AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3057, 12 pp., 25 figures March 24, 1993

South American Panurgine Bees (Apoidea: : ), Part I. Biology, Mature Larva, and Description of a New Genus and Species

LUISA RUZ1 AND JEROME G. ROZEN2

ABSTRACT

Parasarus Ruz, new genus, is described and il- atacamensis includes: pollen sources, nest archi- lustrated. This genus ofsmall, black, ground nest- tecture, provisioning, oviposition habits, egg de- ing panurgines is found in Chile and Catamarca scription, larval defecation habits, and cuckoo bee Province, Argentina. It is based on P. atacamensis parasitism. The mature larva is described taxo- Ruz, new species (described herein), from north- nomically, illustrated, and compared with larvae ern Chile, and contains a number ofother species, of , , and Pterosarus. yet to be named. Biological information about P.

RESUMEN

Se describe e ilustra un nuevo genero de Pan- nadas. Informacion biol6gica de P. atacamensis urginae, Parasarus Ruz. Las especies de este ge- incluye: fuentes de polen, arquitectura de nido, nero son pequeiias, negras, anidan en el suelo y se aprovisionamiento, habitos de oviposicion, des- encuentran en Chile y en la Provincia de Cata- cripcion del huevo, hibitos de defecacion de la marca, Argentina. Parasarus, basado en P. ata- larva y abejas parasitas. La larva madura es des- camensis Ruz, n. sp. (especie que aqui se describe), crita taxonomicamente, ilustrada y comparada con se encuentra en el Norte de Chile, y contiene ade- larvas de Protandrena, Pseudopanurgus y Ptero- mas varias otras especies que aun deben ser nomi- sarus.

' Professor, Laboratorio Zoologia, Universidad Cat6lica de Valparaiso, Valparaiso, Chile. 2Curator, Department of Entomology, American Museum of Natural History.

Copyright K American Museum of Natural History 1993 ISSN 0003-0082 / Price $1.40 2 AMERICAN MUSEUM NOVITATES NO. 3057

INTRODUCTION This is the first of a series of papers in structures were inked by Carmen Tobar, which we shall describe and name new genera Universidad Catolica of Valparaiso, Chile and subgenera of South American panurgine (UCV). Beatrice Brewester, American Mu- bees and provide information about their seum of Natural History (AMNH), helped nesting behavior, ontogeny, floral prefer- prepare the manuscript, and Eric Quinter ences, and other aspects of their natural his- (AMNH) assisted in editing it. tory. Where possible we will also describe We thank Professor Haroldo Toro and Dr. their immature stages, especially the last-stage Terry L. Griswold for carefully reviewing the larva. Because the Panurginae tend to be manuscript and making a number ofvaluable small-bodied, they were generally overlooked suggestions for its improvement. by early workers who concentrated on col- lecting and describing larger South American SYSTEMATICS bees. As a consequence much ofthe very rich panurgine fauna of that continent has yet to In the following descriptions ofadults, pas- be studied. This paper deals with a panurgine sages in italics indicate the most important genus that at this time contains perhaps three diagnostic features. T and S refer to num- unnamed species, from Chile and Catamarca bered metasomal terga and sterna (e.g., T5, Province, Argentina. The commonest species S4). Fore, mid, and hind legs are numbered and the one for which we have the mature 1, 2, and 3, respectively. Measurements were larva and biological information is described made with an ocular micrometer in a Wild herein and serves as the type of the genus; M5 stereomicroscope with a 25 x objective. the other species await description. Ratios are expressed in units of the ocular Because there are many new genera and micrometer: 25 units = 1 mm. Figures of subgenera of panurgines to be described, the male genitalia and associated sterna are ori- first efforts ofthe series will be to characterize ented so that their bases point toward the these taxa and validate their names as infor- bottom of the page and their apices toward mation becomes available; therefore, the se- the top. Collection data have been tran- quence of presenting taxa in the series does scribed to correspond to the recent reorga- not necessarily imply relationships. We hope nization ofthe political regions and provinces to analyze their phylogenetic relationships in of Chile; names in capital letters refer to a final paper of the series. Revisions of the regions, and specific localities are in italics. genera identified in the present series will be undertaken separately. To do so now would PARASARUS Ruz, new genus be a prohibitively long task. Figures 1-18 In this paper, the first author (LR) provides TYPE SPECIES: Parasarus atacamensis Ruz, the descriptions ofthe genus and species based new species. on adults, and the second author (JGR) con- DiAGNOSIS: Parasarus superficially resem- tributes the sections on Biology and Mature bles Heterosarus since both genera have a Larva. rather flattened face, slightly depressed lateral Specimens of immature stages and nest foveae on metasomal tergum 2 (T2), and components are in the collection of the rather short, thin, and apparently simple hairs American Museum of Natural History. De- on the tibial scopa. Nevertheless they can be position of adult specimens is treated below. easily separated because Parasarus shows no ACKNOWLEDGMENTS yellow on the face of the male, has sparse mesosomal punctation, and its metasoma is Information about the ecology, behavior, usually as wide as the mesosoma. Males of and immatures ofParasarus atacamensis was Parasarus with a wide head look like Ce- collected on field trips supported by grants to phalurgus males, but in Cephalurgus the su- JGR from the National Geographic Society praclypeal area is distinctly swollen, the para- (No. 3844-88) and from the National Science ocular areas are clearly convex, the lateral Foundation (GB5407). Drawings of adult foveae ofT2 are well defined, and the genital 1993 RUZ AND ROZEN: PANURGINE BEES, PART I 3 capsule has a small gonobase. Parasarus dif- than 2nd abscissa M + Cu. Hind wing with fers from Heterosarus and Cephalurgus as cu-v about 1/4 or 1/5 as long as second abscissa follows: subantennal area about as wide as M + Cu (fig. 2). Basitarsus 1 somewhat more length of inner subantennal suture, small tu- than 4 times longer than broad in male and bercle present between antennal sockets, male more than twice as wide in female. Femur 2 genitalia and associated sterna as illustrated, of female basally with comb on ventral bor- and female tibial spurs distinctly curved api- der not well defined. Middle tibial spur of cally. female almost as long as basitarsus 2, distally DESCRIPTION: Body small (3-5 mm), black withfine, evenly spaced teeth (fig. 15); in male or dark brown, with some yellow markings more than half as long as basitarsus 2 and especially on legs. Pubescence in general short with minute, fine, dense teeth. Basitarsus 2 and whitish, minute on most parts of meta- about as long as 1 and shorter than 3. Tibia somal terga, rather sparse although usually 3 of female less than twice as long as basi- denser on lower halfofface. Integument are- tarsus 3, with keirotrichia on most of inner olate but smooth in some undescribed species surface except close to dorsal and ventral from Chile and Argentina. Punctation in gen- margins; male with dorsal margin of tibia 3 eralfine, sparse, usually dense on clypeus. serrate (fig. 12), with keirotrichia as in female. Head varying from narrower to wider than Scopal hairs on tibiafine, apparently simple, mesosoma. Mandible simple. Glossa shorter rather sparse, somewhat longer than hairs at than prementum (much shorter in an unde- apex ofmetasoma, and bent toward their api- scribed Chilean species). Segment I oflabial ces. Hind tibial spurs with fine teeth; in male, palpus shorter than 2-4 together. Maxillary outer one somewhat shorter than inner one palpus six-segmented. Galeal comb well de- and more curved toward apex; infemale spurs veloped (6-12 bristles) (fig. 6). Labrum (fig. subequal and strongly curved at apex. Basi- 3) with basal part smooth, impunctate, usu- tibial plate ofmale well defined, with margins ally delimited by ridge or carina; distal part somewhat elevated, carinate. Tarsus 3 un- usually flat, strongly reflexed (slightly convex modified. Claws bifurcate, rather deeply cleft; in an undescribed species from Argentina). rami subequal in male, inner shorter than Tentorial pit in intersection between episto- outer in female. mal suture and outer subantennal suture (fig. T2 with lateralfovea extremely weak (ab- 4). Antennal socket (fig. 4) just above middle sent in one undescribed Chilean species). Py- offace (below in an undescribed Chilean spe- gidial plate rounded at apex in female, absent cies). Lower mesal paraocular area rather in male except for small median glabrous area flattened. Facialfovea elongate, shallow, and distally on T7. T8 of male trapezoidal. S5 about half as long as scape (or shorter) in and S6 with hairs short, appressed, and di- male, deeperand much longer infemale. Mid- rected obliquely toward midline. S6 of male dle ocellus above orbital tangent (frontal view). distally with rather large membranous area Vertex convex. Gena of variable width. and V-shaped median emargination. S6 of Pronotum, on dorsal margin, with rounded female with basal margin straight, margin ridge laterally. Preepisternal groove well de- bordering apodeme strongly sclerotized, and veloped, extending below scrobe (weak in lateral margin with strong curved ridge; distal some undescribed species). Pterostigma lon- margin slightly concave in middle; duplica- ger than prestigma, with middle area yellow- tion (on dorsal side ofsternum) membranous ish; side basal to vein r clearly divergingfrom (see Michener, 1944; Ruz, 1991); exposed costa; side within marginal cell curved (fig. 1). distal area of S6 basally with minute sparse Marginal cell (fig. 1) truncate at apex, slightly hairs, which are longer and denser on pre- longer than distance from its apex to wing marginal area (especially dense laterally). S7 tip. Two submarginal cells. First submarginal ofmale with two distal projections; proximal cell longer than second (cell 2 = 2 + 3 ac- arms widely open. S8 of male with gradually cording to Ruz, 1991). Second cell M (max- tapered distal projection; body of sternum imum length) shorter than first one. First re- with strong longitudinal salient on dorsal sur- current vein not interstitial with first face (figs. 10, 11). Genital capsule with wide, transverse cubital. Forewing with cu-v longer inflexed gonocoxal apodeme; gonocoxites 4 AMERICAN MUSEUM NOVITATES NO. 3057

1

2 3

7

Figs. 1-10. Parasarus atacamensis, n. sp., male. 1, 2. Forewing and hindwing. 3. Labrum. 4. Head, frontal view. 5. S6, dorsal view left; ventral view right. 6. Mouthparts. 7. Head, lateral view. 8. Genital capsule, dorsal view left; ventral view right. 9. S7. 10. S8, dorsal view left; ventral view right.

usually elongate, fused proximally only; gon- surpassing stylus), first valvula well sclero- ostyli fused to gonocoxite, not articulated. tized, valve present. Volsellae apparently free mesally, with den- The species ofthis genus are distributed in ticles; penis valvae simple, acute distally, northern Chile and in Catamarca Province, fused to each other and to membranous pe- Argentina. nis, except at apex. Sting elongate (slightly ETYMOLOGY: The prefix is from Greek 1993 RUZ AND ROZEN: PANURGINE BEES, PART I 5

11

14 15

1

Figs. 11-18. Parasarus atacamensis, n. sp. Figs. 11, 12, male. 11. S8, lateral view. 12. Hind tibia. Figs. 13-18, female. 13. S6, dorsal view left; ventral view right. 14. T6. 15. Middle tibial spur. 16. Hind tibial spurs. 17. Sting. 18. T7 (= hemitergite). meaning "akin to," and the stem has been laterally, with hairs slanting toward very well- used in the names ofa number ofpresumably delimited, glabrous midline area; integument related genera. generally areolate (dull on dorsal part offace); labrum with premarginal ridge which is lat- Parasarus atacamensis Ruz, new species erally delimited by distinct clypeal projec- Figures 1-18 tion; genitalia and associated sterna as illus- DIAGNOSIS: Males can be recognized by the trated. Females have a clypeus with a median following: relatively dense pilosity on the longitudinal groove. In both sexes the lateral lower half offace; metasomal sternum 2 (S2) parts of the clypeus are somewhat concave. apparently almost bare; S3-S5 clearly pilose DESCRIPTION: MALE. Length about 4 mm; 6 AMERICAN MUSEUM NOVITATES NO. 3057 head width 1 mm; mesosoma width 1 mm; width narrower than eye in lateral view (8: forewing length 3 mm. COLORATION. 11). Pterostigma twice as wide as prestigma Black, with following parts yellow: ventral (4:2). Marginal cell 1.1 times longer than dis- part of flagellum (dorsal part brown), base of tance from its apex to wing tip (22:20) (fig. mandible, apex offemur, outer surface offor- 1). Basitibial plate narrowing slightly toward etibia, apex of second and third tibia, basi- apex. Tibia 2 with spur shorter than basitar- tarsus (other tarsomeres testaceous). Wings sus, finely serrate (fig. 12); tibia 3 with inner hyaline; tegula, veins, and stigma testaceous. spur a little longer than outer one. Fovea of PUBESCENCE. In general short, sparse, T2 weakly depressed. Pygidial plate absent. whitish and with tiny, almost invisible S6 weakly membranous distally, with small branches, denser on lower part offace (below V-shaped emargination (fig. 5). S7 with distal antennal socket), sparse on scutum, scutel- processes triangular in shape, shorter than lum, and postscutellum; hairs simple on dis- basal ones (fig. 9). Proximal half of S8 with tal fourth of labrum. Metasomal terga with longitudinal median carina ventrally, and very short, simple hairs; some sparse strongly developed laminar process dorsally branched hairs on premarginal and lateral ar- (figs. 10, 11). Genital capsule as illustrated eas; hairs almost absent on T1; branched hairs (fig. 8). becoming denser toward apex of metasoma. FEMALE. Length about 5 mm; head width Metasomal sterna with fine hairs, but S3 and 2 mm; mesosomal width 2 mm; forewing S6 each with group oflateral stiffhairs point- length 3.5 mm. As described for male except: ed toward glabrous mesal area, these hairs COLORATION. Black, with following parts very short and very fine on S6. PUNCTA- yellow: anterior surface of foretibia, apex of TION. Head clearly areolate above antennal middle tibia, and posterior basitarsus. Meta- socket, with fine, somewhat shallow punc- somal sterna brown. PUBESCENCE. Clypeal tures irregularly distributed; lower part of pubescence sparse. Labrum pubescent on head and gena almost smooth, shining, im- apical half. Metasomal sterna with proximal punctate. Mesosomal integument like that of part glabrous mesally; S5 with denser hairs upper part of face, but punctures larger and distally; S6 densely pubescent laterally, mesal sparser. Propodeal triangle finely striate ba- area apparently glabrous, but group of short, sally; posterior part smooth, shining, im- dense, yellow hairs apically on each side. punctate; lateral parts sparsely punctate and STRUCTURES. Head width to length 46:37. slightly areolate. Metasoma smooth; terga Labrum width to length 15:9. Clypeus with finely and rather densely punctured, though median longitudinal sulcus; clypeal width to punctures irregularly scattered, usually coars- length 28:15. Facial fovea somewhat broader er on posterior terga than on anterior terga. than that of male; its length to inner suban- Metasomal sterna weakly areolate, with fine tennal suture 11:0.5. Supraclypeal area punctures which tend to be deeper laterally prominent. Distance from inner orbits to an- toward apex. STRUCTURES. Head 1.2 times tennal socket compared to distance between broader than long (42:34) (head width versus sockets 0.8:0.9. Genal width to eye width 1 1: length varying from 1.2 to 1.5 times) (fig. 4). 10. Marginal cell length to distance from its Labrum a little more than twice as broad as apex to wing tip 23:18. Basitibial plate some- long (14:6), with ridge delimiting marginal what expanded distally. Middle tibial spur area (fig. 3). Clypeus little more than twice about as long as basitarsus, becoming more as broad as long (23:1 1) (varying from 2.08 denticulate apically (fig. 15). Posterior tibial to 2.6 times broader than long). Facial fovea spurs subequal in length, their apices clearly narrow, slightly longer than inner subanten- curved (fig. 16). Lateral fovea of T2 larger nal suture (0.5:0.4). Supraclypeal area scarce- than that of male, sometimes oblique, ex- ly produced between antennal sockets. Fron- panded distally. S5 with wide V-shaped tal line with a small protuberance at apex. emargination apically. S6 as in figure 13. Py- Distance from inner orbit to antennal socket gidial plate with longitudinal swollen ridge; a little shorter than distance between sockets apex narrowed and rounded (fig. 14). Sting (5:8). Eye length varying from 0.75 to 0.65 and T7 as illustrated (figs. 17, 18). of head length (fig. 7). Gena with maximum Holotype male and allotype: Paipote, At- 1 993 RUZ AND ROZEN: PANURGINE BEES, PART I 7 acama Region, Chile, October 11, 1971 (J. Los Choros: 49, X-12-1977 (J. C. Maguna- G. Rozen and L. Pefna); in the collection of celaya); (Choros, Canyon, 10 km S Incahuasi) the American Museum of Natural History. 16, 19, X-13-1969 (J. G. Rozen, L. Pe-na). Paratypes: CHILE.-ATACAMA: Paipote: Carretera Panamericana al Norte de La Se-

896, 229, X-15-1969, 38, 19, X-20-1971 , 46, rena: 26, 1970-72, Int. Biol. Program (A. 19, X-1-1 971, 56, 19, X- 12-1971 (all J. G. Moldenke). La Serena: 29, X-29- 1966 (R. W. Rozen, L. Pefna); 106, 29, X-10-1977 (V. Ca- H.). Totoral: 16, X-21-1980 (L. Pefia). Los bezas); same date: 16 (H. Flores), 168, 19 (J. Loritos (N of Choros Bajos): 76, X- 1983 (L. C. Magunacelaya), 56 (L. Ruz), 106 (E. De La Pefia). Pangue: (22 km S of Vicufna) 66, 79, Hoz), 106 (H. Toro) and 48 (E. Balart). Al- X-16-1971 (J. G. Rozen, L. Peiia). garrobal (Paipote): 56, X- 11- 1977; same date: VALPARAISO: Aconcagua Province: 36, 29 (E. Balart), 16 (H. Flores), 19 (H. Toro). Punta Molles: 16, X-14-1977 (R.W.H.). Canto del Agua: 146, 29, X-9-1977 (H. FPo- Most of the paratypes are deposited in the res); same date: 246 (J. C. Magunacelaya), AMNH and UCV. Other paratypes will be 366, 89 (H. Toro), 86, 19 (E. Balart), 266, 69 deposited in the following collections: Museo (V. Cabezas); 1506, 109, X-21-1969, 29, X- 1 3- Nacional de Historia Natural, Santiago, Chile; 1971 (J. G. Rozen, L. Pefna). Caldera: (14 km Snow Entomological Museum, University of N) 466,249, X-9- 1971 (J. G. Rozen, L. Penia). Kansas, Lawrence, Kansas; Bee Biology and Copiapo: (42 km S) 146, 49, X-19-1969 (J. Systematics Laboratory, Utah State Univer- G. Rozen, L. Peiia); (44 km S) 46, 89, X-2 1- sity, Logan; National Museum of Natural 1971, (55 km S) 39, same date, (66 km S) 16, History, Washington, D.C.; the Museum of X-19-1971,(10-20 miS), 16, X-18-1969 (all Natural History, London, England; Museum J. G. Rozen, L. Pei'na); (NW) 26, 1100 m, X-4- fir Naturkunde der Humboldt Universitiit zu 1980 (L. Penia); (100 km S) 48, X-1-1980 (L. Berlin, Germany; Museo Argentino de Cien-

Penia). Between Copiapo and Juntas: 76, 1 9, cias Naturales "Bernardino Rivadavia," 1600 m, X-2,3-1980 (L. Pe-na). Between Pai- Buenos Aires, Argentina. pote and Las Juntas: 126, 29, X- 15-1969 (J. CoMMENTs: This species is highly variable, G. Rozen, L. Pefna). LasJuntas: 136,39, X- 15- especially with respect to head width, clypeal 1969 (J. G. Rozen, L. Pefia); 26, 19 (on Rio width, length and position of eyes (from di- Huasco), 1250 m X-27-1967 (L. Pefia); 66, vergent above to more or less subparallel), IX-1968 (E. De La Hoz); same date: 56 (H. integumental texture, and puncture density Toro), 46 (E. Montenegro), 28 (L. Ruz). Tra- on head and scutum. Because of this varia- vesia: 786, 59, X- 1969, (H. Toro); same date: tion, species boundaries are uncertain and 46, 1 , (G. Dazarola), 36 (E. Montenegro), 46, therefore some specimens (in AMNH) col- 29 (L. Ruz); 16, 19, X-11-1977 (H. Toro); lected near Vicunia, Elqui Province, Coquim- same date: 26 (J. C. Magunacelaya), 26 (L. bo Region, were not designated as paratypes. Ruz). Vallenar (10 km S): 66, X-13-1971 (J. They might not be conspecific with those from G. Rozen, L. Pefia); Conai (E. Vallenar): 126, the Atacama Region; however, there are no X-25,26-1980 (L. Pefia). Q. Maitencillo (SW significant differences in the genitalia be- of Vallenar): 16, X-l 1- 1980 (L. Penia). Pinte tween them. Also, 676 and 249 from Pinte (SE of Vallenar): 396, 169, 1600 m, X-25- (SE Vallenar), Atacama Province, Chile (L. 1980 (L. Pefia). Vicinity of Freirina: 46, 29, Pejna) (in AMNH), were not designated para- X-14-1969 (J. G. Rozen, L. Peina). Obispito: types because they lacked collection dates. 19, X-17-1969 (J. G. Rozen, L. Penfa). Individuals ofboth sexes ofP. atacamensis COQUIMBO: Llano de la Higuera: (N of have been found visiting the following plant Tofo) 366, 329, X-14,15-1971 (J. G. Rozen, species: Baccharis paniculata, Gutierrezia L. Penia); Choros Bajos: 166, 139, X- 12-1977 taltalensis, Mesembrianthemum crystallin- (H. Flores); same date: 96, 19 (J. C. Magun- um, and Calandrinia sp. Further host plant acelaya), 76 (H. Toro), 26 (V. Cabezas), 36, data are given in the section on Biology, be- 79 (E. Balart), 38 (E. De La Hoz) and 16, 19 low. (M. Rojas). Los Choros: 76, 229, II-2-1972 DISTRIBUTION: This species is found in (L. Ruz); same date: 26, 89 (E. Montenegro), northern Chile. 19 (M. Pasten) and 19 (H. Flores). Quebrada ETYMOLOGY: The specific name refers to 8 AMERICAN MUSEUM NOVITATES NO. 3057 the Atacama Desert in Chile, where many with small yellow flowers. This plant, which specimens have been collected. was not identified and which grew for about 10 m next to the shoulder of the road, was BIOLOGY the food source at this locality. Soil of very fine, dry silt containing numerous large rocks DEScRIPrION OF SITES: Nests of Parasarus was faintly moist at the cell level. The nest atacamensis were encountered at three sites. area was unshaded except for the low-grow- Two nests, about 100 m apart, were discov- ing pollen plant. Only one female seemed to ered at 6 km south ofVicu-na, Elqui Province, be associated with this nest, as was also sug- Chile, in October 1991. This area is pictured gested by the single open cell. and characterized in Rozen (in prep. 1) and The third nesting site, consisting ofa single further discussed in Rozen (in prep. 2). As nest among nests ofLiphanthus alicahue Ruz indicated in the Systematics section, speci- and Toro at Paipote, Copiapo Province, was mens from this region differ from type spec- mentioned in Ehrenfeld and Rozen (1977) imens of this species, suggesting that Vicu-na and Rozen (1989). When it was excavated on material may eventually be found to be non- October 20, 1972, the only significant infor- conspecific. Most information about nesting mation noted was the apparent association biology comes from this site. ofKelita tuberculata as a nest parasite ofPar- Both nests were composite, being used by asarus. a number of females. One, excavated on Oc- NEST ARCHITECTURE: The two nests stud- tober 19, 1991, was in rather sandy soil (faint- ied (from near Vicu-na and near Copiapo) were ly moist at the cell level) on a nearly hori- similar. Both had simple entrances, without zontal shoulder of an unpaved road. Four tumuli, about 2.0 mm diameter. In each, the females as well as a single female of Kelita main tunnel meandered downward. The main tuberculata Ehrenfeld and Rozen were col- tunnel ofthe nest from Vicuina remained open lected from it before it was dissected. JGR its entire length and descended along a crack unsuccessfully attempted to excavate the oth- at first so that its diameter did not become er nest (on a mostly unshaded surface, sloping constant until it reached a depth of 8-9 cm. about 200) in November after it had become Cells were arranged in all directions from the dormant. Notes below refer only to the nest burrow and were connected to it by filled studied in October. At this locality, females laterals, some ofwhich ranged in length from bearing pollen were collected from Alonafil- 4.5 to 6 cm. These cells were found at depths ifolia and Pleurophora pungens.3 Because the from 13 to 16 cm, and some may have been color of the pollen carried by females from even deeper. the two plants was plant specific, there is little The main tunnel from the Copiapo nest doubt that both plant species are used for remained at a diameter of 2 mm and was larval food. plugged with loose fill 4 to 8 cm down. At 8 A single nest ofParasarus atacamensis was cm the tunnel divided, with one branch end- also examined at 28 mi south of Copiapo, ing several cm away and the other one me- Copiapo Province, Chile (a site within the andering mostly horizontally for 11 cm. The known range of typical P. atacamensis), latter branch opened into a cell that was being alongside a paved highway on October 20, provisioned. Three complete cells, all at about apparent exit holes occur- 1969. Numerous 10 cm in depth, were closely associated with ring near the nest entrance as well as an abun- the open one and were connected to the open dance ofboth males and females on the flow- burrow by filled laterals. Hence this nest was ers suggested that other nests were present. somewhat shallower and tended to -extend The nest entrance, on horizontal ground, was farther laterally than the other. at the base of a low-growing succulent plant Arranged singly, cells of both nests were nearly horizontal, tilting at most 10 to 200 toward the rear. The entrance diameter of a 3Plants kindly identified by C. Marticorena and M. cell from each nest was 2.0 mm. Cell dimen- Quezada, Departamento de Botanica, Universidad de sions at the Vicunia site were 4.7-5.2 mm long Concepcion, Chile. and 2.7-2.8 mm in maximum diameter (N 1993 RUZ AND ROZEN: PANURGINE BEES, PART I 9

= 3); at the Copiapo site, 5.0-5.5 mm long ences, as is also suggested by host labels on and 2.2-2.7 mm in maximum diameter (N adult specimens (above). = 3). One cell from Vicufia may have had a Eggs, found only in the Copiapo nest, were slightly flatter floor than ceiling, but cells from placed on top of the flattened food spheres Copiapo appeared symmetrical around their parallel to the long axis of the cell. Shiny, long axes. Cells from both sites were normally white, with a smooth chorion, the egg was round at the rear and, in other ways, unre- slightly arched and touched the food mass at markable in shape. On the Copiapo samples, both ends. Two eggs measured 1.35 mm long surfaces of cell walls were smooth and only and 0.7 mm in maximum diameter. The pos- faintly shiny. Droplets ofwater placed at var- terior end, toward the rear of the cell, was ious points on the wall indicated that the rear somewhat more pointed than the rounded and floor were waterproof, but the ceiling near anterior end. The single larva observed at the cell entrance readily absorbed droplets. Copiapo rested on top of the provisions and Beneath the waterproof, transparent lining, ate the food beneath its head; it was unable the cell wall may have been composed of a to crawl. Small larvae from Vicunia were found very thin layer ofspecially worked soil in the in the same position. One cell from there, Copiapo material, but this layer was not no- with feces plastered on the rear of the cell as ticed in the Vicunia material. At least at Vi- a single concave mass, contained an actively cuna, cell walls were slightly harder than the moving postdefecating larva. Another cell substrate. All cell closures were concave spi- with feces in the same position was empty, rals on the inside with three to four coils. A suggesting either that the occupant ofthe cell closure from Vicufia (but probably charac- had been of the same generation as its larval teristic of all closures) clearly revealed a con- nest mates but had already matured, or that cave, rather smooth outer surface, with a clo- the nest was being used by more than one sure thickness of 2 mm at the periphery and generation. Pupal structures were developing approximately 0.5 mm at the center, as seen in at least one of the postdefecating larvae in cross section. Loose soil had filled the lat- and in several of the predefecating larvae, eral leading to the closure. indicating that the larvae were nearing pu- PROVISIONING AND DEVELOPMENT: Pollen pation. Hence the species probably has more was transported in a distinctly moist condi- than one generation per year. tion on the anterior surfaces of the female's ADULT ACTIVITY: Males flew actively hind tibiae. Completed food masses at both around the flowers at both sites and were sites were flattened spheres. Those at Copia- presumably searching for mates. po were 1.6-1.7 mm high (N = 3) and 2.1- PARASITISM: Rozen (in prep. 2) definitely 2.3 mm (N = 2) in maximum diameter. The associated Kelita tuberculata with Parasarus mass was mealy-moist throughout and placed atacamensis at the Vicu-na site and tenta- about 0.5 mm from the rear ofthe cell. It was tively at Paipote. In addition, several partly not "glued" to the cell floor by secretions or to fully grown meloid larvae were found in nectar, and it was not coated either by a wa- the nest at Vicufna. terproof covering or a layer of nectar. Pro- visions in an open cell at Copiapo indicate that food loads from foraging trips are stored MATURE LARVA first as paired irregular masses at the rear of Figures 19-25 the cell rather than as a single shaped small DiAGNOSIS: References to descriptions of mass as is the case with some of the more mature larvae of other panurgines can be specialized panurgines. Because females were found in McGinley (1988). gathering pollen from two species of plants The mature larvae ofthis species resemble in the Vicunia area, this species does not ap- larvae ofProtandrena, Pseudopanurgus, and pear to be narrowly oligolectic. Another fe- Pterosarus in that they possess well-devel- male bearing moist pollen was collected on oped maxillae and large maxillary palpi and Calandrinia at Puquios, Copiapo Province, a strongly recessed labial region. They also on October 24, 1991, further suggesting that exhibit somewhat forward projecting dorsal the species may have broad pollen prefer- thoracic tubercles as described below and, 10 AMERICAN MUSEUM NOVITATES NO. 3057

2 4 t7 VENTRAL LA tEAL PHUNOTAL SWELLING APICAL EDGE 23 ~~~~24 2

Figs. 19-25. Parasarus atacamensis, n. sp., mature larva. 19. Predefecating larva, lateral view. 20. Postdefecating larva, lateral view. 21. Head, lateral view. 22. Head, frontal view. 23-25. Right mandible, dorsal, adoral, and ventral views, respectively. Scale (= 1.0 mm) refers to figures 19 and 20. with the exception of Protandrena, small dorsal apical edge.4 The greatly enlarged, glo- spiracles, the atria of which do not project bose antennal prominences in conjunction above the body wall. However, this species with the curved vertex as seen in lateral view can be distinguished from the few species in these other genera whose larvae are known 4 The mandible may actually possess a subapical tooth the lower on the dorsal apical edge homologous with the subapical in that its mandible is serrate along tooth of Protandrena, Pseudopanurgus, and Pterosarus. apical edge (also true for Protandrena), and However, because the teeth on the mandible are large the mandible does not appear to bear a sub- (relative to mandible size), many of the teeth along the apical tooth larger than other teeth along the dorsal apical edge appear equally large. 1993 RUZ AND ROZEN: PANURGINE BEES, PART I I1I

(fig. 21) is unique to Parasarus, so far as is characteristic of Panurgus. Labrum normal known now. While larvae ofthese other gen- in size and shape for Panurginae, without era have somewhat projecting antennal sclerite; paired labral tubercles arising from prominences, they are not as globose as those labral disc, large; epipharynx spiculate lat- of Parasarus and their vertices tend to be erally. angled or projecting as seen from the side. Mandible (figs. 23-25) moderately slender; The larva ofthis species should be compared dorsal mandibular spiculation very fine, with larvae of Heterosarus, Xenopanurgus, sharp-pointed, nonsetiform; outer surface Pseudosarus, Rhophitulus, Chaeturginus, Ce- without tubercle; apex tapering, slender, sim- phalurgus, Metapsaenythia, and Anthemur- ple (i.e., with subapical tooth subequal in size gus, which have yet to be described. This to other teeth on dorsal apical edge; ventral assemblage, including Parasarus, is a largely apical edge with distinct serrations; cusp unresolved clade in the Panurginae according moderately strongly produced, with moder- to Ruz (1986). ately large, evenly spaced teeth, not produced Characters that are particularly helpful in ventrally as in Panurgus and without large distinguishing mature larvae of Parasarus cuspal tooth as in Melitturga. Labiomaxillary atacamensis from those of other panurgines region recessed, and, except for maxillary are presented in italics. apices, fused. Maxilla as seen in lateral view HEAD (figs. 21, 22): Integument of head (fig. 21) projecting well beyond labial apex; capsule with very few sensilla, which are not cardo and stipes represented as unpigmented, setiform; sensilla of mouthparts also not se- vague integumental thickening (best seen on tiform; integument of both pre- and post- cleared specimen); articulating arm ofstipital defecating larvae unpigmented except for sclerite perhaps evident on sides of hypo- mandibular apices. pharyngeal groove; palpus large, at least as Head size moderate in relationship to rest large as labral tubercle, not directed down- ofbody ofpostdefecating larva; head capsule wardas in Panurginus; dorsal surface ofmax- much wider than maximum length from ver- illa spiculate, with some spicules extending tex to lower clypeal margin. Tentorium in- onto dorsal surface of maxilla. Labium not cluding dorsal arms well developed; anterior divided into prementum and postmentum; pits in normal position on face; posterior ten- premental sclerite not evident; labral apex torial pit normal in position; posterior thick- recessed by comparison with maxillary api- ening of head capsule well developed, only ces, unlike those in Camptopoeum; labial pal- slightly curving forward medially as seen in pus only vaguely represented as slight swell- dorsal view; posterior margin ofhead capsule ing, much smaller than maxillary palpus. normal in position; median longitudinal Salivary opening in weakly expressed, curved thickening of head capsule absent; hyposto- groove. Hypopharynx slightly bulging but well mal ridge well developed, without ramus; recessed under labrum, spiculate. pleurostomal ridge well developed; epistomal BODY: Integument without obvious setae; ridge well developed between anterior man- dorsally integument not spiculate; ventrally dibular articulations and anterior tentorial most segments with small, widely spaced, fine pits, absent mesad of these pits; epistomal spicules, except posteroventral area on ab- depression moderately pronounced between dominal segment X with band of large spic- pits. Parietal bands distinct though hidden in ules; lateral pronotal swelling strongly spic- frontal view (fig. 22) by enlarged antennal ulate. Body form (figs. 19, 20) moderately prominences. Antennal prominences very robust; intersegmental lines only moderately strongly developed, globose; antennal disc and incised on pre- and postdefecating larvae; in- papilla moderate in size, with three sensilla. trasegmental lines not evident; paired dorsal Vertex in lateral view (fig. 21) evenly rounded, body tubercles (figs. 19, 20) moderately small; without projections or elevations and, by con- dorsal prothoracic tubercles apically some- trast, accentuating large antennal promi- whatforwardprojecting; meso- and metatho- nences; frontoclypeal area normal in length racic dorsal tubercles subapically somewhat and configuration, not projecting strongly, as forward projecting; hence thoracic tubercles 12 AMERICAN MUSEUM NOVITATES NO. 3057 similar to those of Protandrena, Pterosarus, south of Vicuina, Elqui Province, Chile, Oc- Pseudopanurgus, Melitturga, but unlike those tober 19, 1991 (J. G. Rozen). of Meliturgula, Camptopoeum, Panurgus, most Calliopsis (sensu Ruz, 1991), and Pan- REFERENCES urginus; dorsal abdominal tubercles rounded Ehrenfeld, J., and J. G. Rozen, Jr. (i.e., not transverse), nonspiculate, but arising 1977. The cuckoo bee genus Kelita, its system- abruptly from each segment, hence well de- atics, biology, and larvae. Am. Mus. fined; prothorax (figs. 19-22) with pair oflat- Novitates 2631: 24 pp. eral swellings that are strongly spiculate,5 sug- McGinley, R. J. gestive ofprothoracic flaps of Camptopoeum 1988. A catalog and review ofimmature Apo- (Rozen, 1988: figs. 5-7) except not so flat; idea (). Smithson. Con- pleural region elsewhere not produced; only trib. Zool. 494: 24 pp. abdominal segment X without paired tuber- Michener, C. D. segment X with dorsal somewhat 1944. Comparative external morphology, cles; surface phylogeny, and a classification of the elevated as seen in lateral view (figs. 19, 20); bees (Hymenoptera). Bull. Am. Mus. segment X attached centrally to IX, not pro- Nat. Hist. 82: 151-326. duced posteriorly or ventrally. Spiracles ex- Rozen, J. G., Jr. tremely small, unpigmented, virtually impos- 1966. Systematics of the larvae of North sible to detect on uncleared specimen under American panurgine bees (Hymenop- normal magnification; anterior spiracles tera, Apoidea). Am. Mus. Novitates somewhat larger than posterior ones; peri- 2259: 22 pp. treme not certainly present; atrium not pro- 1988. Ecology, behavior, and mature larva of jecting above body wall, globose, without rim, a new species of the Old World bee ge- apparently unornamented; primary tracheal nus Camptopoeum (Andrenidae: Pan- opening with collar; subatrium of moderate urginae). Am. Mus. Novitates 2925: 12 PP. length, consisting ofapproximately 10 cham- 1989. Life history studies of the "primitive" bers; in general spiracle, except for non- panurgine bees (Hymenoptera: Andren- grooved atrial wall, similar to that of Ptero- idae: Panurginae). Am. Mus. Novitates sarus boylei (Cockerell) (Rozen, 1966: fig. 4). 2962: 27 pp. Male sex characters consisting of pale, trans- In prep. 1. Phylogenetic relationships of Eu- verse, dumbbell-shaped imaginal disc in herbstia with other short-tongued bees middle of venter of abdominal segment IX; (Hymenoptera: Apoidea). Am. Mus. female sex characters, paired paramedian, Novitates. pale imaginal discs on venter of abdominal In prep. 2. Biology and immature stages of the South American cuckoo bee genera Bra- segments VII, VIII, and IX; those of segment chynomada and Kelita (Hymenoptera: VII farthest apart, those ofIX closest togeth- Anthophoridae: Nomadinae). Am. Mus. er. Novitates. MATERIAL STUDIED: 4 predefecating, Ruz, L. 2 postdefecating, nondiapausing larvae, 6 km 1986. Classification and Phylogenetic Rela- tionships of the Panurgine Bees (Hy- menoptera-Andrenidae). Ph.D. thesis, sCursory examination of larvae of a number of pan- Univ. Kansas, Lawrence, KS, 312 pp. urgine taxa suggests that spiculate lateral prothoracic lobes 1991. Classification and phylogenetic rela- immediately behind the head may have been overlooked tionships of the panurgine bees: The in previous treatments ofpanurgine larvae. This feature Calliopsini and allies (Hymenoptera: seems variably developed in different taxa and should Andrenidae). Univ. Kansas Sci. Bull. 54: be addressed in future descriptions. 209-256.