Phylogenetic Relationships Of A New Genus Of Calliopsine Bees From Peru, With A Review Of Spinoliella Ashmead (Hymenoptera: Andrenidae) Author Gonzalez, Victor H. Author Smith-Pardo, Allan H. Author Engel, Michael S. text Bulletin of the American Museum of Natural History 2017 2017-04-13 2017 412 1 72 http://www.bioone.org/doi/10.1206/0003-0090-412.1.1 journal article 10.1206/0003-0090-412.1.1 0003-0090 5407146 Spinoliella rufiventris Toro and Ruz Figure 20A–E Spinoliella ( Peniella ) rufiventris Toro and Ruz, 1972a: 151 ( holotype ♂ , SEMC, seen: Las Trancas, Ñuble, Chile ). Spinoliella ( Peniella ) karhadra Rodríguez, Toro, and Ruz, 2001: 104 ( holotype ♂ , PUCV, seen: Hualqui, Concepción, Region VIII, Chile ). New synonymy. DIAGNOSIS: This species can be recognized by the following combination of traits: female pygidial plate broad, with lateral margins converging toward apex at a 50° angle, apically bifid; frons, mesoscutum, and mesoscutellum with minute, contiguous punctures (fig. 20A, B), often shallow and faint on mesoscutellar disc; female metabasitibial plate distinct (barely indicated in the male: fig. 20D), slightly depressed on disc but not delimited by a strong border or carina (fig. 20C); and metasoma light reddish brown, often distinctly contrasting with dark brown to black head and mesosoma. It resembles S. nomadoides and S . ruzi in the body color and sculpturing of frons, mesoscutum, and mesoscutellum; however, in S. nomadoides the female pygidial plate is apically truncate, not bifid; the metabasitibial plate is delimited by a strong carina in both sexes; the female outer metatibial spur is distinctly curved apically (straight in S. rufiventris ); and the male mandible is arcuate, with a small preapical tooth (straight and without a preapical tooth in S. rufiventris ). The broader pygidial plate FIGURE 20. Spinoliella rufiventris Toro and Ruz (A-E) from Ñuble, Chile (SEMC) and male of Spinoliella ruzi Compagnucci (F) from Nahuel Huapi, Argentina (PCYU). A. Detail of female frons of S. rufiventris . B. Detail of female mesoscutum and mesoscutellum in dorsal view of S. rufiventris . C. Female metabasitibial plate of S. rufiventris . D. Male metabasitibial plate of S. rufiventris . E. Male metasomal S8 of S. rufiventris . F. Male metasomal S8 of S . ruzi . of the female and more robust median process of male S8 (cf. figs. 20E and 20F) separate S . rufiventris from S . ruzi . TYPE MATERIAL EXAMINED ( n = 10♀♀ , 8♂♂ ): Holotype ♂ [ rufiventris ], Chile , Las Trancas , Prov. Ñuble, 12-14-52 [ 14 December 1952 ], L.E. Peña ( SEMC ). Allotype ♀ [ rufiventris ], Las Trancas , Prov. Ñuble, 12-14-52 [ 14 December 1952 ], L.E. Peña ( SEMC ). Holotype ♂ [ karhadra ], Chile , Región III, Hualqui , 2-xi-1985 , C. Vial ( PUCV ). Allotype ♀ [ karhadra ], same data as holotype of S . karhadra (PUCV) . Paratypes [ rufiventris ]: O’Higgins (R-VI): 4♀♀ , 2♂♂ , Las Cabras [ Region VI ], XII 10–23 1954 [ 10–23 December 1954 ], 1100–1460 m , L.E Peña ( SEMC ) ; Bío Bío (R-VIII): 4♀♀ , 4♂♂ , Las Trancas , Prov. Ñuble, 12-14-52 [ 14 December 1952 ], L.E. Peña ( SEMC ) . MATERIAL EXAMINED ( n = 26♀♀ , 44♂♂ ): CHILE : O’Higgins (R-VI): 2♂♂ , Las Cabras [ Region VI ], XII 10–23 1954 [ 10–23 December 1954 ], 1100–1460 m , L.E Peña ( SEMC ) ; Maule (R-VII): 1♀ , Region VII, W of Laguna del Maule , 35°57′461″, 070°34′844″, 29.xii.06 [ 29 December 2006 ], L. Packer ( PCYU ) ; 1♂ , Region VII, W of L. Maule , S35.96901 , W70.56940 , 1982 m , 4.i.2009 [ 4 January 2009 ], L. Packer , PCYU- CHI09-6-4-002 ( PCYU ) ; 1♀ , E of Laguna del Maule , 1359 m , -35.90163 , -70.64261 , 6.i.2013 [ 6 January 2013 ], L. Packer , ex: Primula (PCYU) ; 1♂ , same as previous except 160.120 km , 2505 m , -35.99282 , -70.39911 , 1.i.2013 [ 1 January 2013 ], blue pan traps , L. Packer , R. Smith ( PCYU ) ; 1♂ , Laguna del Maule , -36.020519 , -70.500163 , 2364 m , 29.xii.2006 – 27.i.2007 [ 29 December 2006 – 27 January 2007 ], pans, L. Packer , A.I. & M. Gravel ( PCYU ) ; 1♂ , same as previous except 1955 m , -35.96881 , -70.56934 , 6.i.2012 [ 6 January 2012 ], L. Packer , R. Smith ( PCYU ) ; 1♂ , N of Laguna del Maule , 29. xii.2006 – 27.i.2007 [ 29 December 2006 – 27 January 2007 ], pans, L. Packer , A.I. & M. Gravel ( PCYU ) ; 2♂♂ , Laguna del Teno , -35.10666 , -70.53926 , 26.i.2016 [ 26 January 2016 ], 2548 m , L. Packer ( PCYU ) ; 4♂♂ , Curicó prov. El Planchón , 34°08′57.8″S , 70°31.49.2″W, 2400, 1–5. ii.2003 [ 1–5 February 2003 ], leg. A. Ugarte P. ( PCYU ) ; Bío Bío (R-VIII): 7♀♀ , 12♂♂ , Shangri-La [ Region VIII], -36.89764 , -74.76026 , 1335 m , 27.xii.2006 [ 27 December 2006 ], pan traps , L. Packer ( 6♀♀ , 11♂♂ PCYU , 1♀ , 1♂ SEMC ) ; 1♂ , same as previous except 1331 m , -36.89719 , -71.47636 , 5.i.2013 [ 5 January 2013 ], L. Packer , R. Smith ( PCYU ) ; 1♀ , 1♂ , Region VIII, Shangri-La, 4436 ft , 36°53′897″, 071°28′637″, 11.xii.06 [ 11 December 2006 ], L. Packer pans ( PCYU ) ; 1♂ , Shangrilá , XII-1998 [ December 1998 ], A. Ugarte P ( SEMC ) ; 2♀♀ , 1♂ , Ñuble , Chillán , Las Trancas , 1100 m , Jan 1987 , L. Peña ( AMNH ) ; 2♂♂ , Las Trancas , Prov. Ñuble , 12-14-52, L.E. Peña [ 14 December 1952 ] ( SEMC ) ; 5♀♀ , 4♂♂ , Termas de Chillan , -36.916642 , -71.425488 , 1543 m , 24.i.2007 [ 24 January 2007 ], L. Packer ( PCYU ) ; 4♀♀ , 4♂♂ , Las Trancas , 1200 m , L. Peña , on cactus flowers ( SEMC ) ; 1♀ , 2♂♂ , Las Trancas , 78 km E Chillan ; 36°54.5′S , 71°29′W , 12.XII.2003 [ 12 December 2003 ]; pan trap; FD Parker; FDP#3748, 3762, 3772 ( BBSL ); 2♂♂ , Las Trancas , 1300 m , December 13–16, 1976 , L.E. Peña ( SEMC ) ; 1♀ , Las Trancas , SE Recinto , in Chillan area , January 1987 , Luis E. Peña ( SEMC ) ; 1♀ , Chillan area , Shangri La , Las Trancas E. Recinto , Jan 20, 1979 , Luis E. Peña ( SEMC ) ; 1♀ , same as previous except January 19–22, 1979 ( SEMC ) ; Araucanía (R-IX): 1♀ , Region IX, 37.809°S , 71.016°W , Parque Nacional de Nahuelbuta , 3860 ft , 6–9.i.2000 [ 6–9 January 2000 ], W. Webb & D. Yeates ( PCYU ) ; Región Metropolitana ( RM ): 1♂ , Region Metro [Santiago Metropolitan Region, Cordillera Province], Termas Valle de Colima , S33.81533 , W70.00726 , 2355 m , 7.i.2009 [ 7 January 2009 ], L. Packer ( PCYU ) . DISTRIBUTION: Chile : O’Higgins (R-VI): Cachapoal , Colchagua ; Maule (R-VII): Curicó ; Bío Bío (R-VIII): Concepción , Ñuble ; Araucanía (R-IX): Malleco ; Región Metropolitana ( RM ): Cordillera . FLORAL RECORDS: Primula sp. (Primulaceae) ; one series of males and females were captured on flowers of an unidentified cactus ( Cactaceae ). COMMENTS: This species seems to be highly variable in body size, maculation (e.g., distal margin of mesoscutellum and metanotum often without maculation), shape of the facial fovea, clypeal basal margin, and inner orbits of the compound eyes. The features originally used to distinguish S . karhadra as separate species from S . rufiventris are all among these variable features and with larger series it is apparent that the two intergrade. Spinoliella karhadra was initially conceived to be slightly larger, with ventrally divergent compound eyes, and a clypeal basal margin slightly convex. These are all highly variable across S. rufiventris , even among specimens from the same locality as indicated by Toro and Ruz (1972a: 154) and observed in our study. For example, we found that among the paratypes of S. rufiventris deposited at SEMC, larger bees tend to have the inner compound eye orbits diverging ventrally and a shorter clypeus (more than 4× broader than long), basally straight. The same phenomenon was observed in males of S. herbsti . We also found that in some females the facial fovea is about the same size across its length, with the clypeus basally slightly convex. Thus, such specimens could be identified as S. karhadra , but we did not find other features (i.e., sculpturing, shape of pygidial and metabasitibial plates) that reliably separate them from the typical S. rufiventris . Additionally, there are specimens with intermediate features, such as a normal body size and inner orbits slightly divergent ventrally. These observations suggest that the types of S. karhadra are nothing more than large specimens of S. rufiventris , principally in terms of head width (head widths of S . karhadra are 2.8 mm in the holotype male and 2.2 mm in the allotype female, while in smaller S. rufiventris these measurements are 2.1 mm and 1.8 mm in the male and female, respectively, but intermediates are known). Intraspecific variations in body size, particularly in the head and often resulting from allometry, are not unusual across bees (e.g., Sakagami and Moure, 1965 ; Danforth, 1991 ; Packer et al., 2003 ; Engel, 2008 ; Engel et al., 2012 ). Therefore, it is likely that the ventrally divergent compound eyes as well as the clypeus basally more pronouncedly convex are perhaps modifications related with a large body size. Given that the holotype and allotype of S . karhadra fall within the overall variation, there are no differences in the male terminalia, and this putative species is sympatric with S . rufiventris , we have considered the former to be a junior synonym of the latter.