,r.': ' ■ • COMMITTEE ON PUBLICATION George E. Lindsay, Chairman Edward L. Kessel, Editor OCCASIONAL PAPERS OF THE California Academy of Sciences No. 109, 36 pages ; 6 figures A REVISION OF THE GENUS ANCYLANDRENA (Hymenoptera:Andrenidae) By Thomag J. Zavortink j li/larina Blo!i;gic2i LaL. I LIBRARY APR 2 1974 [j Woods Hole, Masf ■***■"" ""''^'^^^***^ SAN FRANCISCO PUBLISHED BY THE ACADEMY March 14, 19 74 COMMITTEE ON PUBLICATION George E. Lindsay, Chairman Edward L. Kessel, Editor OCCASIONAL PAPERS OF THE California Academy of Sciences No. 109, 36 pages, 6 figures. A REVISION OF THE GENUS ANCYLANDRENA (HymenopteraiAndrenidae) By Thomas J. Zavortink Department of Biology, University of California, Los Angeles, California 90024 Abstract: Anaylandvena is a genus of rarely col- lected, vernal, andrenine bees found in the south- western United States and northwestern Mexico. It comprises four species, A. atoposoma (Cockerell, 1934), A. lavveae (Timberlake, 1951), A. koehelei (Timberlake, 1951), and A. timberlakei Zavortink, new species. Detailed descriptions of the genus and all four species are given; keys to the species are presented; the face, right mandible, pygidial plate, distal sterna, and genitalia of the male of each species are illustrated; the variation, biology, and distribution of the species are summarized; and the collection data for the 391 specimens examined are given. CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers Introduction Anaylandrena is a small genus of rarely collected bees found in the extreme southwestern United States and north- western Mexico. The group is poorly known: there is no complete description or diagnosis of the genus; there is no key to the species; there are no illustrations of the male genitalia or associated sterna of any species; and there is no summary of the biology or distribution of the genus or any of the included species. The primary purpose of the present study, based on the examination of 391 speci- mens (170 males, 221 females), is to provide as much of this information as possible. Cockerell (1930, pp. 5-6) originally described Anoyl- andvena as a subgenus of Andrena and later (Cockerell, 1941, p. 347) raised it to generic rank. Michener (1944, pp. 241- 243) treated Anoylandvena as a distinct genus and placed it with Andvena and Megandrena in the subfamily Andreninae of the Andrenidae. He implied a close relationship between Anoylandvena and Megandrena by stating that they were "gen- era of doubtful distinctness." Lanham (1949, p. 192) subse- quently reduced Anoylandvena to subgeneric rank under Megandvena. Although this interpretation has been followed by some (e.g. Timberlake, 1951, pp. 411-414; Rozen, 1971, p. 2), its acceptance has not been universal and Anoyl- andvena has been considered a separate genus by others (e.g. Stephen, Bohart, and Torchio, 1969, pp. 70, 125; Zavortink, 1972, pp. 66-68) . During the present study I examined numerous genera of short- tongued bees in an attempt to determine the affinities of Anoylandvena. Although I found many similarities, par- ticularly in the male genitalia and distal sterna, between Anoylandvena and genera presently placed in the subfamilies Melittinae and Macropidinae of the Melittidae and the sub- family Dufoureinae of the Halictidae, I do not believe I have discovered sufficient evidence to change the existing placement of Anoylandvena at this time, and I am leaving it in the subfamily Andreninae. I am, however, removing Megandvena from the Andreninae and placing it in the sub- family Panurginae of the Andrenidae, where it is related to the complex of genera including Calliopsis , Hypomaovoteva , and Nomadopsis . The evidence for this change will be presented separately. The morphological terminology used in the present study follows Michener (1944) . The male genitalia and distal sterna were studied in glycerol at a magnification of 200 diameters with a compound microscope. The plants collected by me have been determined by using A Calif ovnia Flova (Munz and Keck, 1959) and Supplement to a Calif ovnia Flova (Munz, 1968) . With one exception the names of plants recorded on the labels of specimens not collected by me are given un- altered in the lists of flower records and lists of speci- mens examined even though some of the species referred to are currently synonymized or otherwise known by different names. The one exception is in the genus Lavvea, where the No. 109] ZAVORTINK: ANCYLANDRENA names Larrea, Larrea divaricata , Lavrea glutinosa , and creosote bush have all been emended to Lavvea tridentata. Except as noted under Anoytandvena koehelei , I have not analyzed the pollen loads of females. The abbreviations used for the depositories of the specimens are indicated in the acknowledgments. Acknowledgments For loans of specimens or permission to study material in their care, I am indebted to: P. H. Arnaud, Jr., Califor- nia Academy of Sciences (CAS); G. C. Eickwort, Cornell Uni- versity (CU) ; P. D. Hurd, Jr. , National Museum of Natural History (USNM) ; W. E. LaBerge, Illinois Natural History Survey, who siobloaned material from the Academy of Natural Sciences, Philadelphia (ANSP) , Cornell University (CU) , Texas A & M University (TAM) , the University of Arizona (UA) , the University of California at Berkeley (UCB) , the Uni- versity of Kansas (UK) , and the Wild Bee Pollination Investigations, USDA, Logan, Utah (UTAH); U. N. Lanham (UNL) ; E. G. Linsley, University of California at Berkeley (UCB) ; C. D. Michener, University of Kansas (UK) ; J. G. Rozen, Jr. , American Museum of Natural History (AMNH) ; R. R. Snelling, Natural History Museum of Los Angeles County (LACM) ; Gerald I. Stage (GIS); R. W. Thorp, University of California at Davis (UCD) ; P. H. Timberlake, University of California at Riverside (UCR) ; P. F. Torchio, Wild Bee Pollination Investigations, USDA, Logan, Utah (UTAH); and F. G. Werner, University of Arizona (UA) . I thank Caryle Stallard for typing part of the first draft of the manuscript; L. Margaret Kowalczyk for pre- paring part of the preliminary drawings and all of the final illustrations; and my wife, Judy, for assisting with the collecting and labeling of specimens, the recording of data, and the preparation of some of the preliminary drawings, for typing most of the first and final drafts of the manuscript, and for handling all the correspondence relating to this study. I am grateful to John N. Belkin for many favors and assistance, Roy R. Snelling for innumerable courtesies and much freely given advice, Marjorie S. Favreau and Paul D. Kurd, Jr. , for searching for the holotype of Andrena hetevodoxa , and Henry J. Thompson for the outline maps that are used here in slightly altered form. Finally, I am indebted to John N. Belkin, Charles L. Hogue , and Roy R. Snelling for reading and criticizing the manuscript. Taxonomic Treatment ANCYLANDRENA COCKERELL Andrena {Anoylandrena) Cockerell, 1930, pp. 5-6; 1934, p. 82. Anoylandrena of Cockerell, 1941, p. 347; Michener, 1944, pp. 242-243; Linsley, 1951, p. 1086; Mitchell, 1960, CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers p. 86; Stephen, Bohart, and Torchio, 1969, pp. 70, 125; Zavortink, 1972, pp. 66-68. Megandrena (Ancylandrena) of Lanham, 1949, pp. 192, 193; Timberlake, 1951, pp. 411-414; Krombein, 1958, p. 216. TYPE SPECIES. Andrena (Anaylandrena) heterodoxa Cockerell, 1930; monobasic. BOTH SEXES. Medium sized species; length, exclusive of antenna, 8.0-14.7 mm. Head. Eye with scattered short hair. Ocelli normal or enlarged; in frontal view distance from lateral ocellus to summit of vertex less than or subequal to diameter of lateral ocellus. Ocellar triangle broad, distance between lateral ocelli subequal to or greater than distance from lateral ocellus to eye. Frontal line elevated below, usually im- pressed above, and extending dorsad to or nearly to median ocellus. Subantennal area more or less trapezoidal to some- what triangular, about as broad as length of inner subanten- nal suture. Clypeus moderately to strongly protuberant; disk flattened; with slightly to prominently elevated median longitudinal ridge or impunctate line; with moderately strong to very strong preapical, transverse, setaceous groove that is slightly to conspicuously curved or angled away from apex laterally. Malar space very short. Genal area subequal to or narrower than eye, without carina. Paramandibular carina without specialized setae. Pubescence abundant to very abundant, loose, especially dense and/or long on face and lower portion of gena; whitish to silver-white except some- times on upper part of face and vertex. Punctation fine and close to very fine and very close in center of face, becoming sparser in lower paraocular area, and sparser and slightly to distinctly coarser in upper portion of face; area dorsolaterad of lateral ocellus impunctate or nearly so. Integument between punctures smooth or roughened. Antenna. Scape, pedicel, flagellar segment 1, and dorsal surface of flagellar segments 2 to 10 or 11 dark brown to black, ventral surface of flagellar segments 2 to 10 or 11 tan or ferruginous to brown. Scape with long, entirely or predominantly white pubescence on inner, outer, and lower surfaces; pedicel with short, light hair. Mouthparts . Labrum subrectangular , width about 1.5-2.0 length; median basal portion slightly to strongly elevated and more or less emarginate around tuft of setae arising from de- pression; additional setae in lateral submedian group; black or slightly reddened. Mandible similar in both sexes; with or without basal blister; with subapical tooth on inner margin; moderately long, apices overlapping when closed. Proboscis without specialized bristles. Galea 0.6-0.8 length of prementum. Maxillary palpus extending beyond apex of galea; segments not markedly or constantly differ- entiated in length, segment 5 and sometimes segments 2 and 3 slightly shorter than others; segments 4-6 more slender than basal segments. Labial palpus shorter than maxillary palpus; segment 1 slightly elongate, segments 2 and 3 No. 109] ZAVORTINK: ANCYLANDRENA frequently shortened; segments 1-3 slightly flattened, segment 4 cylindrical and more slender than basal segments; apex of segments 2 and 3 produced beyond base of following segment laterally. Glossa acute; 0.3-0.4 length of prementum. Mesosoma. Pronotum with relatively strong suture ex- tending cephalodorsad from midportion of anterior margin of posterior pronotal lobe. Dorsolateral angle of pronotum scarcely produced, rounded. Metanotum slightly to moder- ately swollen. Metapseudosternal area longer than broad, rather narrow, the ventral hind coxal articulations closely approximated. Propodeal enclosure not bordered by carina; basal portion of enclosure with weak transverse striations, distal portion (on posterior surface of propodeum) smooth or minutely tesselate. Integument nonmetallic, entirely black or suffused with red or brown laterally. Pubescence very abundant, loose, usually somewhat shortened on meso- scutum. Punctation predominantly moderately fine and moder- ately close to very fine and close in anterior portion of mesoscutum and moderately fine to very fine but more widely spaced on disk of scutellum, becoming finer and closer in lateral portion of mesoscutum, in lateral and posterior portions of scutellum, and on metanotum. Integument smooth to slightly roughened on anterior portion of mesoscutum and disk of scutellum. Posterior center of mesoscutum more sparsely punctate and smoother than rest. Tegula elongate, cephalolateral margin usually concave; light brown or reddish brown to black; pubescent except caudolaterally . Legs. Hind coxa shortened. Tibiae not expanded. Malus of fore tibia and posterior spur of hind tibia not modified. Hind tarsus 0.9-1.0 length of mid tarsus. Basitarsi about 0.5-0.6 length of entire tarsus of corresponding leg. Distitarsi slightly enlarged. Claws normal, bifid. Arolia cleft dorsally, U- or V-shaped. Wings. Fore wing with pterostigma short, about 0.40-0.52 as long as distance from its apex to apex of marginal cell, narrow, about as broad as or slightly broader than distance from inner edge of prestigma to costal margin; apex of pterostigma considerably distad of base of vein r. Apex of marginal cell slightly bent away from costal margin, not or but minutely appen- diculate. Three submarginal cells; second longer than broad, trapezoidal; third slightly shorter than or subequal to first along posterior margin. First recurrent vein end- ing in distal 0.4 of posterior margin of second submarginal cell. Basal vein nearly interstitial with cu-v or arising distinctly distad of it. Pterostigma black to brown; wing veins black and brown, R and C darkest; wing membrane colorless or nearly so. Hind wing with jugal lobe broad, separated from vannal lobe by deep incision. Metasoma. Slightly to moderately flattened. Broad to near base in dorsal aspect. Anterior face of first tergum longer than dorsal face; concave. Integument nonmetallic, brown to black, sometimes suffused with red; apical margin of terga colorless to amber. Punctation of terga moderately CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers fine to very fine, moderately close to close. Integument of terga roughened basally, roughened or smooth dis tally. MALE. Head. Facial quadrangle longer than broad. Eyes subparallel to convergent ventrally. Facial fovea poorly defined, the entire upper paraocular area slightly to moderately depressed. Clypeus not or slightly pro- tuberant beyond level of supraclypeal area. Anterior mandibular articulation in front of lower anterior eye margin. Integument of head nonmetallic, black except for cream colored to yellow mark in lower paraocular area and sometimes on clypeus. Pubescence more abundant and longer than in female; clypeus moderately to very densely pu- bescent. Clypeus more sparsely and sometimes more coarsely punctate toward apex, impunctate and polished near apex. Antenna. Longer than in female. Flagellar segment 1 shorter than combined length of segments 2 and 3; segment 2 shorter than segments 1 or 3; segments 3-11 longer than broad. Mouthparts . Labrum with lateral submedian hair not arising from groove or depression. Mesosoma. Propodeum long to short, the basal portion moderately declivous to nearly vertical in profile. Pu- bescense whitish to silver-white except on dorsum where it is usually partially pigmented. Legs. Fore trochanter without spine. Basitibial plate moderate in size, oblong, its upper surface concave and with short appressed hair. Tarsi not unusually shortened; fore tarsus 1.4-1.5 length of fore tibia; mid tarsus 1.5-1.6 mid tibia; hind tarsus 1.1 hind tibia. Hind basitarsus slightly broadened. Legs dark brown or black basally, slightly lighter distally. Tibial spurs testaceous yellow. Pubescence abundant, loose, some hair elongate; predominantly white to silver-white. Wings. Fore wing sparsely to moderately pubescent basally. Metasoma. Less flattened than in female. Slightly to conspicuously curled under. Terga II-V moderately to conspicuously swollen cephalad of apical depression. Apical depressions punctured. Tergum VII with large, distinct, pointed pygidial plate with elevated edge and narrow, weakly to moderately developed median keel. Pubescense very abundant, loose; terga with hair predominantly long and erect, entirely silver-white to white or some pigmented; apical margin of terga II-V sometimes with weakly to moder- ately well developed white or silver-white fascia or fringe; sterna with hair predominantly erect, elongate on at least sternum I and lateral or apicolateral portion of sterna II-V, predominantly silver-white to white or some weakly pigmented. Distal sterna and genitalia. Sternum VI with broad, deep emargination in caudal margin; gradulus present, conspicuously bowed cephalad medially, not conspicuously bent caudad laterally; lobes at side of emargination with simple, thick setae with swollen bases and often blunt tips. Sternum VII with long, diverging anterior arms and with very strongly sclerotized, thick, distinctly Y-shaped median caudal process with lateral membranous hairy flap. Sternum VIII with large, more or less quadrangular internal portion No. 109] ZAVORTINK: ANCYLANDRENA with undulate surface, each caudal angle upturned and bearing a strong apodeme , the midbasal portion strongly deflexed; without spiculum; with moderately long, strongly sclerotized, median caudal process that has a flat-topped, glabrous basal plate with a strong longitudinal keel and a very strong longitudinal supporting apodeme. Gonobase large; dorsal surface very long; with deeply invaginated, thin, middorsal longitudinal apodeme; foramen of gonobase large, opening ventrally, its margin smooth, nearly oval or circular in outline in ventral aspect; apodeme surrounding foramen narrow. Gonocoxal apodeme very large ventrally, partly occluding foramen of gonobase, narrowing laterally, nearly absent dorsally; weakly sclerotized and without supporting ridges. Gonoforceps large, long, nearly straight in dorsal aspect; with single massive apical projection directed mesad and curved ventrad distally and with ventral or ventrolateral rounded, subapical angle; contiguous for 0.25-0.30 of length at base dorsally, then excavated around aedeagus ; deeply inflected at base dorsally, the resulting apodeme immovably attached to base of aedeagus; ventral surface membranous where it overlies volsella. Volsella small, approximately oval in outline; cuspus and digitus fused, neither elongate; nearly horizontal, the notch between cuspus and digitus oriented dorsoventrally ; nearly transverse, the digiti closely approximated; with relative- ly few, strongly sclerotized sensilla basiconica (not spicules); not fused with mate midventrally ; with firm yet membranous attachment to gonoforceps; dorsal surface of digitus firmly attached to and articulating with ventral lobe of aedeagus; without small, separate, dorsal sclero- tization. Penis and penis valves fused, the resulting aedeagus more or less completely sclerotized, slender, without dorsolateral flange or more basal lateral process and not excavated laterally; aedeagus more or less sickle- shaped in lateral aspect, directed dorsad basally, sharply bent caudad and then ventrad; distal portion of aedeagus more strongly sclerotized than proximal portion, deeply cleft, the resulting lobes contiguous or nearly so; penis not emergent or inflatable, the gonopore between bases of distal lobes; base of aedeagus, between apodemes of penis valves , abutting on and immovably attached to dorsal in- flection of gonoforceps; apodeme of penis valve extended caudad and produced into a large, strongly sclerotized ventral lobe that articulates with dorsal surface of digitus. FEMALE. Head. Facial quadrangle longer than broad to as long as broad. Eyes slightly divergent to slightly convergent ventrally. Facial fovea broad, extending mesad to or beyond level of outer edge of lateral ocellus, and deeply impressed above, becoming narrower and shallower below; short, extending ventrad only to level of upper margin of antennal socket; pubescence outstanding and greyish brown, chocolate brown, or dark brown in upper portion, becoming appressed and silvery in lower portion. Clypeus slightly to conspicuously protuberant beyond level of supraclypeal area. Anterior mandibular articulation below CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers or in front of lower anterior eye margin. Integument non- metallic, usually black, sometimes dark brown, sometimes slightly reddened ventrally. Pubescence less abundant and shorter than in male; clypeus sparsely pijbescent. Disk of clypeus nearly impunctate except for submedian longitudinal group of punctures; polished. Antenna. Shorter than in male. Flagellar segment 1 slightly shorter than to slightly longer than combined length of segments 2 and 3; segment 2 shorter than segments 1 or 3; segments 3-9 broader than long to longer than broad. Mouthparts . Labrum with all or part of lateral submedian hair arising from groove or depression. Mesosoma. Propodeum long to moderately long, the basal portion moderately to steeply declivous. Propodeal corbicula poorly developed; anterior margin without fringe; dorsal margin with dense vestiture of long, straight to curved, barbed or short-plumose hair; interior with short to long hair. Pubescence whitish to silver-white except on dorsum and sometimes upper portion of pleuron and propodeum, where it is partially or entirely pigmented. Legs. Basitibial plate large, broad, its upper surface more or less plane and with a central patch of short, dense tomentum. Tarsi shortened; fore tarsus 1.1-1.3 length of fore tibia; mid tarsus 1.3-1.4 mid tibia; hind tarsus 0.9-1.0 hind tibia. Mid basitarsus not expanded medially; slightly broadened and flattened; anterior and posterior margins subparallel. Hind basitarsus short, 0.5-0.6 hind tibia and 0.9-1.0 mid basitarsus; broad, width about 0.32-0.42 length; anterior and posterior margins subparallel; apex conspicuously produced beyond base of tarsal segment 2 posteriorly. Hind coxa and trochanter with hair dense to very dense, more or less straight, short to moderately long; longer hair, particularly on trochanter, predominantly simple, shorter hair predominantly barbed or short-plumose. Hind tibia with scopal hair dense, long, slender, flexible, simple, and gently curved toward apex of segment; longest hair much longer than apical width of tibia. Hind basi- tarsus with scopal hair similar to that of hind tibia; longest hair more than 0.5 length of basitarus. Legs reddish brown to dark brown or black basally, usually lighter distally. Tibial spurs testaceous yellow to deep amber, frequently darkened apically. General pubescence of legs abundant, predominantly loose. Mid femur with abundant short hair on posterior surface. Hind femur with abundant long hair on upper portion of anterior surface. Fore basitarsus with hair of outer surface long, simple, and gently curved toward apex of segment. Mid basitarsus with hair of outer surface dense, moderately long, plumose. Wings. Fore wing moderately pubescent basally. Metasoma. More flattened than in male. Not curled under. Terga II-IV slightly to moderately swollen cephalad of apical depression. Apical 0.36-0.44 of middorsal post- gradular area of terga II and III depressed, the depression narrowing and becoming more distinct laterally. Tergum VI with large pygidial plate with elevated edge and broad. No. 109] ZAVORTINK: ANCYLANDRENA flat- topped, strongly elevated median keel basally; pygidial plate longer than broad and with lateral margin usually concave dis tally. Pubescence abundant; tergum I with hair predominantly long, erect and loose, silver-white or weakly pigmented; terga II-IV with hair predominantly short and outstanding, pigmented; apical margin of terga I-IV with conspicuous, well developed, complete or medially inter- rupted white fascia; prepygidial fimbria pigmented, emarginate, the notch with short dense tomentum; pygidial fimbria pigmented; sterna I to III or IV with scopa; scopal hair simple and erect, strong and elongate on sterna I and II; apicolateral margin of sterna II-V with elongate, plumose hair. DISCUSSION. Anoylandrena can be distinguished from Andvena as follows: in both sexes by (1) the moderately to strongly protuberant clypeus , (2) the shortened hind coxa, (3) the slightly enlarged distitarsi, and (4) the long, concave anterior face of metasomal tergum I; in the male by (1) the large, distinct pygidial plate, (2) the long, diverging anterior arms and the strongly sclerotized, distinctly Y-shaped process of metasomal sternum VII, (3) the large, more or less quadrangular internal portion and the glabrous, flat- topped, keeled plate at the base of the process of sterniom VIII, (4) the characteristically shaped gonoforceps that are contiguous for 0.25-0.30 their length at the base dorsally, excavated around the aedeagus , and continued apically as undivided, massive projections, (5) the immovable attachment of the aedeagus to the dorsal inflection of the gonoforceps, and (6) the articulation of the volsella with a ventral lobe of the aedeagus; and in the female by (1) the short to moderately long, more or less straight hair on the hind coxa and trochanter, (2) the shortened hind tarsus and basitarsus, and (3) the scopa on the basal metasomal sterna. In addition, Ancylandvena can be distinguished from most species of Andvena by (1) the absence of specialized setae on the paramandibular carina, (2) the short and narrow pterostigma, and (3) the broad, median keel in the basal portion of the pygidial plate of the female. In spite of the small number of included forms, the genus Anoylandrena is rather varied structurally and the four species differ in such features as size, robustness, length and protuberance of the clypeus, length and declivity of the propodeum, concaveness of the anterior face of the first metasomal tergum, and length of the antennae. This structural diversity, in association with the diversity of pollen sources, suggests that the group is an old one. BIOLOGY. Relatively little is known about the biology of Anoylandrena. All species are vernal. Males are seldom found sleeping in flowers; those of at least A. koehelei , A. lavreae , and A. timberlakei fly extremely rapidly between and around shrubs frequented by females. The foraging behavior of females has not been observed, Anoylandrena koehelei , the one species with enlarged ocelli, is active primarily only in the very early morning and at least one 10 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers of the remaining species, A. timberlakei , is active through- out most of the day. Anaylandrena larreae is definitely oligolectic, on Larrea tridentata , while A. timbevlakei is apparently polylectic. The remaining two species may be oligolectic on legumes, Lupinus and Lotus in the case of A. atoposoma and Dalea in the case of A. koebelei . Mating of Anoylandvena has not been observed and nests have not been found. DISTRIBUTION. Ancylandrena is presently known from only the states of California, Nevada, and Arizona in the United States and Baja California and possibly Sonora in Mexico. One species, A. atoposoma, is largely restricted to coastal areas while the other three occur in the Mohave, Colorado, and Sonoran deserts. KEYS TO THE SPECIES OF ANCYLANDRENA Males 1. Integument of clypeus partially cream colored or yellowish; mandible with large, whitish or yellowish, nearly circular blister at base of outer surface; fore femur with very short hair 6n posterior and ventral surfaces 4. A. koebelei Integument of clypeus black; mandible without blister or with small, yellowish, tan, or brown, transverse blister at base of outer surface; fore femur with long hair on posterior and ventral surfaces 2 2. Mandible with small, yellowish, tan, or brown, transverse blister at base of outer surface; preapical, transverse groove of clypeus moderately strong; metasomal sternum II with pubescence long and conspicuous and sterna III and IV without conspicuous band of dense, shortened hair midapically 3. A. larreae Mandible without blister at base of outer surface; preapical, transverse groove of clypeus very strong; metasomal sterna not as above, either sternum II with pubescence short and inconspicuous or sterna III and IV with conspicuous band of dense, shortened hair midapically 3 3. Clypeus moderately long, strongly protuberant and densely pubescent; apex of labrum gently convex; metasomal sternum II with pubescence of middle portion moderately long and relatively conspicuous and sterna III and IV with a conspicuous band of dense, shortened hair midapically 2. A. timberlakei No. 10 9] ZAVORTINK: ANCYLANDRENA 11 Clypeus short, moderately protuberant and moderately pubescent; apex of labrum subtruncate; metasomal sternum II with pubescence of middle portion predominantly short and inconspicuous and sterna III and IV with poorly developed band of shortened hair midapically 1. a. atoposoma Females Mandible with large, whitish or yellowish, nearly circular blister at base of outer surface; scopal hair of hind tibia and basitarsus light to moderate amber in color; apical fascia of metasomal terga II-IV broadly interrupted medially 4. A. koebelei Mandible without blister or with small, yellowish, tan, or brown, transverse blister at base of outer surface; scopal hair on outer surface of hind tibia and basitarsus white, cream colored, greyish brown, brown, or black; apical fascia of metasomal terga II-IV complete or very narrowly interrupted or indented medially 2 Mandible with small, yellowish, tan, or brown, transverse blister at base of outer surface; preapical, transverse groove of clypeus moderately strong; metasomal sterna III and IV with definite scopa, the hair nearly, as long and strong as hair on sterna I and II 3. A. larreae Mandible without blister at base of outer surface; preapical, transverse groove of clypeus very strong; metasomal sternum III with weakly developed scopa, the hair much shorter and finer than hair on sterna I and II; sternum IV without definite scopa, the hair short and fine 3 Scopal hair on outer surface of hind tibia and basitarsus predominantly greyish brown, brown, or black; clypeus moder- ately long, strongly protuberant; apex of labrum gently convex 2. A. timbevlakei Scopal hair on outer surface of hind tibia and basitarsus predominantly white to cream colored; clypeus short, moderately protuberant; apex of labrum subtruncate. . . 1. A. atoposoma 12 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers 1. Ancytandvena atoposoma (Cockerell) . (Figures 1 , 3. ) Andrena (Anoylandrena) heterodoxa Cockerell/ 1930, pp. 6-7. Junior primary homonym of Andrena heterodoxa Perez, 1903. Andrena atoposoma Cockerell, 1934, p. 82. Nomen novum for A. heterodoxa Cockerell, 1930. Anoylandvena atoposoma of Cockerell, 1941, p. 347; Michener, 1944, p. 242; Linsley, 1951, p. 1086. Megandrena (Ancytandvena) atoposoma of Lanham, 1949, p. 193; Timberlake, 1951, pp. 411, 412, 413. TYPE. Holotype male. Riverside, Riverside County, California, 25 April 1930, P. H. Timberlake, on Cryptantha intermedia (USNM 55307). BOTH SEXES. Head. Greatest width 1.24-1.35 length from top of vertex to apex of clypeus. Ocelli normal. Clypeus moderately protuberant; short; preapical transverse groove very strong, slightly curved away from apex laterally. Punctation fine and close in center of face, becoming dis- tinctly coarser dorsad. Integument smooth. Mouthparts . Apex of labrum subtruncate. Mandible without basal blister. Mesosoma. Pubescence of mesoscutum relatively sparse. Punctation predominantly moderately fine and moderately close in anterior portion of mesoscutum. Integument smooth on anterior portion of mesoscutum and disk of scutellum. Metasoma. Integument of disk of terga II-IV predomi- nantly smooth. MALE. Length, exclusive of antenna, 8.7-11.1 mm. Length of fore wing 6.5-7.3 mm. Width of head 3.1-3.4 mm. Head (fig. 3) . Width of face at level of clypeal base 0.90-0.9 3 length from lower edge of median ocellus to apex of clypeus. Eyes convergent ventrally. Pale mark of lower paraocular area usually yellowish to yellow; usually scarcely broadened below and nearly crescentic; clypeus black. Pubescence of clypeus usually moderately dense, long. Hair on upper part of face and vertex whitish to brown, dark brown, or black. Antenna. Flagellum short, segment 1 slightly shorter or slightly longer than segment 3, segment 2 broader than long, segments 3-10 1.16-1.32 times as long as broad. Mouthparts . Mandible black with reddened apex. Mesosoma. Propodeum long, the basal portion moderately declivous. Dorsal mesosomal hair rarely entirely silver- white, usually some pigmented, the darkest, varying from brown to black, on posterior half of mesoscutum and scutellum; sometimes additional light brown hair on pos- terior lateral edge of mesoscutum, axilla, and edges of scutellum. Legs. Fore femur with long hair on posterior and lower surfaces. Wings. Fore wing moderately pubescent basally . Metasoma. Concavity of first tergum moderately con- spicuous. Pygidial plate (fig. 3) more or less deltoid; short, length 1.0-1.3 basal width; lateral margin straight to slightly concave; median keel moderately developed. No. 109] ZAVORTINK: ANCYLANDRENA 13 Pubescense of terga entirely white to silver-white, or much or most dorsal, long, erect hair tan or brown; apical or apicolateral margin of terga II-V with poorly developed fascia of shortened, semierect or decumbent, white to silver-white hair; hair of middle portion of sternum II much shorter than hair of sternum I and scarcely longer or stronger than hair of sternum III; sterna III and IV without a complete apical fringe, with a poorly developed band of shortened hair midapically; pubescence of proximal sterna predominantly silver-white. Punctation of terga moderately fine, moderately close. Distal sterna and genitalia (fig. 3) . Sternum VI with gradulus not reaching antecosta medially. Sternum VII with emargination between arms of caudal process parabolic in shape; basal portion of emargination sparsely pubescent; membranous flap of process narrow, with moderately dense pubescence, the proximal hair short and simple to long and sparsely plumose, the distal hair long, densely plumose. Sternum VIII with caudal process moderately broad, usually slightly expanded at apex which is subtruncate to broadly rounded; dorsolateral edge of process without flange; pubescence of side and apex of process moderately dense. Gono forceps slightly curved ventrad dis tally; apex pointed in ventral aspect; subapical angle directed ventrolaterally. Volsella with digitus and cuspus separated by deep notch with sensilla basiconica on its walls. Aedeagus strongly bent, the distal portion directed ventroposteriorly ; distal lobe of aedeagus not expanded. FEMALE. Length, exclusive of antenna, 10.1-11.8 mm. Length of fore wing 6.8-7.2 mm. Width of head 3.3-3.5 mm. Head. Width of face at level of clypeal base 0.93-1.01 length from lower edge of median ocellus to apex of clypeus. Eyes slightly convergent ventrally. Preapical transverse groove of clypeus usually densely setaceous laterad to near level of lateral edge of labrum. Integument uniformly black or slightly reddened ventrally. Some hair on upper part of face and vertex tan to dark brown or black. Antenna. Flagellum short, width of segments 3-9 subequal to or greater than length. Mouthparts . Mandible black with reddened apex. Mesosoma. Propodeum long, the basal portion moderately declivous. Brown to black hair on posterior portion of mesoscutum, scutellum, and sometimes center of metanotum; fulvous, tannish, or greyish hair on anterior portion and edge of mesoscutum, edge of scutellum and metanotum, and sometimes on axilla, posterior lobe of pronotum, upper portion of pleuron, and upper lateral portion of propodeum. Legs. Pubescence of coxae, trochanters, and femora predomi- nantly white to silver-white; pubescence of outer surface of fore and mid tibiae light chocolate brown to brown; elongate hair of fore basitarsus brown; pubescence of outer surface of mid basitarsus white and/or light brown; scopal hair on outer surface of hind tibia and basitarsus predomi- nantly white to cream colored. 14 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers Metasoma. Concavity of first tergum moderately con- spicuous. Long hair of tergum I silver-white; short outstanding hair of terga II-IV predominantly dark brown to blackish, that at base of segments whitish, forming a narrow, pale band; apical fascia of terga II-IV complete or very narrowly interrupted or indented medially on segments II and III; hair of apical fasciae usually not closely appressed to integument; prepygidial and pygidial fimbriae greyish brown, chocolate brown, or dark brown medially, becoming lighter or white laterally; scopa poorly developed, present on sterna I-III; scopal hair of sterna I and II sparse; scopal hair of sternum III conspicuously shorter and finer than that of sterna I and II; sternum IV with hair short and fine, similar to hair of sternum V; pubescence of proximal sterna predominantly white to silver-white; hair of sternum VI chocolate brown to brown. Punctation of terga moderately fine to fine, moderately close to close. DISCUSSION. Ancylandvena atoposoma is separated from the most similar species, A. timberlakei , as follows: in both sexes by (1) the shorter and less protuberant clypeus , (2) the subtruncate apex of the labrum, and (3) the smoother integument of the metasomal terga; in the male by (1) the sparser, shorter pubescence of the clypeus, (2) the shorter and less conspicuous pubescence of metasomal sternum II and the poorly developed band of dense, shortened hair midapi- cally on sterna III and IV, (3) the usually yellowish to yellow, more or less crescentic, integumentary mark in the lower paraocular area, (4) the stouter flagellum of the antenna, and (5) the broader caudal process on metasomal sternum VIII; and, in the female by the predominantly white to cream colored scopal hair on the outer surface of the hind tibia and basitarsus. This species is considerably variable. The males from Ventura County, California, are smaller than average and have predominantly or entirely whitish pubescence on the head and mesoscutum; the males from Deep Creek, on the north side of the San Bernardino Mountains, San Bernardino County, California, also have the pubescence of the head and mesoscutum lightened and have the pale integumentary mark of the lower paraocular area of the face slightly broadened below; some males from the extreme western part of the Colorado Desert northeast of Palm Springs, Riverside County, and at Morongo Valley, San Bernardino County, California, have the face marks larger, more triangular, and/or more cream colored than usual, some have dense pubescence on the clypeus, and some have the metasomal integument roughened; and, finally, the males from east of Socorro in the Sierra San Pedro Martir of Baja California, Mexico, have the face marks whitish. The single female from Monterey County, California, has the pubescence of the upper part of the head and mesoscutum predominantly tannish or fulvous, the hair of the metasomal fasciae closely appressed to the integument, the brown hair of the No. 109] ZAVORTINK: ANCYLANDRENA 15 prepygidial fimbria reduced in extent, and the punctures of the metasomal terga finer and closer than usual. Anoylandrena atoposoma and A. timberlakei are closely related, basically allopatric species. At the one locality where they have been taken together, Deep Creek, on the north side of the San Bernardino Mountains, the only possi- ble evidence of interbreeding between them is the slightly broadened face marks in the males of A. atoposoma. The variations in the males of A. atoposoma from the extreme western part of the Colorado Desert northeast of Palm Springs and at Morongo Valley listed in the previous para- graph are in the direction of A. timberlakei and probably indicate that both species occur in this region and occasionally hybridize. Of the four species of Anoylandrena , A. atoposoma is the most generalized anatomically, the least dimorphic sexually, and the only one largely restricted to a mesic habitat. For these reasons I believe it is the most primitive species of the group. BIOLOGY. The recorded seasonal flight period of A. atoposoma extends from 5 March to 21 June. No information is available on the daily flight period. Females exhibit a preference for flowers of the legumi- nous genera Lupinus and Lotus. A complete list of flower records for 44 males and 47 females is: Lupinus hallii , 4 d, 24 V ; Lupinus formosus , 5 9; Lupinus paynei , 4 9; Lupinus species, 2 9; Lotus saoparius, 5 6, 39; Lotus glaber, 2 9; Lotus species, 1 c5 , 19; Calochortus splendens , 8 6; Calochortus species, 6 6; Ceanothus species, 1 <^ , 3 9; Convolvulus species, 1 cJ ; Cryptantha intermedia , 2 6, 1 9; Eriodiotyon triohocalyx , 12 i ; Eriogonum fas oiculatum , 1 <5 ; Gilia virgata, 1 9; Penstemon species, 1 cJ ; Phaoelia ramosissima , 1 9; Sphaeraloea ambigua , 2 d . DISTRIBUTION (fig. 1). Anoylandrena atoposoma occurs in cismontane southern California, cismontane northern Baja California, and the adjacent desert regions. There is also one distant, uncertain record from Sonora, Mexico. SPECIMENS EXAMINED. 66 males, 54 females, as follows: MEXICO BAJA CALIFORNIA. Ensenada, 15 April 1941, T. D. A. Cockerell, on Convolvulus , 1 d (USNM) . Socorro (6 km. E. ) , Sierra San Pedro Martir, 27 May 1958, J. Powell, 2 6 (UCB) . SONORA. San Bernardo, 25 March 19 35, G. E. and R. M. Bohart, 1 6 (UTAH) . UNITED STATES CALIFORNIA. Inyo County: Mountain Springs Canyon, Argus Mts., 22 May 1937, C. D. Michener, 1 9 (UK). Los Angeles County'. Claremont, 5 March 192 8, M. McFadden, 1 9 (LACM) . Monterey County'. Stone Canyon, 27 April 1919, E. P. Van Duzee, 1 9 (CAS). Riverside County'. Banning, 15 May 1926, W. D. Pierce, on Calochortus , 6 6 (LACM). The Gavilan, 15 May 19 30, C. M. Dammers and P. H. Timberlake, on Calochortus splendens, 6 6 (UCR) ; 18 May 1939, P. H. Timberlake, on Calochortus splendens, 1