A. Roig Alsina 2000 (MACN). 1 female and 2 males: R. A. Río Negro S. A. Oeste (60 Km So.) 24/11/1973 J. L. Neff on flowers of Monttea aphylla 88207 (♀) 88205 (♂) y 88204 (♂) Paratype Centris vogeli det. Moldenke’76\ Centris vardyorum J. L. Neff 02 ♀ (CTMI). 1 female and 2 males: Pr. Río Negro Km 1200 S. A. oeste coll. J. L. Neff 88432 (♀) 88433 (♂) y 88431 (♂) 26/11/1973 #refers to host and date on flowers of Monttea aphylla\ Paratype Centris vogeli det. Moldenke’76\ Centris vardyorum J. L. Neff 02 ♀ (CTMI). FIGURES 25–28. Centris vardyorum Roig-Alsina, 2000. Fig. 25. Female (El Challao, Argentina): Head, frontal view. Fig. 26. Habitus, lateral view. Fig. 27. Male (El Challao, Argentina): Head, frontal view. Fig. 28. Habitus, lateral view. Scale bars 1 mm. Key to the species of the subgenus Centris (Wagenknechtia) Females 1 2 Metasoma with metallic blue or greenish reflections (Figs. 2, 18, 22 and 26) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Metasoma without metallic reflections (Figs. 6, 10 and 14) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Mesoscutum and scutellum with whitish or yellowish brown to orange hair (Figs. 2 and 18); metasoma with bluish or greenish reflections; eyes black (in live specimens); fore elaiospathe located on the distal half of the basitarsus (Fig. 29); hind elaiospathe with the hairs of the primary comb separate (Fig. 30); distal edge of 4th labial palpomere truncate . . . . . . . . . . . . . . . . . 3 REVISION OF THE BEE SUBGENUS CENTRIS (WAGENKNECHTIA) Zootaxa 3683 (5) © 2013 Magnolia Press · 521 - 3 4 5 6 - Mesoscutum and scutellum with dark brown hairs (Figs. 22 and 26); metasoma with blue violet reflections; eyes red (in live specimens); fore elaiospathe extending for the entire length of the basitarsus or strongly modified with abundant, short and dense pilosity on its ventral surface (Fig. 33); hind elaiospathe formed by overlapping hairs; distal edge of 4th labial palpomere rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Mesoscutum and scutellum with whitish hairs (Fig. 2); metasoma with bluish metallic reflections; fore elaiospathe with the secondary comb formed by three giant spatulate hairs (Fig. 29) . . . . . . . . . . . . . . . . . . . . . . . . . . Centris cineraria Smith, 1854 Mesoscutum and scutellum with light brown to orange hairs (Fig. 18); metasoma with blue greenish metallic reflections; fore elaiospathe with the secondary comb formed by four giant spatulate hairs . . . . . . . . . . . . . . . . . . . Centris orellanai Ruiz, 1940 Clypeus with denser punctation in the upper half, sparser on the lower half; labrum triangular (Fig. 25); fore elaiospathe highly modified with short, dense and branched hairs on the ventral surface and on the anterior margin of the basitarsus (Fig. 33) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centris vardyorum Roig-Alsina, 2000 Clypeus with dense and relatively uniform punctation; labrum semicircular; fore elaiospathe with compact primary and secondary combs positioned along the basitarsus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centris rhodophthalma Pérez, 1911 Body covered mainly with whitish pilosity, at least on head and mesosoma (Fig. 14); basitibial plate with one concavity near the anterior and one near the posterior margin, without secondary plate (Fig. 37) . . . . . . . . . Centris muralis Burmeister, 1876 Body completely covered with dark brown or yellowish pilosity (Figs. 6 and 10); basitibial plate with a central concavity in the distal half of the primary plate, secondary plate present, with the lower edge concave . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Body completely covered with dark brown hairs (Fig. 6); maxillary palpus five segmented; fore and hind elaiospathe normally developed with compacted combs; distal edges of terga and sterna not broadly translucent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centris escomeli Cockerell, 1926 Body covered with yellowish pilosity (Fig. 10); maxillary palpus four segmented; fore and hind elaiospathe vestigial, with both primary combs formed by separated hairs (Figs. 31 and 32); distal edges of terga and sterna broadly translucent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centris moldenkei Toro & Chiappa, 1989 FIGURES 29–34. Oil-collecting structures. Figs. 29, 31 and 33. Fore elaiospathes. Figs. 30, 32 and 34. Hind elaiospathes. Figs. 29 and 30. Centris cineraria Smith, 1854. Scale bars 500 µm. Figs. 31 and 32. Centris moldenkei Toro & Chiappa, 1989. Scale bars 200 µm. Figs. 33 and 34. Centris vardyorum Roig-Alsina, 2000. Scale bars 500 µm. Males 1 - Metasoma with greenish, bluish or violet metallic reflections (Figs. 4, 20, 24 and 28) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Metasoma without greenish, bluish or violet metallic reflections (Figs. 8, 12 and 16) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 522 · Zootaxa 3683 (5) © 2013 Magnolia Press VIVALLO 2 3 4 5 6 - Mesoscutum and scutellum with whitish or light brown to orange hairs; metasoma with bluish or greenish reflections (Figs. 4 and 20). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Mesoscutum and scutellum with dark brown hairs; metasoma with blue violet reflections (Figs. 24 and 28) . . . . . . . . . . . . . . 4 Mesoscutum and scutellum with whitish hairs; metasoma with bluish metallic reflections (Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centris cineraria Smith, 1854 Mesoscutum and scutellum with light brown to orange hairs; metasoma with greenish metallic reflections (Fig. 20) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centris orellanai Ruiz, 1940 Labrum semicircular (Fig. 23); clypeal disc with strong and dense punctation; apex of third mandibular tooth relatively acute (Fig. 23); eyes red (in live specimens) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centris rhodophthalma Pérez, 1911 Labrum triangular (Fig. 27); clypeal disc with dense and not very strong punctation; apex of third mandibular tooth relatively straight or truncate (Fig. 27); green eyes (in live specimens) . . . . . . . . . . . . . . . . . . . . . . Centris vardyorum Roig-Alsina, 2000 Head and mesosoma with grayish to whitish hairs (Fig. 16), usually spread throughout the body; posterior edge of second mandibular tooth and anterior edge of the third one forming an angle; apex of the third mandibular third tooth concave (Fig. 15) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centris muralis Burmeister, 1876 Head and mesosoma with dark brown (Fig. 8) or yellow hairs (Fig. 12) spread throughout the body; posterior margin of second mandibular tooth and anterior edge of the third one broadly curved; apex of the third tooth acute (Figs. 7 and 11). . . . . . . . . . 6 Body completely covered with yellow hairs (Fig. 12); mandible dark brown with an orange area in the distal half (Fig. 11); terga and sterna light brown with broad translucent distal margins . . . . . . . . . . . . . . . Centris moldenkei Toro & Chiappa, 1989 Body completely covered with dark brown hairs (Fig. 8); mandible uniformly dark brown (Fig. 7); terga and sterna dark brown without broad translucent distal margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centris escomeli Cockerell, 1926 TABLE 1. Floral records of the species of the subgenus Centris (Wagenknechtia). Species Floral record Family Reference Centris cineraria Adesmia brachysemeon Phil. Fabaceae Kalin et al. (1982)1 Adesmia conferta Hook. & Arn. Fabaceae Kalin et al. (1982) Adesmia radicifolia Clos Fabaceae Kalin et al. (1982) Adesmia sp. Fabaceae Camousseight & Barrera (1998) Vivallo et al. (2002) Anarthrophyllum cumingii F. Phil. Fabaceae Kalin et al. (1982) Rozzi et al. (1997) Alonso-Amelot (2008) Anarthrophyllum gayanum (A. Gray) Jackson Fabaceae Kalin et al. (1982)2 Astragalus curvicaulis (Clos) Reiche Fabaceae Kalin et al. (1982) Astragalus sp. Fabaceae Camousseight & Barrera (1998) Vivallo et al. (2002) ☼Calceolaria ascendens Lindl. Scrophulariaceae Vogel (1974) ☼Calceolaria corymbosa Ruiz & Pav. Scrophulariaceae Etcheverry & Valenzuela (1960) Vogel (1974) Chiappa et al. (2000) Vivallo et al. (2002) ☼Calceolaria integrifolia L. Scrophulariaceae Vogel (1974) ☼Calceolaria polyrhiza Cav. Scrophulariaceae Cosacov et al. (2012) ☼Calceolaria pritchardii Rowlee Scrophulariaceae Vogel (1974) Vivallo et al. (2002) ☼Calceolaria sp. Scrophulariaceae Ruiz (1940) Wagenknecht (1971) *Guindilia trinervis Gilles ex Hook. & Arn. Sapindaceae This work Hypochoeris sp. ☼Jovellana violacea G. Don. Asteraceae Kalin et al. (1982) Scrophulariaceae Etcheverry & Valenzuela (1960)3 Vogel (1974) Chiappa et al. (2000)3 Vivallo et al. (2002)3 ......continued on the next page REVISION OF THE BEE SUBGENUS CENTRIS (WAGENKNECHTIA) Zootaxa 3683 (5) © 2013 Magnolia Press · 523 TABLE 1. (Continued) Species Centris escomeli Floral record Family Reference Lathyrus maritimus (L.) Fr. Fabaceae Holmberg (1903) Lathyrus subandinus Phil. Fabaceae Kalin et al. (1982) Madia sativa Mol. Asteraceae Kalin et al. (1982) Phacelia secunda J. F. Gmel. Hydrophyllaceae Kalin et al. (1982) Robinia pseudoacacia L. Fabaceae Herbst (1922) Ruiz (1923, 1940) Wagenknecht (1971) Vivallo et al. (2002) Robinia viscosa Vent. Fabaceae Wagenknecht (1971) Schizanthus grahamii Gillies Solanaceae Kalin et al. (1982) Schizanthus hookeri Gillies ex Graham Solanaceae Kalin et al. (1982) Senecio bustillosianus J. Rémy Asteraceae Kalin et al. (1982) Sisyrinchium philippi Klatt. Iridaceae Kalin et al. (1982) Stachys albicaulis Lindl. Lamiaceae Kalin et al. (1982) Stachys grandidentata Lindl. Lamiaceae Etcheverry & Valenzuela (1960) Chiappa et al. (2000) Vivallo et al. (2002) Stachys macraei Benth Lamiaceae Wagenknecht (1971)4 Vivallo et al. (2002)4 Teucrium bicolor Sm. Lamiaceae Herbst (1922) Ruiz (1940) Etcheverry & Valenzuela (1960) Wagenknecht (1971) Chiappa et al. (2000) Vivallo et al. (2002) Viola portalesia Gay Violaceae Espinoza et al. (2012) Wisteria sinensis (Sims) DC. Fabaceae Etcheverry & Valenzuela (1960) Cistanthe celosioides (Barnéoud) Carolin ex Portulacaceae Hershk. Toro et al. (1996)5 Vivallo et al. (2002)6 Cristaria integerrima Phil Malvaceae Toro et al. (1996)7 Vivallo et al. (2002)7 Ipomoea purpurea L. (Roth) Convolvulaceae Wagenknecht (1971) Vivallo et al. (2002) Loasa fruticosa Urb. & Gilg. Loasaceae Toro et al. (1996) Vivallo et al. (2002) *Loasa sp. Loasaceae This work *Medicago sativa L. Fabaceae This work Nolana paradoxa Hook. Solanaceae Toro et al. (1996) Vivallo et al. (2002) *Prosopis chilensis Stuntz Leguminosae This work Caesalpiniaceae Toro et al. (1993) Toro & Chiappa (1989) Toro (2002) Vivallo et al. (2002) Medicago sativa L. Fabaceae Toro & Chiappa (1989) Prosopis chilensis Stuntz Leguminosae Toro & Chiappa (1989) Centris moldenkei Caesalpinia aphylla Phil. ......continued on the next page 524 · Zootaxa 3683 (5) © 2013 Magnolia Press VIVALLO TABLE 1. (Continued) Species Centris muralis Centris orellanai Floral record Family Reference Prosopis tamarugo F. Phil. Leguminosae Toro (2002) Toro & Chiappa (1989) Toro et al. (1993) Acacia furcata Gilles ex Hook. Leguminosae Jörgensen (1909, 1912a) Vogel (1974) Acacia sp. Leguminosae Jensen-Haarup (1908) Acalypha sp. Euphorbiaceae Cilla & Rolón (2012b) Atriplex lampa Gill ex. Moq. Chenopodiaceae Jörgensen (1909, 1912a) Vogel (1974) Bulnesia retama (Gilles ex Hook. & Arn.) Zygophyllaceae Jörgensen (19098, 1912a) Vogel (1974) Michelette & Camargo (2000) Cilla & Rolón (2012b) Caesalpinia gilliesii (Hook) D. Dietr. Caesalpiniaceae Jörgensen (1909, 1912a) Cucurbita pepo L. Cucurbitaceae Jörgensen (1912a, b) Geoffroea decorticans (Gilles ex Hook. & Arn.) Burkart Fabaceae Jörgensen (1909, 1912a, b)11 Vogel (1974)12 Grabowskia obtusa Arn. Solanaceae Jörgensen (1912a, b)13 Vogel (1974)13 Hoffmannseggia glauca (Ortega) Eifert Caesalpiniaceae Jörgensen (1912a)14 Vogel (1974) Hoffmannseggia sp. Caesalpiniaceae Jörgensen (1909)15 Larrea cuneifolia Cav. Zygophyllaceae Cilla & Rolón (2012b) Rolón & Cilla (2012) Larrea divaricata Cav. Zygophyllaceae Jörgensen (1909, 1912a) Vogel (1974) Michelette & Camargo (2000) Cilla & Rolón (2012b) Rolón & Cilla (2012) Medicago sativa L. Fabaceae Jörgensen (1909, 1912a) Mimosa sp. Leguminosae Jensen-Haarup (1908) Parkinsonia praecox (Ruiz & Pav) J. Hawkins Caesalpiniaceae Jörgensen (1909, 1912a)9 Vogel (1974)9 Prosopis alpataco Phil. Leguminosae Jörgensen (1909, 1912a) Vogel (1974) Prosopis campestris Griseb. Leguminosae Jörgensen (1909, 1912a) Vogel (1974) Psoralea higuerilla Gilles ex Hook. Fabaceae Jörgensen (1909, 1912a) Vogel (1974) Salix babylonica L. Salicaceae Jörgensen (1909, 1912a) Salix humboldtiana Willd. Salicacaeae Jörgensen (1909, 1912a)16 Senna aphylla (Cav.) H. S. Irwin & Burkart Caesalpiniaceae Jörgensen (1912a, b)10 Vogel (1974)10 Solanum elaeagnifolium Cav. Solanaceae Jensen-Haarup (1908) Vitis vinifera L. Vitaceae Jörgensen (1909, 1912a) Adesmia aphylla Clos Fabaceae Wagenknecht (1971) Vivallo et al. (2002) ......continued on the next page REVISION OF THE BEE SUBGENUS CENTRIS (WAGENKNECHTIA) Zootaxa 3683 (5) © 2013 Magnolia Press · 525 TABLE 1. (Continued) Species Centris rhodophthalma Floral record Family Reference Adesmia hystrix Phil. Fabaceae Wagenknecht (1971) Vivallo et al. (2002) Adesmia trijuga Gillies ex Hook. & Arn. Fabaceae Wagenknecht (1971) Vivallo et al. (2002) ☼Calceolaria arachnoidea Graham Scrophulariaceae Giannini et al. (2013) ☼Calceolaria lepida Phil. Scrophulariaceae Wagenknecht (1971) Vivallo et al. (2002) Acacia sp. Fabaceae Chiappa (1998) Chiappa et al. (2000) Vivallo et al. (2002) Adesmia bedwellii Skottsb. Fabaceae Wagenknecht (1971) Vivallo et al. (2002) Adesmia cinerea Clos Fabaceae Ruiz (1940) Wagenknecht (1971) Chiappa et al. (2000) Vivallo et al. (2002) Adesmia glutinosa Hook. & Arn. Fabaceae Chiappa (1998) Chiappa et al. (2000, 2000) Chiappa & Rodríguez (2001) Vivallo et al. (2002) *Adesmia melanthes Phil. Fabaceae This work Adesmia microphylla Hook & Arn. Fabaceae Wagenknecht (1971) Vivallo et al. (2002) Adesmia monosperma Clos Fabaceae Chiappa et al. (2000) Adesmia pedicellata Hook & Arn. Fabaceae Chiappa et al. (2000) Chiappa & Rodríguez (2001) Vivallo et al. (2002) Adesmia trijuga Gilles ex Hook. & Arn. Fabaceae Wagenknecht (1939, 1971) Ruiz (1940) Chiappa (1998)17 Chiappa et al. (2000)17 Vivallo et al. (2002) Balsamocarpon brevifolium Clos Leguminosae Wagenknecht (1971) Caesalpinia angulicaulis Clos Caesalpiniaceae Wagenknecht (1939, 1971) Ruiz (1940) Chiappa (1998) Chiappa et al. (2000) Vivallo et al. (2002) Dinemagonum gayanum A. Juss. Malpighiaceae Wagenknecht (1971)18 Simpson (1989) Vivallo et al. (2002)18 Dinemandra ericoides A. Juss. Malpighiaceae Cocucci et al. (1996) Fuchsia lycioides Andrews Onagraceae Wagenknecht (1971) Vivallo et al. (2002) Geoffroea decorticans (Gill. ex Hook. & Arn.) Burkart Fabaceae Wagenknecht (1971) Chiappa (1998) Chiappa et al. (2000) Chiappa & Rodríguez (2001) Vivallo et al. (2002) ......continued on the next page 526 · Zootaxa 3683 (5) © 2013 Magnolia Press VIVALLO TABLE 1. (Continued) Species Centris vardyorum Floral record ☼Krameria cistoidea Hook. & Arn. Family Reference Krameriaceae Wagenknecht (1971) Simpson et al. (1977) Vivallo et al. (2002) Lathyrus odoratus L. Fabaceae Wagenknecht (1971) Vivallo et al. (2002) Lathyrus sp. Fabaceae Ruiz (1940) Chiappa et al. (2000) Loasa tricolor Ker Gawl. Loasaceae Wagenknecht (1971) Vivallo et al. (2002) Medicago sativa L. Fabaceae Wagenknecht (1939) Oxalis gigantea Barnéoud Oxalidaceae Wagenknecht (1971) Vivallo et al. (2002) Pelargonium radula L’Hér. Geraniaceae Wagenknecht (1971) Vivallo et al. (2002) Prosopis chilensis Stuntz Leguminosae Wagenknecht (1971) Vivallo et al. (2002) Robinia pseudoacacia L. Fabaceae Ruiz (1940) Wagenknecht (1971) Chiappa (1998) Chiappa et al. (2000) Vivallo et al. (2002) Stachys albicaulis Lindl. Lamiaceae Chiappa et al. (2000) Chiappa & Rodríguez (2001) Stachys grandidentata Lindl. Lamiaceae Chiappa (1998) Vivallo et al. (2002) Tropaeolum azureum Miers Tropaeolaceae Wagenknecht (1971) Vivallo et al. (2002) Vicia benghalensis L. Fabaceae Wagenknecht (1971)19 Vivallo et al. (2002)19 Wisteria sinensis (Sims) DC. Fabaceae Wagenknecht (1939, 1971) Ruiz (1940) Chiappa et al. (2000) Hoffmannseggia glauca (Ortega) Eifert Caesalpiniaceae Jensen-Haarup (1908)20 Jörgensen (1909, 1912a, b)20 ☼Monttea aphylla (Miers) Hauman Scrophulariaceae Simpson et al. (1990) Sérsic & Cocucci (1999) Tadey (2011) Prosopis alpataco Phil. Leguminosae Tadey (2011) * New records. ☼ Oil flowers. 1 Cited as Adesmia brachysemon [sic= brachysemeon]. Uncertain identity. 3 Cited as Calceolaria violacea. 4 Cited as Stachys macroei [sic= macraei]. 5 Cited as Philippiamra celosioides. 6 Cited as Cistanthe celsoloides [sic= celosioides). 7 Cited as Cristaria foliosa Phil. 8 Cited as Buluesia [sic= Bulnesia] retama. 9 Cited as Caesalpinia praecox Ruiz & Pav. ex Hook. 2 REVISION OF THE BEE SUBGENUS CENTRIS (WAGENKNECHTIA) 10 Cited as Cassia aphylla Cav. 11 Cited as Gourliaea decorticans Hook. & Arn. 12 Cited as Gourliea [sic= Gourliaea] decorticans. 13 Cited as Grabowskya [sic= Grabowskia] obtusa. 14 Cited as Hoffmannseggia falcata [sic= falcaria]. 15 Cited as Hoffmanseggia [sic= Hoffmannseggia]. 16 Cited as Salix chilensis Mol. 17 Cited as Adesmia trifuga [sic= trijuga]. 18 Cited as Dinemagonum maculigerum Phil. 19 Cited as Vicia atropurpurea Desv. 20 Cited as Hoffmannseggia falcaria Cav. Zootaxa 3683 (5) © 2013 Magnolia Press · 527 FIGURES 35–38. Figs. 35 and 36. Centris orellanai Ruiz, 1940 (Neotype female): Fig. 35. Basitibial plate. Fig. 36. Pygidial plate. Fig. 37. Centris muralis Burmeister, 1876: Basitibial plate. Fig. 38. Centris cineraria Smith, 1854: Male genital capsule (dorsal view). Scale bars 0.5 mm. Discussion Adaptations for oil collection. Although there are no published data demonstrating the monophyly of Centris (Wagenknechtia), some characters related to the morphology of the basitibial and pygidial plates of females and the genital capsule of the males could indicate that species of this subgenus form a monophyletic group (Vivallo, in preparation). This lineage would be phylogenetically related to a small group of species, known as the “hyptidis group”, which is distributed from northeast Argentina and Paraguay to northeast Brazil (Vivallo & Melo, 2009). As observed in the species of that group (Vogel & Machado, 1991; Machado et al., 2002; Vivallo & Melo, 2009), in Centris (Wagenknechtia) is also found interesting interspecific variation in the elaiospathes. Even though this subgenus is composed of only seven species, it is possible to identify five different conformations in the combs of both pairs of elaiospathes: i) Both elaiospathes with compact combs, the former disposed along the basitarsus (Centris muralis and C. rhodophthalma share this type of oil-collecting structures, but both primary combs of the former are clearly reduced relative to those of the latter species). ii) Both elaiospathes with compact combs, the former disposed only in the distal half of the basitarsus and the latter with non-overlapping apices (Centris escomeli). 528 · Zootaxa 3683 (5) © 2013 Magnolia Press VIVALLO iii) Fore elaiospathe disposed in the distal half of the basitarsus, hind elaiospathe with the comb formed by separate hairs (Figs. 29 and 30) (Centris cineraria and C. orellanai). iv) Fore elaiospathe highly modified, virtually absent, with the primary comb disposed only on the upper half of the basitarsus, with short, dense and branched hairs on the ventral surface; hind elaiospathe with compact comb (Figs. 33 and 34) (Centris vardyorum). v) Both elaiospathes vestigial, with the hairs of both primary combs widely separated (Figs. 31 and 32) (Centris moldenkei). The morphology of the elaiospathes is related to the type of elaiophores present in the flowers that the females visit for obtaining oil (Neff & Simpson, 1981). The great diversity of forms observed in the oil-collecting structures found in Centris (Wagenknechtia), suggests the existence of a high degree of specialization in the host plants visited to obtain this resource. An interesting case is observed in the relationship between Centris and plants of the genus Calceolaria (Scrophulariaceae) (Vogel, 1974; Rasmussen & Olesen, 2000). As seen in Table 1, there are records of C. cineraria and C. orellanai visiting different species of Calceolaria throughout the distributional range of both species. Both species of bees share the same morphological pattern of oil-collecting apparatus (type iii) with the primary comb of the fore elaiospathe located in the distal half of the fore basitarsus and the comb of the hind elaiospathe formed by separate hairs (Figs. 29 and 30). This pattern was originally recognized as “type α” by Vogel (1974) and it is also found in other species groups of different subgenera that have also been recorded visiting flowers of that genus, as Centris neffi Moure, 2000, C. nigerrima and others (Vogel, 1974; Neff & Simpson, 1981; Rasmussen & Olesen, 2000; Vivallo, in preparation). On the other hand, the pattern found in Centris muralis and C. rhodophthalma (type i), originally recognized as “type β” by Vogel (1974), is broadly found in the genus. Despite the morphological similarity observed in the oilcollecting structures of both species, they have some differences, mainly in the reduction of both primary combs in C. muralis relative to those of C. rhodophthalma. As observed in Table 1, this latter species visits oil flowers, being a classic four-leg oil collector, while C. muralis apparently lost the habit of collecting this resource (Neff & Simpson, 1981). Among the many floral records of this species, there is none of oil secreting plants, although in this case the morphology of both elaiospathes indicates that structurally they may still have the capacity to collect this resource. It is interesting to note the vestigial condition of the elaiospathes of Centris moldenkei, an endemic species found in xeric areas of northern Chile. In this species, all combs of both pairs of elaiospathes are formed by widely spaced hairs (Figs. 31 and 32), making them non-functional for oil collection. The vestigial condition of the oilcollecting structures is present in several other species of Centris found in deserts of North and South America (Neff & Simpson, 1981; Zanella, 2002; Vivallo in preparation), where all those species apparently lost the habit of collecting oil (Neff & Simpson, 1981; Cappellari et al., 2011). Biogeography. The species of Centris (Wagenknechtia) are exclusively distributed in southern South America, from Arequipa (Peru) to Patagonia, on both sides of the Andes. This distribution covers various South American biogeographic subregions: south and midwest of the South American transition zone, Chacoan (south), Patagonian, Sub-Antarctic and Central Chilean (Morrone, 2001), the maximum diversity for the subgenus is found in the latter region (Coquimbo province). Centris escomeli is the species with the northernmost distribution range of the subgenus, occurring from Arequipa (Peru) to the Quebrada de Paipote (III Atacama Region, Chile), including areas near the coast to 2900 m.a.s.l. (Fig. 40). This distribution encompasses the Peruvian Coastal Desert and Atacama biogeographic provinces (Morrone, 2001). In this latter province also occurs C. moldenkei, an endemic Chilean species distributed in the north of the country between Quebrada Vitor (XV Arica y Parinacota Region) and San Pedro de Atacama (II Antofagasta Region) (Fig. 41). Both biogeographic provinces are on the western slope of the Andes, near the coast of northern Chile and southern Peru. These areas are characterized by the presence of relatively sparse vegetation, abundant xeric flora and herbaceous and/ or shrub vegetation of occasional emergence (Dinerstein et al., 1995). The vegetation of this region is favored by occasional rainfall and by the effect of the Humboldt Current in the form of fog coming from the Pacific Ocean, which permit the maintenance of areas with relatively rich plant communities located in both gullies and hills near the sea. REVISION OF THE BEE SUBGENUS CENTRIS (WAGENKNECHTIA) Zootaxa 3683 (5) © 2013 Magnolia Press · 529 Centris cineraria is the species with the largest range within the subgenus, extending from Coquimbo (IV Coquimbo Region, Chile) to Laguna Amarga (XII Magallanes y de la Antártica Chilena Region, Chile) and Santa Cruz (Argentina) (Fig. 39). These latter two localities are the southernmost distribution records not only of the species, but also for the genus. The range of this species includes the biogeographic provinces of Coquimbo, rich xeric shrub lands; Santiago, with abundant small Mediterranean forests and xeric shrub lands; and Central Patagonia, characterized by the presence of shrub steppes (Morrone, 2001). Morphologically, Centris cineraria is very similar to C. orellanai, which has a relatively restricted and partially allopatric distribution with respect of that species. Centris orellanai occurs from Baños del Toro (IV Coquimbo Region, Chile) to Los Andes (V Valparaíso Region, Chile) (Fig. 40), mainly in areas of high altitude and associated with plant species of the families Fabaceae and Scrophulariaceae. Although there are no published data to support this claim, both Centris orellanai and C. cineraria appear to be sister-species, being separated altitudinally. Distribution records of C. cineraria range from sea level to about 2000 m.a.s.l. (with a few exceptions in southern Chile), while all distribution records of C. orellanai are between 2000 and 3700 m.a.s.l. (Wagenknecht, 1971). FIGURE 39. Distribution records for Centris cineraria Smith, 1854 and C. muralis Burmeister, 1876. The Chilean species Centris rhodophthalma is distributed from Parque Nacional Llanos de Challe (III Atacama Region) to Putaendo (V Valparaíso Region), occurring almost exclusively in the biogeographic province of Coquimbo, with a few distribution records in the adjacent province of Santiago (Central Chilean Subregion) (Fig. 41). This species is found mainly in xeric areas of the central valleys and the foothills of Central Chile, associated with plants such as Adesmia pedicellata (Fabaceae), Loasa tricolor (Loasaceae) and Stachys albicaulis (Lamiaceae), among others. 530 · Zootaxa 3683 (5) © 2013 Magnolia Press VIVALLO FIGURE 40. Distribution records for Centris escomeli Cockerell, 1926 and C. orellanai Ruiz, 1940. Centris muralis and C. vardyorum are both found in Central Argentina. Centris muralis is distributed from Cafayate and Chuscha (Salta) to El Carmen (Río Negro) (Fig. 39), covering the biogeographical provinces of Chaco and Monte, which are historically closely related (Morrone, 1993, 2001). According to Cabrera (1953) and Dinerstein et al. (1995) both provinces are characterized by strata of grasses and bromeliads, along with open shrub lands dominated by species of the genera Larrea and Bulnesia (Zygophyllaceae), among others. Centris vardyorum is distributed from Chacras de Coria (Mendoza) to San Antonio (Río Negro) Argentina (Fig. 41), in the biogeographical provinces of Pampa and Monte, which are characterized by scrub-type vegetation and open savanna with grasses, herbs and shrubs (Morrone, 2001). In those provinces is possible to find plants such as Hoffmannseggia glauca (Caesalpiniaceae) and Monttea aphylla (Scrophulariaceae), which are visited by C. vardyorum for obtaining different floral resources. Conducting cladistic and biogeographic analyses of Centris (Wagenknechtia) would allow an in depth discussion of the phylogenetic relationships among the species that make up this subgenus, along with providing new evidence for increasing the knowledge of the biogeography of southern South America. REVISION OF THE BEE SUBGENUS CENTRIS (WAGENKNECHTIA) Zootaxa 3683 (5) © 2013 Magnolia Press · 531 FIGURE 41. Distribution records for Centris moldenkei Toro & Chiappa, 1989, C. rhodophthalma Pérez, 1911 and C. vardyorum Roig-Alsina, 2000. Acknowledgments I want to thank Jerome Rozen, John Ascher (AMNH), Vince F. Lee, Wojciech J. Pulawski, Robert Zuparko (CAS), John Neff (CTMI), Gabriel A. R. Melo (DZUP), Hector Vargas (IDEA), Arturo Roig-Alsina (MACN), Mario Elgueta, Fresia Rojas (MNHN), Sergio Quiroz (MHNV), Clair Villemant (MNHP), Roberto Brandão (MZUSP), Luisa Ruz (PUCV), Eduardo A. B. Almeida (RPSP), Sean G. Brady, David G. Furth, Brian Harris (USNM) and Lars Bjørn Vilhelmsen (ZMUC) for the loan of specimens and/or for the information provided about some specimens deposited in their collections. I am also grateful to David Notton for his help during my visit to the Natural History Museum (London, England), José Salamanca (CSPN), Juan Pablo Botero (MNRJ), Fernando Fernández (Universidad Nacional de Colombia, Bogotá, Colombia) and Mariana Taniguchi for their help during the development of this publication and to Vitor Nardino (Taxonline) for taking the pictures of the specimens studied here. I am also grateful to John Neff and Laurence Packer for comments and suggestions that helped to improve this article. Financial support was provided by Fundação de Apoio à Pesquisa do Rio de Janeiro, Brasil (FAPERJ, grant INST-110.793/2012). This publication is dedicated to J.M. References Alonso-Melot, M.E. (2008) High altitude plants, chemistry of acclimation and adaptation. 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