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Novttates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET NEW YORK, N.Y AMERICAN MUSEUM Novttates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET NEW YORK, N.Y. 10024 U.S.A. NUMBER 2551 OCTOBER 10, 1974 JEROME G. ROZEN, JR., AND RONALD J. MCGINLEY Systematics of Ammobatine Bees Based on Their Mature Larvae and Pupae (Hymenoptera, Anthophoridae, Nomadinae) ., 1, 'm -; ;f, I1, Systematics of Ammobatine Bees Based on Their Mature Larvae and Pupae (Hymenoptera, Anthophoridae, Nomadinae) JEROME G. ROZEN, JR. Deputy Director for Research and Curator of Hymenoptera The American Museum of Natural History RONALD J. MCGINLEY Research Assistant, the American Museum of Natural History Present Address: Department of Entomology and Parasitology University of California, Berkeley AMERICAN MUSEUM NOVITATES NUMBER 2551, pp. 1-16, figs. 1-33 ISSN 0003-0082 Issued October 10, 1974 Price. $1.00 Copyright © The A -erican Museum of Natural History 1974 ABSTRACT SYSTEMATICS AND PHYLOGENY Mature larvae of representatives of four genera The Ammobatini were divided by Popov of the Ammobatini are taxonomically de- (1951) into the Ammobatini (Ammobates, Par- scribed-Pseudodichroa, A mmobates, Morgania, ammobatodes, and Oreopasites) and the Pasitini and Oreopasites. A key is provided for their iden- (including among others, Morgania and presum- tification and the tribe is characterized on the ably Pseudodichroa). In reviewing Pseudodi- basis of the mature larvae. Pupae of representa- tives of two genera are described-Oreopasites chroa, Rozen (1968) used the older, more con- and Morgania. A preliminary key to the pupae of servative classification employed by Michener the subfamily Nomadinae is also presented. (1944) and others because there was some evi- dence that a more detailed study of male geni- The study was undertaken to illuminate the talia might necessitate a revision of Popov's higher classification and phylogeny of antho- classification. Recently Baker (1971) has fol- phorid bees, to provide means for identifying the lowed Popov's division and correctly pointed out tribe Ammobatini and its genera on the basis of that in the Pasitini the labrum of the adult ex- immature characters, and to establish a back- tends only to the mandible, whereas in the Am- ground for a forthcoming evolutionary study of mobatini, sensu stricto, it extends well beyond the ammobatine genus Oreopasites. the mandibles. However, we note that within the The present paper may be considered a "third Pasitini there is a difference among the genera generation" investigation on bee larvae. Miche- with respect to whether the mandibles of adults ner's work on larvae (1953) was the original com- are directed mesiad so that their points overlap in *prehensive treatment of larval forms but did not repose (e.g., Morgania) or whether the mandibles describe any of the Ammobatini. Since its publi- are directed somewhat posteriorly so that their cation various workers, with new and more ex- apexes cross in repose (e.g., Pseudodichroa; see tensive material, readdressed various taxa, and Rozen, 1968, fig. 2). Furthermore, an analysis of specifically Rozen produced a series of papers larval features does not suggest a grouping of treating the larvae of the Anthophoridae. Rozen Morgania and Pseudodichroa on the one hand in 1966 covered the Nomadinae and discussed and Oreopasites and Ammobates on the other. the only genus of ammobatines (Oreopasites) Indeed a phenetic grouping would seem to place then available. The present paper is yet another Oreopasites and Morgania close together, with refinement, made possible by recent acquisitions Ammobates somewhat divergent from them and of larval specimens of three more genera of the with Pseudodichroa apparently farthest away. Ammobatini. Pupae of the tribe have gone un- With respect to phylogeny, such larval features as described until now. the anterior placement of the posterior tentorial All nomadine bees are parasitic, the larvae pits in Pseudodichroa, the navicular shape of the feeding in the cells of various solitary bees. Am- predefecating larva ofPseudodichroa, the swollen mobatine bees even as adults are rare in collec- mandibular bases ofMorgania, and the strong an- tions. The acquisition of immature specimens for terior projection of the head features of Ammo- the study, therefore, would have been virtually bates are probably derived. However, too few lar- impossible without the excellent support of the val characters can be classified as being primitive National Science Foundation (grants GB-5407 or specialized to make possible an analysis of the and GB-32508) which we wish to acknowl- phylogeny of the tribe. We also note the simi- edge with appreciation. We also thank Ms. Liliane larity of the pupae of Morgania and Oreopasites. Floge and Mrs. Barbara Rozen who assisted in For all of these reasons we continue to follow the illustrations and Ms. Phyllis Browne who the more conservative classification, i.e., placing typed the manuscript. All specimens are in the the Ammobatini, sensu stricto, and Pasitini into a collection of the American Museum of Natural single tribe at least for the present, with the hope History. that a broad comprehensive analysis of all avail- 3 4 AMERICAN MUSEUM NOVITATES able characters will lead to a clear understanding and Scrapter seem paradoxical since both feed on of the phylogeny of the group and to an appro- the same semiliquid provisions. Loss or reduction priate classification. of the abductor apodeme is found among certain other colletids with watery nest provisions, e.g., Colletes and Euryglossa. LARVAE Another interesting point concerning Pseudo- The anatomy of bee larvae and techniques for dichroa is that both the anterior and posterior studying larvae have been discussed by Michener points of articulation of the mandible are some- (1953) and more recently by Rozen and McGin- what separate from the articulating points on the ley (1974). The posterior margin of the head head capsule. This fact plus the missing adductor capsule of the Ammobatini (excepting Pseudo- mandibular apodeme suggests that the mandibu- dichroa) and ofNomada possesses two transverse lar mechanism at least of the last instar may be weak intemal ridges, one anterior to the other structurally weak and the mandible may not be ("apparent posterior thickening" and "posterior required to function very actively during feeding. thickening," figs. 17, 24). This characteristic is This deduction would seem to be supported by also weakly developed in Paranomada but has the very short mandibles, the apexes of which not been investigated in Kelita. We presume the hardly reach the mouth area. On the other hand more anterior ridge to be the anatomical poste- larval mandibles of almost all the Nomadinae rior margin of the head capsule because the cap- (this apparently is not true of Paranomada and sule, so defined, would be normal in shape in Isepeolus) have nearly equally short mandibles lateral view for most bee larvae, and because the with adductor apodemes. This situation empha- sclerotization of the capsule ends at the anterior sizes that we know very little about how the ridge in Morgania. If this is the true anatomical short mandibles function, about what role the boundary of the head, then the sclerotized area swollen epipharynx of Nomada, Pseudodichroa, behind the posterior thickening in Ammobates, Ammobates and perhaps other Nomadinae play Pseudodichroa, and Oreopasites represents an ex- in the ingestion of food, and in general about the tension of the functional head capsule. Perhaps it feeding mechanisms of these larvae (or any bee is an ontogenetic holdover of an elongate head larvae). capsule of the first instar, an adaptation that Description of the Ammobatini could accommodate massive mandibular muscula- Based on Mature Larvae ture. In Pseudodichroa (fig. 8) the head exhibits Diagnosis. Mature larvae of the Ammobatini only a single transverse ridge. The placement of key readily to couplet 8 in Rozen (1966, the posterior tentorial pit midway along the hy- pp. 8-10) where Oreopasites and Nomada appear. postomal ridge seems to suggest that the true Because all ammobatines have a nonspiculate hy- posterior thickening has become obliterated but popharynx they can be separated from Nomada that the area from the posterior pits to the appar- which they tend to resemble. After the publica- ent posterior thickening is still homologous with tion of Rozen (1966), mature larvae of two other the extension of the head capsule found in other genera of Nomadini, Kelita and Paranomada, Ammobatini and in Nomadini. were collected. These demonstrate that the tribe We note that the mandible of Pseudodichroa is remarkably diverse (also reflected in the pupa (figs. 4-6) is peculiar because the abductor apo- of Paranomada) and raise intriguing questions re- deme is missing although the adductor apodeme garding the origin and monophyly of both the is reasonably well developed. The mandible of tribe and the subfamily. Although these larvae Scrapter (Colletidae), the host ofPseudodichroa, have not been studied in detail they can be has a well-developed adductor apodeme but the clearly distinguished from the Ammobatini abductor apodeme is absent, although the point because Kelita and Paranomada possess distinct of attachment of the abductor muscle is evident paired dorsal tubercles on most body segments. as a cuticular invagination of the mandibular cor- Furthermore their hypopharnyxes seem to be ium. These differences between Pseudodichroa spiculate as in the hypopharnyx ofNomada. ROZEN
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