Hymenoptera: Apidae Sl: Protandrenini
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ZOOLOGIA 31 (5): 503–515, October, 2014 http://dx.doi.org/10.1590/S1984-46702014000500011 Taxonomic revision of Parasarus (Hymenoptera: Apidae s.l.: Protandrenini), a South American genus of bees Kelli dos Santos Ramos Museu de Zoologia, Universidade de São Paulo. Avenida Nazaré, 04263-000 São Paulo, SP, Brazil. E-mail: [email protected] ABSTRACT. Parasarus Ruz, 1993 comprises small black bees (3-5 mm long) endemic to xeric regions of South America, mainly along of the Andean Cordillera. Prior to this study, the genus included only the type-species P. atacamensis Ruz, 1993 (from northern Chile) which has mesoscutum strongly reticulated and inner hind tibial spur curved apically. In this paper, a taxonomic revision of Parasarus is presented and two new species are described: P. specularis sp. nov. (from central to northwest Argentina) diagnosed mainly by pygidial plate of female extremely acute apically and labral plate yellow in male; and P. spiniventris sp. nov. (only recorded from central portion of Chile) diagnosed by antennal socket below middle of face, subantennal area as long as broad, metapostnotum smooth, and sternum 3 of male with a tuft of stiff hairs. The morphological variation related to the type-species was studied and not considered sufficiently to recog- nized distinct species into P. atacamensis. Distribution maps, floral associations, key to species of the genus, and illustra- tions of general external morphology and genitalia are also provided. KEY WORDS. Andreninae; Apoidea; new species; taxonomy. Protandrenini is the most diverse group of Andreninae MATERIAL AND METHODS bees (Hymenoptera: Apidae s.l.) in the Neotropical region by number of genera with more than 400 species described (MOURE The material examined belongs to the following collec- et al. 2012, ASCHER & PICKERING 2014). The tribe is restricted to the tions: AMNH – American Museum of Natural History, New York, Americas, being diverse in temperate and xeric areas (MICHENER USA; DZUP – Departamento de Zoologia, Universidade Fed- 2007). Among Protandrenini genera of South America, Parasarus eral do Paraná, Curitiba, Brazil; IMLA – Fundación Miguel Lillo, Ruz, 1993 comprises small black bees (3-5 mm long) endemic to San Miguel de Tucumán, Argentina; MZSP – Museu de Zoologia, xeric regions of Argentina and Chile (RUZ & ROZEN 1993, MICHENER Universidade de São Paulo, São Paulo, Brazil; PCYU – Packer 2007). Parasarus was proposed by RUZ (in RUZ & ROZEN 1993) for Collection at York University, Toronto, Canadá; PUCV – a single species – P. atacamensis Ruz, 1993. Nests of this species Pontificia Universidad Católica de Valparaíso, Valparaíso, Chile; are built in soil and shared by a number of females, and parasit- and RPSP – Departamento de Biologia, Faculdade de Filosofia, ized by bees of the genus Kelita Sandhouse, 1943 (Apidae: Ciências e Letras de Ribeirão Preto, Universidade de São Paulo, Nomadini) (RUZ & ROZEN 1993). Species of Parasarus can be col- Ribeirão Preto, Brazil. lected from September to February and are apparently polylectic General morphological terminology follows URBAN (1967) in relation to floral hosts. Males fly actively around flowers near and MICHENER (2007); labral plate is characterized by an elevated nesting sites, searching for mates (RUZ & ROZEN 1993). Although and glabrous area in the central portion of the labrum; antennal the mature larvae exhibit unique features (such as a globose flagellomeres are indicated as F1-F11, and metasomal terga and antennal prominence and minute spiracles), they generally re- sterna as T1-T7 and S1-S8, respectively. The antennal flagellomeres semble other Protandrenini larvae in having dorsal thoracic tu- and labial palpomeres were numbered from the base to the apex. bercles, well-developed maxillae, large maxillary palpi, and The density of punctation refers to the relative puncture diam- strongly recessed labial region (RUZ & ROZEN 1993). eter, pd (e.g., <2 pd: less than 2x the puncture diameter between Parasarus remained monotypic until the present study, punctures). All measurements are given in millimeters. although RUZ & ROZEN (1993) mentioned the existence of addi- The labels of the specimens examined are exactly tran- tional undescribed species from Chile and Argentina, and called scribed in the sections Type material and Additional material, attention to morphological variability observed among popu- where one inverted bar (\) indicates different lines on the la- lations of P. atacamensis. In this paper, a taxonomic revision of bel and quotation marks (“) indicate different labels of the same Parasarus is presented with description of two new species and specimen. Photographs were taken with a Leica video camera the study of morphological variation related to the type-spe- (DFC 295) attached to a Leica stereomicroscope (M205C). Se- cies. Floral associations, distribution map, illustrations, and a ries of images were combined in the software LEICA LAS (Leica key to species are also provided. Application Suite V3.6.0) and CombineZP to produce confo- 2014 Sociedade Brasileira de Zoologia | www.sbzoologia.org.br | www.scielo.br/zool All content of the journal, except where identified, is licensed under a Creative Commons attribution-type BY-NC. 504 K.S. Ramos cal images. Male genitalia and associated sterna were detached lum of males as long as maximum head width (similar to fe- from the metasoma, cleared in a 10% KOH solution for 16h, males); pronotal lobe almost black in both sexes; metapostnotum neutralized in acetic acid, and stored in a vial with glycerin. slightly depressed with some minute hairs laterally (Figs 15-20); Floral records for species of Parasarus were compiled from la- lateral line of T1 evident; metasomal terga of males with bels of examined specimens and cited references in the postgradular area weakly depressed (Fig. 28); S8 of males with catalographic list. The plant names are according to the Inter- enlarged disc (Figs 31, 34, and 37); volsella with short cuspis; national Plant Names Index (www.ipni.org). Distribution maps gonostylus with sparse pilosity (Figs 39-47); penis valve acute were produced using the software Quantum GIS (http:// distally and fused basally. www.qgis.org); the locality coordinates were traced using the Remarks. The phylogenetic relationships among genera Global Gazetteer Version 2.2 (http://www.fallingrain.com/ of Protandrenini are poorly understood and the current generic world/index.html) when not provided on specimen labels. classification is not consensual. Parasarus was recognized as a subgenus of the North American genus Protandrena Cockerel, TAXONOMY 1896 by MICHENER (2000, 2007). However, morphological and molecular phylogenetic analyses provided substantial support Parasarus Ruz, 1993 for recognizing Parasarus as a phylogenetic independent lineage from Protandrena (J.S. Ascher, unpubl. data, K.S. Ramos, unpubl. Figs 1-50 data). Parasarus differs from Protandrena s.s. mainly in having Parasarus Ruz, 1993 in Ruz & Rozen, 1993: 2. Type-species: integument with shallow and sparse punctures, face without Parasarus atacamensis Ruz, 1993 by original designation. yellow marks, mid tibial spur of females finely serrate, metasoma Parasarus; Neff & Rozen, 1995: 13 (comparative note); Rozen & of males broader than mesosoma, metasomal terga without basal Ruz, 1995: 9 (citation); Rozen, 1997: 346 (citation); Moure et hair bands, postgradular area of metasomal terga of males weakly al., 2007: 34 (catalog); Montalva & Ruz, 2010: 20 (distribution). depressed, and S8 of males with broad disc (not constricted in Protandrena (Parasarus); Michener, 2000: 269 (systematics, distri- the middle). Males of Parasarus exhibit intraspecific variation bution); Fernández, 2001: 105 (citation); Fernández, 2002: 127 in the width of head, which may vary from narrower to broader (citation); Ruz & Chiappa, 2004: 790 (comparative note); Ramos than mesosoma. These wide-headed males look like those of & Melo, 2006: 43 (citation); Michener, 2007: 281 (systematics, the genus Cephalurgus Moure & Lucas de Oliveira, 1962, but they distribution); Gonzalez & Engel, 2011: 59 (comparative note). differ principally by the lateral foveae of T2 weakly depressed, Diagnosis. The diagnostic characters presented by RUZ & supraclypeal area slightly produced, genital capsule without basal ROZEN (1993) and MICHENER (2000, 2007) that are common to all sclerite, and gonocoxite without an oblique lateral impression species of Parasarus here recognized are the following: small body (Figs 39-47). size (3-5 mm); integument predominantly black or dark brown Variation. As mentioned above, males of Parasarus show with some yellow markings on legs; pubescence in general short variation in head width. In comparison to males with small head, and white; face without yellow marks in either sex; glossa shorter wide-headed specimens have more sparsely distributed punctures than prementum; segment 1 of labial palpus with similar length and hairs, distinctly longer mandibles, vertex more swollen above or shorter than segments 2-4 combined; lower half of face of ocelli, and gena broader than compound eyes (Figs 5 and 6). males densely pilose; anterior tentorial pit at intersection be- Distribution. Species of Parasarus are restricted to xeric tween outer subantennal and epistomal sutures; supraclypeal regions of South America, occurring from lowlands to high area weakly produced between antennal sockets; mesosoma mountains (above 4000 m) of northern Chile to southern Ar- sparsely punctate; submarginal