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Home , Diadasia, Emphorini, Ptilothrix

Revista de la Sociedad Entomológica Argentina ISSN: 0373-5680 [email protected] Sociedad Entomológica Argentina Argentina

ROIG ALSINA, Arturo A review of the albidohirta group of (: : ) Revista de la Sociedad Entomológica Argentina, vol. 66, núm. 3-4, 2007, pp. 113-119 Sociedad Entomológica Argentina Buenos Aires, Argentina

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How to cite Complete issue Scientific Information System More information about this article Network of Scientific Journals from Latin America, the Caribbean, Spain and Portugal Journal's homepage in redalyc.org Non-profit academic project, developed under the open access initiative ISSN 0373-5680 Rev. Soc. Entomol. Argent. 66 (3-4): 113-119, 2007 113

A review of the albidohirta group of Ptilothrix (Hymenoptera: Apidae: Emphorini)

ROIG ALSINA, Arturo

Museo Argentino de Ciencias Naturales «Bernardino Rivadavia,» Av. Angel Gallardo 470, 1405 Buenos Aires, Argentina; e-mail: [email protected]

Revisión del grupo albidohirta de Ptilothrix (Hymenoptera: Apidae: Emphorini)

„ RESUMEN. El grupo albidohirta de Ptilothrix Smith incluye tres especies: Ptilothrix albidohirta Brèthes, P. concolor sp. nov. y P. nemoralis sp. nov. Este grupo se conoce del oeste y norte de la Argentina y del centro de Bolivia. Una de sus especies, P. albidohirta, llega hasta los 3.600 m s.n.m. en la cordillera de los Andes, en tanto que las otras dos especies habitan en el llano. Se caracteriza el grupo y se describen e ilustran las especies.

PALABRAS CLAVE. Abejas. Emphorini. Argentina. Bolivia.

„ ABSTRACT. The albidohirta group of Ptilothrix Smith includes three species: Ptilothrix albidohirta Brèthes, P. concolor sp. nov., and P. nemoralis sp. nov. The group is known from northern and western Argentina and from central Bolivia. One of its species, P. albidohirta, reaches high altitudes in the Andes mountains, up to 3600 m a.s.l., while the other two species occur in the lowlands. A characterization of the group, descriptions of the species, and illustrations are provided.

KEY WORDS. . Emphorini. Argentina. Bolivia.

INTRODUCTION occurs in temperate South America, but the number of species is difficult to assess, since The Ptilothrix Smith is a member of Ptilothrix has not been reviewed to date. the Emphorini, a tribe of bees restricted to Species of Ptilothrix are solitary, the Western Hemisphere (Michener, 2000). burrowing bees. Many species are The tribe comprises ten genera (Roig-Alsina, oligolectic, and have been associated with 1999), six of which are restricted to South plants mainly of the families , America, two are broadly distributed , , and throughout the continent, and two ( Cactaceae (Linsley et al., 1956; Rust, 1980; Patton and Ptilothrix Smith) have numerous Hazeldine, 1997). species in North and South America, but are According to Roig-Alsina (1994) and Sipes scarce or not represented in Central America. & Tepedino (2005), Ptilothrix shares derived The species of Ptilothrix occur from north- features with Diadasina Moure and central eastern U.S.A. to the state of Morelos Alepidosceles Moure, suggesting that these in Mexico (Michener et al., 1994), and from three genera are closely related. Ptilothrix is northern Brazil to northern Patagonia in distinguished, among other emphorines, by Argentina. The largest diversity of the genus a suite of characters that includes the lack of

Recibido: 19-IV-2007; aceptado: 28-VIII-2007 114 Rev. Soc. Entomol. Argent. 66 (3-4): 113-119, 2007

arolia, the elongate first flagellomere, the MATERIAL AND METHODS bowed, elevated upper margin of the clypeus coupled with a short proboscis, the reduced Terminology for structures follows apical projection of the female hind Michener (1944, 2000), except that basitarsus, and the type of scopal hairs on metapostnotum is used instead of propodeal the outer surface of the hind tibia, which have triangle (Brothers, 1976). The maximum the branches not retrorse, but arising at acute diameter of the median ocellus (MOD) is used angles to the axes of the hairs and directed as a reference to express the length of the apicad. pubescence and other structures. In the The present contribution deals with a descriptions, the metasomal terga (T) and small group of species restricted to northern sterna (S) are identified with Arabic numerals. Argentina and Bolivia. Although the three Specimens studied are deposited in the species included in the group are following institutions: Museo Argentino de morphologically homogeneous, they are Ciencias Naturales «Bernardino Rivadavia,» strikingly different in their color pattern and Buenos Aires (MACN); Museo de La Plata, vestiture. La Plata (MLP); Instituto y Fundación Miguel

Figs. 1-9. Figs. 1-3, Ptilothrix concolor n. sp., male. 1, S7, ventral (left) and dorsal (right) views. 2, S8, ventral view. 3, apex of T7. Figs. 4-6. Ptilothrix albidohirta Brèthes, male. 4, S7, ventral (left) and dorsal (right) views. 5, S8, ventral view. 6, apex of T7. Figs. 7-9, hairs of center of outer surface of hind tibia of female. 7, P. albidohirta Brèthes. 8, P. concolor n. sp. 9, P. nemoralis n. sp. Scale bars = 0.1 mm. ROIG ALSINA, A. The albidohirta group of Ptilothrix 115

Fig. 10. Distribution map of the species of the albidohirta group: P. albidohirta Brèthes (dots), P. concolor n. sp. (stars; open star indicates imprecise locality in La Rioja province), and P. nemoralis n. sp. (diamonds).

Lillo, Tucumán (IFML); American Museum of and P. nemoralis in the Chaco region. Natural History, New York (AMNH); The group is characterized by the slender, Zoologische Staatssammlung, München short-branched scopal hairs on the outer (ZSM). Acronyms are used to indicate surface of the hind tibia of the female, by the depositories of the specimens. lack of an apical white brush of hairs on the hind tibia, and by the digitiform projection RESULTS of the inner lobe of the seventh sternum of the male (although males of P. nemoralis are albidohirta group as yet unknown). The three species differ considerably in This species-group includes Ptilothrix their appearance. Ptilothrix albidohirta albidohirta Brèthes and the two new species resembles Diadasia baeri (Vachal), another P. concolor and P. nemoralis. The group is hirsute, pale-colored emphorine, with which known from northern and western Argentina it is sympatric in the Andes mountains. and from central Bolivia (Fig. 10). One of the Ptilothrix concolor resembles a large, species, P. albidohirta, occurs in the Puna ordinary Diadasia, due to its short, appressed, region, reaching high altitudes; records for yellowish-brown vestiture. On the contrary, this species range from 2200 m to 3600 m Ptilothrix nemoralis bears a color pattern that a.s.l. The other two species occur in the is typical for a large number of species of lowlands, P. concolor in the Monte region Ptilothrix, with a rufous dorsum of the thorax 116 Rev. Soc. Entomol. Argent. 66 (3-4): 113-119, 2007

and a yellow banded metasoma. shiny, elevated, giving rugose appearance. Punctation around and above antennal Ptilothrix albidohirta Brèthes, 1910 sockets similar to that of clypeus, but with (Figs. 4-7, 10) tessellate interspaces; rest of head more finely punctured; vertex around ocelli finely Ptilothrix albidohirta Brèthes, 1910: 295-296 tessellate, except small shiny area close to (Holotype male, 7892, Jujuy, Argentina, lateral ocellus. Punctures on scutum, MACN, examined); Schrottky, 1913: 254; scutellum, and mesopleuron dense, on center Hazeldine, 1997: 33-35, figs. 16-17 of scutum 0.2-1.0 diameters apart, with (biology); Roig-Alsina, 1999: 25. weakly tessellate interspaces. Metapostnotum tessellate. Metasomal terga tessellate between Diagnosis. This species is distinguished punctures. by its hirsute vestiture all over the body. The Structure. Inner orbits nearly straight and metasoma of both sexes is clothed with long, parallel. Ocellocular and interocellar erect hairs, lacking the appressed bands distances subequal. Clypeus 1.9-2.0 times characteristic of many other Ptilothrix. It is wider than long, protuberant in lateral view also characterized by the color of the for a distance 0.65 times maximum eye vestiture, greyish on the head and mesosoma, width. Labrum trapeziform, with pointed and yellowish to orange on the metasoma; apex, 1.5 times as broad as median length. the vestiture of the legs is pale in males, but Interantennal distance 2.5-2.8 times mostly black in females. antennocular distance. First flagellomere 2.2 times longer than its apical width; proportion Redescription. Female. Length: 10.3- of lengths of scape, pedicel and proximal 12.8 mm; length of forewing: 8.8-10.3 mm. three flagellomeres 1.7:0.5:1:0.3:0.35. Color: Black, except middle of mandible, apex of tarsi, and translucent outer margin of Male. Length: 8.0-13.0 mm (holotype tegula reddish brown. Membrane of wings: 10.3 mm); length of forewing: 8.5-10.5 mm slightly infuscate; pterostigma and veins: dark (holotype, 8.7 mm). Color, pubescence and brown to black. sculpture: similar to those of female, except Pubescence. Hairs: gray to pale as follows. Hairs: gray to pale yellowish- yellowish-brown on head, mesosoma and T1; brown all over the body, including legs and hairs grayish, partly dusky, on underside of metasoma. Hairs on head and thorax longer, coxae, trochanters and front and mid femora, those on scutum 3.5-4.3 times MOD, on rest of legs with hairs black, including scopal mesopleuron 3.3-4.3 times MOD. Hairs on hairs. Hairs: yellowish brown (with orange metasomal terga: erect, long, on T2 2.2-2.8 hue in fresh specimens) on T2-T6 and on times MOD. S2-S4 with long, erect hairs, and metasomal sterna; prepygidial fimbria orange. apical fringes poorly defined; S5 with hairs Pubescence of head and thorax: long and short medially and longer laterally. S6 bare erect, hairs on center of scutum 2.5-3.0 times basally and laterally, apically with dense MOD, hairs on mesopleuron also 2.5-3.0 median tuft of erect hairs. times MOD, hairs on postero-lateral angles Structure. Labrum rectangular, with of propodeum 3.0-4.0 times MOD. polished, rounded median apical projection. Metapostnotum: clothed with sparse hairs, First flagellomere 2.1 times as long as its 0.8-1.2 times MOD. Tibial scopa with hairs apical width; proportion of lengths of scape, on outer surface 1.8-2.0 times MOD, with pedicel and proximal three flagellomeres rachis slender and 16-20 branches directed 1.9:0.4:1:0.5:0.55. T7 ending in two sharp apicad (Fig. 7). Metasomal terga: densely apical points separated by notch with straight covered with erect hairs, 1.5-2.2 times MOD, base (Fig. 6). S7 and S8 as in Figs. 4-5. completely lacking appressed hairs. Gonostylus filiform. Sculpture. Clypeus and labrum with dense punctures; punctures on disc of clypeus Material examined. ARGENTINA. 0.3-0.5 diameters apart, with interspaces Jujuy: 2 males, Abrapampa, 3400 m, 5-III- ROIG ALSINA, A. The albidohirta group of Ptilothrix 117

1960, R. Golbach (IFML); 1 male, Iturbe, coxae brownish, tibial spurs and tarsi reddish 3300 m, 8-III-1960, R. Golbach (IFML); 1 brown; tegula translucent, yellowish. Wings: female, 1 male, Humahuaca, 24-I-1973, L. hyaline, except slightly infuscate on marginal Stange (IFML); 2 females, 1 male, Tilcara, cell; veins and pterostigma dark brown. 2400 m, 20-III-1967, L. Stange & A. Willink Pubescence. Uniformly pale, gray to (IFML); 1 female, 3 males, Tilcara, 9-III-1965, whitish on face, underside of thorax and A. Willink (IFML); 1 male, Tilcara, II-1989, pleurae, pale yellowish-brown on dorsum of Fritz (AMNH); 1 female, 1 male, Uquía, 2800 thorax, metasoma and most of legs; T2-T4 m, Quebrada de Humahuaca, 17-III-1978, with weakly defined narrow yellow apical A. Willink & P. Fidalgo (IFML); 8 females, 8 bands, due to denser pubescence. Outer males, Maimará, 19-III-1990, A. Roig A. surface of hind tibia without distinct tuft of (MACN); 2 females, 9 males, Maimará, 7-III- white hairs. Prepygidial and pygidial fimbriae 1965, A. Willink (IFML); 8 females, reddish brown. Pubescence of head and Purmamarca, 15-III-1978, A. Willink & P. thorax dense, short on scutum, hairs on center Fidalgo (IFML); 1 female, 5 males,Volcán, of scutum 0.7-1.2 times MOD, hairs on 2100 m, 12-III-1960, R. Golbach (IFML). mesopleuron longer, 1..5-2.5 times MOD, Salta: 4 males, Amblayo, 2500 m, 30-I-1945, hairs on postero-lateral angles of propodeum F. Monrós (MLP); 23 females, 5 males, 3 times MOD. Metapostnotum clothed with Chorrillos, 15-III-1993, P. Hazeldine (MACN); sparser hairs, 0.8-1.5 times MOD. Tibial 8 females, 9 males, Tastil, 3000 to 3600 m, scopa with hairs on outer surface 1.4-1.8 III-1985, III-1986, III-1994, I-1995, III-1996, times MOD, with rachis slender and 14-18 M. Fritz (MACN); 9 females, 4 males, Tastil, branches directed apicad (Fig. 8). Metasomal III-1985, III-1988, II-1992, M. Fritz (AMNH); terga densely covered with appressed, short 2 females, El Alisal, III-1988 and IV-1988, M. hairs, 0.4-0.7 times MOD, intermixed with Fritz (AMNH); 14 females, 7 males, El Alisal, sparse, erect, long hairs; appressed hairs II-1983, III-1985, IV-1988, II-1989, III-1993, denser on posterior margin of terga, forming III-1994, M. Fritz (MACN); 1 female, El weakly defined apical bands. Maray, 2200 m, M. Fritz (MACN). La Rioja: Sculpture. Clypeus and labrum with 2 males, Cuesta de la Aguadita, 9 km S dense, regularly distributed punctures; those Angulos, 25-II-1978, A. Willink & E. on disc of clypeus 0.3-0.5 diameters apart; Domínguez (IFML). BOLIVIA. La Paz: 1 intespaces shiny. Punctation around and female, Irpavi, 3200 m, 4-IV-1938, F. above antennal sockets similar to that of Tippmann (ZSM). Cochabamba: 2 males, clypeus; rest of head more finely punctured; Morochata, 2900 m, R. Zischka (MLP); 1 vertex around ocelli smooth. Punctures on female, Cochabamba, R. Zischka (MLP); 1 scutum, scutellum, and mesopleuron similar male, Cochabamba, 2500 m, W. Wittmer to those of clypeus, 0.5-1.0 diameters apart, (MLP). and also with shiny interspaces. Metapostnotum with shiny interspaces Ptilothrix concolor sp. nov. between punctures. Metasomal terga finely (Figs. 1-3, 8, 10) punctured. Structure. Inner orbits nearly straight, Diagnosis. This species is readily converging below, upper to lower interocular separated from all other Ptilothrix by its distance 1.1:1. Ocellocular distance smaller homogeneous pale-colored vestiture, short than interocellar distance (0.9:1). Clypeus 1.7 and appressed on the metasoma, similar to times wider than long, protuberant in lateral that of many species of Diadasia. view for a distance 0.65 times maximum eye width. Labrum trapeziform, with pointed Description. Holotype female. Length: apex, 1.5 times as broad as median length; 13.5 mm (paratypes, 11.0-12.0 mm); length apex truncate with antero-apical margin of forewing 10.0 mm (paratypes, 9.5-10.3 carinate. Interantennal distance 3 times mm). Color. Black, except base of mandible, antennocular distance. First flagellomere 1.65 underside of flagellum and all legs beyond times longer than its apical width; proportion 118 Rev. Soc. Entomol. Argent. 66 (3-4): 113-119, 2007

of lengths of scape, pedicel and proximal of flagellum, and all legs brownish; tibial three flagellomeres 2.4:0.5:1:0.4:0.4. spurs and translucent tegula yellowish brown. Membrane of wings amber; pterostigma and Male. Length: 12.0 mm; length of veins yellowish. forewing 10.0 mm. Color, pubescence and Pubescence. Hairs whitish on clypeus, sculpture similar to those of female, except labrum, around antennal sockets, and on as follows. Hairs on head and thorax longer, lower half of gena; rufous in front and behind those on scutum 1.0-1.4 times MOD, on ocelli on vertex. Hairs rufous on dorsum of mesopleuron 2.3-3.0 times MOD. Hairs on thorax, propodeum, and upper half of metasomal terga semierect, not appressed, pleurae, becoming paler towards venter of short, on T2 0.6-0.7 times MOD. Hairs of thorax; whitish on underside of coxae, apical fringes of S2-S4 as long medially as trochanters and femora; rest of legs with laterally, but those on S5 very short medially. brown hairs; outer surface of hind tibia S6 bare basally and laterally, apically with without distinct tuft of white hairs. Metasomal dense median tuft of erect hairs. T1 on basal vertical surface with erect Structure. Labrum rectangular, with yellowish hairs; apex of tergum with band of weakly trilobate apical projection. First pale yellow to yellowish brown, appressed, flagellomere nearly twice (1.8 times) as long short hairs. T2-T3 with broad pale yellow as its apical width; proportion of lengths of apical bands occupying most of exposed scape, pedicel and proximal three surface of terga; basad to the bands with short flagellomeres 2.4:0.6:1:0.55:0.6. T7 ending black hairs. T4 with narrower pale yellow in two short apical points broadly separated band broadly interrupted in middle; base and by U-shaped notch (Fig. 3). S7 and S8 as in center of T4 with black hairs and pale yellow Figs. 1-2. Gonostylus filiform. band with sparse, erect, black hairs. Hairs on T5-T6 black, on metasomal sterna brown. Etymology. The name refers to the Hairs on center of scutum 0.9-1.0 times homogeneous pale vestiture of the species. MOD, hairs on mesopleuron longer, 1.5-1.8 times MOD, hairs on postero-lateral angles Type material. ARGENTINA. Catamarca: of propodeum 2.0-2.3 times MOD. Holotype female, San Fernando, 4-6-XI-1989, Metapostnotum clothed with sparser hairs, L. Moffatt (MACN); 1 paratype male,same 1.0-1.5 times MOD. Tibial scopa with hairs data as holotype (MACN); 1 paratype female, on outer surface 1.0-1.4 times MOD, with Tinogasta, 8-III-1990, J.G. Rozen & A. Roig rachis slender and 18-22 branches directed A. (AMNH). La Rioja: 1 paratype female, X- apicad (Fig. 9). Metasomal terga densely 1925, M. Gómez (MACN); 1 paratype female, covered with appressed, short hairs, those on 18-II-1922, M. Gómez (MACN). apical band of T2 0.3-0.35 times MOD. Sculpture. Labrum with dense, regularly Ptilothrix nemoralis sp. nov. distributed punctures. Clypeus on lower third (Figs. 9, 10) with punctures 0.3-0.5 diameters apart, but on upper third 1-2 diameters apart; Diagnosis. This species is distinguished interspaces shiny. Punctation around and among all other Ptilothrix with yellow- above antennal sockets similar to that of banded metasoma by the pattern of the lower third of clypeus; rest of head more bands, covering almost all exposed surface finely punctured; vertex around ocelli on T2-T3 and interrupted on T4, and by the smooth. Punctures on scutum dense, 0.3-1.0 lack of the white tuft of hairs on the apex of diameters apart, with shiny interspaces; on the hind tibia. mesopleuron similar to those of scutum, but interspaces tessellate. Metapostnotum Description. Holotype female. Length: tessellate close to metanotum, remainder of 9.2 mm (paratype, 8.8 mm); length of metapostnotum shiny. Metasomal terga finely forewing: 7.5 mm (paratype, 7.3 mm). Color. punctured. Black, except middle of mandible, underside Structure. Inner orbits nearly straight, ROIG ALSINA, A. The albidohirta group of Ptilothrix 119 converging below, upper to lower interocular Científicas y Técnicas, Argentina, PIP 5755. distance 1.1:1. Ocellocular distance smaller than interocellar distance (0.75:1). Clypeus LITERATURE CITED 1.8 times wider than long, protuberant in lateral view for a distance 0.5 times maximum 1. BRÈTHES, J. 1910. Himenópteros argentinos. An. eye width. Labrum trapeziform, 1.68 times Mus. Nac. Buenos Aires 20: 205-316. 2. BROTHERS, D. J. 1976. Modifications of the as broad as length; apex truncate with antero- metapostnotum and the origin of the ‘propodeal apical margin carinate. Interantennal distance triangle’ in Hymenoptera Aculeata. Syst. 3.4 times antennocular distance. First Entomol. 1: 177 182. flagellomere 1.8 times longer than its apical 3. HAZELDINE, P. L. 1997. Comportamiento de nidificación de cuatro especies de Ptilothrix width; proportion of lengths of scape, pedicel Smith. Physis, Sec. C, Buenos Aires 54: 27-41. and proximal three flagellomeres 4. LINSLEY, E. G., J. W. MACSWAIN & R. F. SMITH. 2.56:0.56:1:0.4:0.4. 1956. Biological observations on Ptilothris sumichrasti (Cresson) and some related groups of emphorine bees. Bull. South. California Acad. Etymology. The specific name, which Sci. 55: 83-101. means «from the forest» in Latin, alludes to 5. MICHENER, C. D. 1944. Comparative external the habitat of the species, as opposed to the morphology, phylogeny, and a classification of habitat of the other two species of the group, the bees. Bull. Am. Mus. Nat. Hist. 82: 151-326. 6. MICHENER, C. D. 2000. The Bees of the World. The which occur in areas with more open and John Hopkins University Press, Baltimore and shrubby vegetation. London. xiv + 1-913 pp. 7. MICHENER, C. D., R. J. McGINLEY & B. N. Type material. ARGENTINA. Formosa: DANFORTH. 1994. The Genera of North and Central America. Smithsonian Institution Holotype female, Pozo de Maza, Press, Washington D.C. x + 209 pp. Departamento Bermejo, II-1989, P. Arenas 8. RUST, R. W. 1980. The biology of (MACN); 1 paratype female, Gran Guardia, (Hymenoptera: Anthophoridae). J. Kansas Entomol. II-1957, J. Foerster (MLP). Soc. 53: 427-436. 9. ROIG-ALSINA, A. 1994. Meliphilopsis, a new genus of emphorine bees, and notes on the relationships among the genera of Emphorina (Insecta: ACKNOWLEDGEMENTS Hymenoptera: Apoidea: Apidae). Reichenbachia 30: 181-188. 10. ROIG-ALSINA, A. 1999. Sinopsis genérica de la tribu I am indebted to the curators of the Emphorini, con la descripción de tres nuevos collections listed in the section Material and géneros (Hymenoptera, Apidae). Physis, Sec. C, methods for the loan of specimens, J. G. Buenos Aires 56: 17-25. Rozen, Jr. (AMNH), M. V. Colomo de Correa 11. SCHROTTKY, C. 1913. La distribución geográfica de los himenópteros argentinos. An. Soc. Cientif. (FIML), J. A. Schnack (MLP), and K. Schönitzer Argentina 75: 225 286. and J. Schuberth (ZSM); to C. D. Michener 12. SIPES, S. D. & V. J. TEPEDINO. 2005. -host and L. Ruz for helpful comments on the specificity and evolutionary patterns of host manuscript. This study was funded by the switching in a clade of specialist bees. Biol. J. Linn. Soc. 86: 487-505. Consejo Nacional de Investigaciones