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Novitates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2861, pp. 1-8, figs. 1-9 October 31, 1986 The Natural History of the Old World Nomadine Parasitic Bee Pasites maculatus (Anthophoridae: Nomadinae) and Its Host Pseudapis diversipes (Halictidae: Nomiinae) JEROME G. ROZEN, JR.1 ABSTRACT In nests of Pseudapis diversipes in Pakistan, a sickle-shaped mandibles used to eliminate the host series ofbranching tunnels surrounded vertical cells larva (or egg) and rival cuckoo bee larvae, as is clustered close to one another in the ground. In- the case with other nomadine parasitic bees. Pas- formation on the tunnel and cell walls, cell size, ites (Old World) and Oreopasites (New World) both eggs, and provisions is given. The cuckoo bee Pas- deposit eggs that are folded into a U as they are ites maculatus flew around the nest entrances; its inserted into the cell wall. This form of oviposi- eggs and first instars were recovered from host tion, a unique feature in the Apoidea, suggests a cells. Information on oviposition and egg size and close relationship between the two genera. structure is presented. The first instar had large INTRODUCTION Recently in Pakistan I discovered nests of cleptoparasite as Pasites maculatus Jurine a Pseudapis being visited by cuckoo bees of (Anthophoridae: Nomadinae). The literature the genus Pasites. Donald B. Baker kindly reveals little information on the nesting bi- identified the host as Pseudapis diversipes ology of the host and on the mode of para- (Latreille) (Halictidae: Nomiinae) and the sitism by Pasites. The following provides ' Deputy Director for Research and Curator, American Museum of Natural History. Copyright © American Museum of Natural History 1986 ISSN 0003-0082 / Price $1.10 2 AMERICAN MUSEUM NOVITATES NO. 2861 some specific data about these bees and per- acters. Variation may be extensive within pop- mits a comparison of the biology of Pasites ulations (the species is rarely taken in numbers with that of other Nomadinae. and appears always to be much less numerous The association of Pasites maculatus with than its host; I have never observed gradation), with other and, generally, variants within one population Pseudapis diversipes and perhaps can be matched by variants within other, dis- Pseudapis has been recorded in the European tant, populations. literature for nearly a century. Recently, Pasites maculatus may, as you suggest, be Hirashima and Nagase (1981) have associ- composite, but I cannot satisfactorily divide it ated Pasites esakii Popov and Yasumatsu on the basis of the material I have seen, and I with Pseudapis mandschurica (Hedicke) in do not think the question will be resolved until Japan. we have adequate series of specimens from Pseudapis, previously regarded as a sub- known hosts from a number of localities: host genus of Nomia, has recently been accorded data are lacking for almost all museum speci- generic status (Baker, personal commun.; mens, and host identities can rarely be inferred Hirashima, 1961; Hirashima and Nagase, from collection data since the species of Pseud- apis are numerous and since several species may 1981; Michener, personal commun.). Its nu- commonly be found flying together within one merous species extend from Europe to south- locality. (Even within the large area where Pseud- ern Africa and to Japan (Michener, 1961), apis diversipes is probably the sole host of mac- and into the Indian region and Indochina ulatus, variability of the latter is considerable; (Baker, personal commun.). Pasites, consist- but under these circumstances one would hardly ing ofonly a few nominal species, ranges from expect the variation to indicate the presence of Europe and North Africa to Japan (Popov, more than one species of parasite.) 1931; Hirashima -and Nagase, 1981). Infor- mation on the genus is limited, and the usage ACKNOWLEDGMENTS of various infraspecific names (see, for ex- ample, Hirashima and Nagase, 1981) sug- Ronald J. McGinley, Smithsonian Insti- gests that a more thorough study ofthe genus tution, Sarfraz Lodhi, American Museum of over its entire geographic range may lead to Natural History, and Ian Stupakoff, City a larger number of species and a more con- University ofNew York, participated on this vincing taxonomy. trip and aided in sampling the bee fauna at Donald B. Baker (in litt.) sent me his notes this nesting site. on Pasites maculatus and its host. They pro- I thank the following persons for carefully vide an informative overview of the taxo- reviewing this manuscript: Donald B. Baker, nomic problems with this cuckoo bee, par- Surrey, England; G. E. Bohart, Logan, Utah; alleling those I have had with the North and Charles D. Michener, Lawrence, Kansas. American ammobatine Oreopasites. The following individuals and their insti- tutions assisted us in various ways in fulfilling The range of Pasites maculatus extends from Northwest Africa to Japan. From Spain to Cen- this and other studies on Pakistan bees: Dr. tral Asia and Pakistan its certain principal host Rafiq Ahmad, National Coordinator, Hon- (and its probable principal host further afield, eybee Research Programme, Pakistan Agri- but not everywhere) is Pseudapis diversipes cultural Research Council, Islamabad, Pa- (Latr.). I have examined several thousand spec- kistan; Dr. Manzoor Ahmed, Department of imens of the host species, which shows rather Zoology, University ofKarachi, Pakistan; Dr. little geographical variation (as one might ex- Abdul Hameed Bajoi, Entomologist, Agri- pect in a species that has a more or less contin- cultural Research Institute, Quetta; and Dr. uous distribution and which usually occurs Riaz Ali Shah, Director General, Pakistan abundantly wherever it is found: I have the Museum ofNatural History, Islamabad. The impression that clinal variation tends to be more marked in species that have a more scattered field trip was made possible through the sup- distribution and smaller populations). I have port ofthe Smithsonian Institution's Foreign examined some hundreds of specimens of the Currency Program and I thank Dr. Gretchen Pasites, which shows considerable variation in Ellsworth, Director, Office ofFellowships and size and in colour, pubescence, sculptural and, Grants, at the Smithsonian, and her staff for to a comparatively small extent, structural char- assisting me in obtaining the support. 1 986 ROZEN: PASITES MACULA TUS AND PSEUDAPIS DIVERSIPES 3 Fig. 1. Nesting area of Pseudapis diversipes at Jalogir, 32 km north ofQuetta, Baluchistan, Pakistan. Ronald J. McGinley examining nest entrances in pathway in front of adobe wall. NESTING BIOLOGY OF a central area that was perhaps 5-8 cm hor- PSEUDAPIS DIVERSIPES izontally and 3 cm vertically. The vertical I found three nests, possibly interconnect- cells, one next to another, were clustered in ed, within 25 cm of one another on May 16, this enclosed area. Similar clusterings have 1984, at Jalogir, 32 km north of Quetta, Ba- been reported for both the Nomiinae and the luchistan, Pakistan. I hastily excavated two Halictinae with the open area surrounding of the nests, recovered immatures of both the cluster varying from tunnels (as here) to hosts and cuckoo bees, made brief notes on a mostly open chamber with a few earthen nest architecture, and preserved samples of pillars supporting the cluster (Stephen et al., cells and immatures for possible future study. 1969; Hirashima, 1961). A single nest may The nest entrances occurred on a packed dirt have included more than one such cluster (as pathway with loose surface soil next to an reported by Hirashima, 1961, for Nomia adobe wall surrounding an agricultural field (Hoplonomia) punctulata Dalla Torre) and (fig. 1) containing many flowering thistles, the possibly other cells not in clusters. Nest ar- possible food source of Pseudapis. The sub- chitecture was not studied in detail and ob- surface soil was fine and had no rocks and viously deserves further investigation. little organic material. Nest entrances were The second nest excavated consisted of a unshaded by vegetation for most of the day. burrow ofabout the same diameter descend- At least three females were associated with ing nearly vertically to the depth of about 6 one nest. cm at which point it started to branch, as was The main tunnel (5.0 mm in diameter) of the case with the first nest. one nest descended obliquely into the hori- The walls ofbranches were rough with nu- zontal surface and at a depth of 2 cm became merous indentations created by the females' vertical, branched at about the depth of 5 cm tamping the walls with their pygidial plates near a single cell, and then continued to de- (fig. 3). The impression of the median carina scend vertically to the depth of 15 cm. At this on the pygidial plate was clearly identifiable level it divided into a number of horizontal in each embossing (fig. 4). The walls, un- branches that extended in various directions, coated, absorbed water droplets immediate- abruptly turned downward and then appar- ly. ently coalesced so that the branches enclosed All cells (figs. 2, 9) were vertical, closure 4 AMERICAN MUSEUM NOVITATES NO. 2861 Fig. 2. Composite scanning micrograph of open cell of Pseudapis diversipes showing faint waves in cell wall and scratches near top of cell probably created by female Pseudapis attempting to remove Pasites egg; hole on right side thought to be puncture created by insertion of Pasites egg that later was destroyed by Pseudapis female. Fig. 3. Pseudapis diversipes, scanning micrograph of section of tunnel circumscribing cell cluster, showing embossings created by pygidial plate of female. Fig. 4. Same, more highly magnified; note median line created by median carina on pygidial plate. Fig. 5. Pasites maculatus, scanning micrograph of operculum embedded in cell wall of Pseudapis diversipes. end up, and spaced close to one another so parent lining that could be peeled in sections that intervening distances ranged from 1.5 to from the wall.
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