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Noqvitatesamerican MUSEUM PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y NoqvitatesAMERICAN MUSEUM PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3066, 28 pp., 45 figures June 11, 1993 Nesting Biologies and Immature Stages of the Rophitine Bees (Halictidae) with Notes on the Cleptoparasite Biastes (Anthophoridae) (Hymenoptera: Apoidea) JEROME G. ROZEN, JR.' CONTENTS Abstract .......................... 2 Introduction .......................... 2 Nesting Biology of the Rophitinae .......................... 3 Sphecodosoma dicksoni .......................... 3 Conanthalictus conanthi .......................... 11 Rophites trispinosus .......................... 15 Profile of the Biology of the Rophitinae .......................... 16 Mature Larvae of the Rophitinae .......................... 17 Key to the Mature Larvae .......................... 18 Sphecodosoma dicksoni .......................... 18 Conanthalictus conanthi .......................... 21 Rophites trispinosus .......................... 23 Pupa of Sphecodosoma dicksoni .......................... 24 Discussion .......................... 24 References .......................... 26 lCurator, Department of Entomology, American Museum of Natural History. Copyright C American Museum of Natural History 1993 ISSN 0003-0082 / Price $2.90 2 AMERICAN MUSEUM NOVITATES NO. 3066 ABSTRACT Information on the nesting biology ofthe ground- The mature larvae of the Rophitinae are char- nesting Sphecodosoma dicksoni (Timberlake) and acterized on the basis of six genera, and a key to Conanthalictus conanthi (Cockerell) from the available species is presented. The mature larvae southwestern United States is added to previously of Sphecodosoma dicksoni and Conanthalictus published data to provide an understanding ofnest conanthi are described taxonomically and com- architecture, cell construction, provisioning, egg pared with larvae ofother Rophitinae. Also treat- deposition, larval feeding behavior, cocoon con- ed is an immature larva (probably last instar) of struction (in the case of S. dicksoni), larval defe- Rophites trispinosus. Whereas the mature larvae cation, and voltinism of these species. No clep- of S. dicksoni and R. trispinosus share many fea- toparasitic bees are associated with either species tures with rophitine genera Dufourea and Xeral- at present. ictus, that of C. conanthi is very different, though Observations on nest provisioning and larval clearly sharing significant synapomorphies with the adaptations ofthe related Palearctic Rophites tris- others. Many ofits differences appear to be related pinosus Perez are included. Recovery of an inter- to the fact that it does not spin a cocoon. The pupa mediate-stage larva of Biastes emarginatus ofS. dicksoni is also described, the first such treat- (Schenck) (Nomadinae: Biastini) from the nest es- ment for any member of the subfamily. tablishes this host association of the cleptoparas- New information on nesting biology and im- ite. Its larva and the mature larva of the related matures of the rophitines, though supporting the genus Neopasites are compared and are found to monophyly of the subfamily, does not seem to share many derived features. Based on informa- demonstrate phylogenetic linkages with the Hal- tion presented here and on published and unpub- ictinae and Nomiinae or with the Andrenidae and lished accounts, a synopsis of the biology of the Melittidae at this time. However, a number of Rophitinae is presented based on 7 genera and 14 characters are identified and discussed that may species. The synopsis identifies features that seem eventually be helpful in resolving these relation- to be characteristic of the subfamily. ships. INTRODUCTION This paper is an attempt to increase our permits a formal (but tentative) overview of understanding ofthe nesting biology and im- the biology ofthe subfamily for the first time. mature stages ofthe Rophitinae (= Dufourei- Larvae ofonly a few taxa ofrophitines have nae), one of the three generally recognized been described and the account of the pupa halictid subfamilies. Such information will of Sphecodosoma dicksoni below is the first eventually contribute to an understanding of treatment of any pupa of the subfamily. To the relationships ofthe rophitines to the other date mature larvae ofonly Dufourea (Torchio subfamilies of Halictidae and to other taxa. et al., 1967 [incomplete]; McGinley, 1981, The research started as a reinvestigation of 1987; and Eickwort et al., 1986) and Rho- the nesting behavior and ecology of Sphe- phitoides (Enslin, 1921) ofthe approximately codosoma dicksoni (Timberlake) and Con- 12 or 13 recognized genera have been treated. anthalictus conanthi (Cockerell). Nests ofboth In addition, Batra and Michener (1966) treat- species had been briefly studied earlier (Roz- ed the immature larva of Systropha. en and McGinley, 1976), but so much new The manuscript for the present paper was information resulted from the current re- initiated as a study ofthe nesting biology and search that an expanded presentation oftheir immatures of S. dicksoni; data on the other biology seems appropriate. In addition, lar- two taxa were subsequently added and com- vae and pollen-nectar masses ofRophites tris- pared with information on S. dicksoni. This, pinosus Perez preserved in the American Mu- and not presumed relationships, accounts for seum of Natural History (AMNH) allowed the sequence ofpresentation ofthe taxa here. interpretation of some aspects of the biology of this species in light of what is known of the other two species. The new information ACKNOWLEDGMENTS on these species, when combined with ac- I thank both George C. Eickwort and Ron- counts of the biology of other Rophitinae, ald J. McGinley for carefully reviewing this 1993 ROZEN: ROPHITINE BEES 3 manuscript and offering a number ofthought- for Sphecodosoma dicksoni. Precipitation in provoking suggestions. the area comes primarily in winter, resulting Roy R. Snelling kindly provided a copy of in a spring blooming period. During dry years a manuscript prepared by him and G. I. Stage the area is without bloom, but the precipi- on the nest and larva of Xeralictus timber- tation of the 1991-1992 season was copious, lakei. He also graciously donated the single and the flowering abundant. Although the known larval specimen of this species to the sand-dwelling Oenothera was past bloom and AMNH, thus permitting me to examine it Larrea tridentata had nearly finished flow- first hand. George C. Eickwort loaned the ering at the April 30-May 1 visit, Nama ap- larva of Dufourea novaeangliae (Robertson) peared to be at maximum flower. The plant and thereby added to the completeness ofthe was still blooming on May 11 although there study. I extend my appreciation to Richard were fewer flowers and a reduced number of H. Kruzansky who analyzed the soil from the bees visiting the plant. In addition to Sphe- nesting area. Noel Holmgren, ofthe New York codosoma, the following bees visited the plant: Botanical Garden, identified the Nama as- Calliopsis (Micronomadopsis) foleyi (Tim- sociated with Conanthalictus conanthi. berlake), Perdita (Pseudomacrotera) turgiceps Andrey Sharkov and Beatrice Brewster as- Timberlake, P. (Heteroperdita) arenaria sisted by translating some of the foreign lit- Timberlake,2 and P. (Perdita)flavicauda for- erature. mosa Timberlake. Sphecodosoma dicksoni nested in a sparse- ly vegetated area at the base of the east side NESTING BIOLOGY OF THE of the sand dune. The nest entrances were ROPHITINAE interspersed with the low-growing (ca. 15 cm high) Nama plants. Although the patches of Sphecodosoma dicksoni plants were scattered along the east side of The biology of this species was studied by the dunes as well as elsewhere in the region, Rozen and McGinley (1976) (Sphecodosoma nests were found only in two areas. The one then considered a subgenus of Conanthalic- site studied (fig. 1) consisted of a loose ag- tus) at 21 mi south of Animas, Hidalgo Co., gregation of nests, approximately 10 in an New Mexico, in August 1975. Information area of 1.5 x 0.5 m with other nests scattered in the earlier report agrees in most respects farther away. The other aggregation, 30 m with the new data, but differences that do away in an ecologically similar area, sug- occur are discussed within the description. gested that, iftime had permitted, nests might Measurements, counts, and other informa- well have been found at the bases of other tion in brackets are data from the 1976 re- sand dunes where there were sufficient num- port, presented here for comparison. bers of plants to support the foraging needs DESCRIPTION OF SITE: The nesting site of of the females. Sphecodosoma dicksoni was studied at 18 The plants and nests were in low elevations miles west of Blythe, Riverside County, Cal- adjacent to the sand dunes, where the ground ifornia, on April 30 and May 1, 1992, and sloped about 150 [20-30° in the 1976 study was again examined on May 11, 1992. This in New Mexico]. This low slope gave the im- locality has been visited by entomologists pression that the species prefers horizontal from the early 1950s to the present and was nesting surfaces, but observations on the nests referred to as Hopkins Well on some early presented below indicate otherwise. Loose on specimen labels. The dominant feature ofthe the surface, the soil was firmer below, es- site is the loose sand dunes that, though pecially where cells were encountered, but somewhat lower, have remained virtually could easily be excavated with a penknife. stationary
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