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PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3645, 12 pp., 19 figures June 25, 2009 Biology of the Bee Hoplitis (Hoplitis) monstrabilis Tkalcu˚ and Descriptions of Its Egg and Larva (Megachilidae: Megachilinae: Osmiini) JEROME G. ROZEN, JR.,1 HIKMET O¨ ZBEK,2 JOHN S. ASCHER,3 AND MOLLY G. RIGHTMYER4 ABSTRACT Herein we describe the nesting biology of the solitary ground-nesting bee Hoplitis (Hoplitis) monstrabilis Tkalcu˚ from eastern Turkey. Its shallow nests in the ground differ from the known nests of members of subgenus Hoplitis, most of which make mortar and pebble nests either on the exposed surfaces of rocks or within stems or other cavities. Cells are not lined with flower petals or other vegetative tissue, as expected for subgenus Hoplitis, but unlike other ground-nesting species of Hoplitis belonging to other subgenera such as Anthocopa. The egg of this bee is also described and illustrated, as is the fifth (last) larval instar. ABSTRAKT Bu c¸alıs¸mada bireysel yas¸ama sahip, toprakta yuva yapan Hoplitis (Hoplitis) monstrabilis Tkalcu˚’in yuva yapma biyolojisi Dog˘u Anadolu (Tu¨rkiye)’da c¸alıs¸ılmıs¸tır. Yuva, toprakta yu¨zeysel bir s¸ekilde yer almaktadır. Aynı alt cinse ait yuvaları bilinen tu¨rlerin yuvalarından az farklılık go¨stermekte, istisna olarak bu alt tu¨rde hu¨crelerin ic¸ cidarları c¸ic¸eklerin tac¸ yaprakları veya dig˘er bitkisel dokularla astarlanmamıs¸ durumdadır. Hoplitis (Hoplitis) monstrabilis’in yumurta ve bes¸inci (son) larva do¨nemi tanımlanmıs¸vec¸izimleri yapılmıs¸tır. 1 Division of Invertebrate Zoology, American Museum of Natural History ([email protected]). 2 Atatu¨rk University, Faculty of Agriculture, Department of Plant Protection, 25240 Erzurum, Turkey (hozbek@atauni. edu.tr). 3 Division of Invertebrate Zoology, American Museum of Natural History ([email protected]). 4 Department of Entomology, Smithsonian Institution, Washington, DC ([email protected]). Copyright E American Museum of Natural History 2009 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3645 INTRODUCTION on the campus (fig. 1). We excavated the two Hoplitis nests on July 11, 2007. The soil was This paper reports on the nesting biology consolidated and compact without visible and describes the mature larva of Hoplitis moisture in the vicinity of the cells, although (Hoplitis) monstrabilis Tkalcu˚ resulting from we found darker, moister soil starting at a depth the excavation of two nests found on the of about 5 cm. campus of Atatu¨rk University in Erzurum, J.S.A. located another Hoplitis nest, possi- Turkey. Andreas Mu¨ller identified the two bly of this species, on a bare (devoid of female voucher specimens, each taken from a vegetation) horizontal surface at 22 km south- nest, and additional females and one male west of Oltu, Erzurum, on July 7, 2007, but collected nearby. This recently described this nest was not studied. A photo of the species belongs to a small species group female at the nest entrance is available ˚ diagnosed by Tkalcu (2000), also including at http://www.discoverlife.org/mp/20p?see5I_ the sympatric H. erzurumensis Tkalcu˚ and H. JSA484. lapidaria (Morawitz). Males of this groups, NEST DESCRIPTION: Both nests on the cam- referred to here as the lapidaria species group, pus of Atatu¨rk University agreed in lacking have highly modified male hind legs (illustrat- tumuli around open entrances and in being ed by Tkalcu˚, 2000), with greatly expanded extremely shallow. Some cells were only 2– hind tibiae and inner tibial spurs. Females appear similar to those of subgenus Annosmia, 2.5 cm below the ground surface. The main and Warncke (1991) therefore placed H. tunnels, open their entire lengths, descended lapidaria, known only from the female, in that more or less diagonally with considerable subgenus. twisting and turning. There were no laterals We present information on H. monstrabilis (i.e., side branches) leading to cells; cells were to expand knowledge of ground-nesting be- attached directly to the main tunnels or to one havior in genus Hoplitis and to provide another. One nest contained five cells arranged evidence pertinent to the phylogenetic place- in two sets of two cells in a linear series and a ment of the lapidaria species group. More single open cell incompletely provisioned. The generally, we hope to augment our limited second nest contained a total of eight cells; the understanding of the biology of the Osmiini three closest to the ground surface were and of the range of larval variation within singletons, while the remaining five cells were Megachilidae. grouped as a descending series, one connected to the other, none separated by intercalary cells or short tunnel lengths (probably arranged like BIOLOGY OF HOPLITIS (HOPLITIS) those pictured by Westrich, 1989: 196, for MONSTRABILIS Hoplitis mocsaryi (Friese) [cited as Osmia DESCRIPTION OF NESTING SITE:OnJuly6, mocsaryi], though without petal linings). 2007, M.G.R. discovered the two nest entrances Cells (fig. 3) were oriented with their long of this species, about 15 cm apart, as both axis approximately horizontal, and all were females repeatedly entered and left their respec- identical in being very broad for their length; tive nests for short periods ranging from 30 sec. one measured 7.0 mm in maximum diameter to about 2 minutes apart. This activity suggest- and 11 mm long. Cells seemed symmetrical ed that they were removing soil and dropping it around their long axes, and their front and rear a short distance away, thus accounting for the ends were equally rounded. Each exhibited a lack of tumuli around the open entrances. The uniformly thin (ca 1–2 mm thick) cell wall ground surface was horizontal with sparse, low (fig. 4) composed of compact soil that tended to herbaceous vegetation between which the sur- separate here and there from the substrate face was barren (fig. 2). The site was within 1 m during our excavation. The inner wall surface of a dense but low patch of vegetation that had was smooth and matt (fig. 4), entirely lacking a grown over the excavated nesting site of visible reflective lining as found in many bee Rophites (Rophitoides) canus Eversmann that cells. However, it was waterproof; during had been studied in 2005 (Rozen and O¨ zbek, testing, a droplet of water remained beaded on 2008) at the edge of a seldom-used soccer field the surface for more than 2 min. None of the 2009 ROZEN ET AL.: BIOLOGY OF THE BEE HOPLITIS (ANTHOCOPA)3 Figs. 1–5. Photographs of nesting site and nest components of Hoplitis monstrabilis. 1, 2. Nesting site (arrow) of Hoplitis monstrabilis on campus of Atatu¨rk University, Erzurum, Turkey, and close-up of nest entrances (marked by dry plaster of Paris) identified by arrows, respectively. 3. Cell showing shape, lateral view from above. 4. Close-up of cell wall showing matt texture; note cell wall composed of compact soil (arrow). 5. Cell closure, inner view. cells was lined with petals or other vegetative approximately 5.0 mm in diameter, had a material as has been reported for other ground- thickness about equal to that of the cell wall, nesting species of genus Hoplitis species (see and were slightly concave on the inside, below). Five cell closures each measured conforming to the general curvature of the 4 AMERICAN MUSEUM NOVITATES NO. 3645 front end of the cell. When observed with a medium class in the classification of eggs/mature stereomicroscope, the inner surface of the oocytes relative to female bee body size, as closure lacked a distinct spiral pattern; it was developed by Iwata and Sakagami (1966: table smooth with some irregularities as if formed 2). The transparent chorionic surface viewed from very moist soil that soon dried after through a stereomicroscope was smooth and deposition (fig. 5). When tested with a droplet somewhat shiny. of water, the inner surface seemed water DEVELOPMENT: Eggs and young larvae were retardant though perhaps not as strongly so as on top of the provisions in the midline of the the cell wall. cells with their anterior ends (as determined by PROVISIONS: We collected no adults on the larvae) pointed toward the cell closures. The flowers, but the single type of pollen in the single fifth instar, still feeding, had cast skins of provisions was almost certainly that of the four previous instars clinging to its venter, Onobrychis (Fabaceae) because of its barrel- proving that there are five larval instars. All like shape. This plant grew abundantly near- instars possess apically bidentate mandibles, by. Of the well known European species of with the ventral tooth longer than the dorsal Hoplitis (Hoplitis), H. loti (Morawitz) and H. one. Only the fifth instar bears conspicuous ravouxi (Pe´rez) are oligolectic on Fabaceae, body setae, as has previously been found for whereas H. adunca, H. anthocopoides, and H. Coelioxys (Rozen and Kamel, 2008). lepeletieri are narrowly oligolectic on Echium CLEPTOPARASITES: None was associated with (Boraginaceae) (Westrich, 1989). Completed the two nests. provisions were an ovoid, homogeneous mass of orange pollen and nectar on the floor of the COMPARISONS WITH NESTING cell. Although appearing solid, the mass was BIOLOGY OF OTHER HOPLITIS so fluid that it deformed with the airflow from the aspirator used to blow away loose soil To what extent is the nesting biology of during our excavations. When the airflow Hoplitis monstrabilis similar to, or different stopped, it reformed as an oblong ovoid, thus from, that of other Hoplitis species? In attempt- suggesting the shape was dictated by the ing to explore this question, we have relied viscous nature of the mixture in combination heavily on the work of Westrich (1989) and with the shape of the lower part of the cell.

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