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. Michener (2007) for published sources of Interestingly, provisions placed in Kahle’s information about the biology of the genus solution with eggs or young larvae tended and on the review of our manuscript by Le Goff not to disassemble as one might expect due to (see Acknowledgments), who kindly directed their semiliquid condition. Freshly deposited our attention to the papers on Hoplitis provisions in the single cell still open consisted (Anthocopa) by Ferton (1891–1911) and who of mostly dry pollen, on top of which had been also supplied pictures of his own nest excava- deposited a large amount of partly clear tions of certain Anthocopa species (cited below nectar. Presumably, after bringing in the final as ‘‘Le Goff, personal commun.’’). A. Mu¨ller provision load, the foraging female mixes and and C. J. Praz generously shared unpublished shapes the mass into its ovoid form. information from their review of osmiine EGG DESCRIPTION AND DEPOSITION: Freshly nesting biology (a collaboration also including deposited eggs are white, i.e., the color of the Le Goff) and phylogeny. In addition to yolk as seen through their transparent chorions. identifying H. monstrabilis, their assistance Their shape (fig. 6) is elongate, gently curved, greatly facilitated our comparative review of parallel sided, and approximately equally its biology. We follow Michener’s (ibid.) rounded at both ends. One measured 3.05 mm classification of the Osmiini, and subsequent long and 0.875 mm in maximum diameter at updates by Praz et al.’s phylogenetic study approximate midlength. This species egg index (2008) and Ungricht et al.’s catalogue of is 77, as calculated by dividing the egg length by Palearctic Osmiini (2008), and compare species the average intertegular distance of 3.95 mm of of genus Hoplitis, especially those in subgenus the two adults collected (Iwata and Sakagami, Anthocopa, because this subgenus includes 1966; Rozen, 2003). This index falls into the many well-known, ground-nesting species, and 2009 ROZEN ET AL.: BIOLOGY OF THE BEE HOPLITIS (ANTHOCOPA)5

Figs. 6–11. Diagrams of immatures of Hoplitis monstrabilis. 6. Egg, lateral view, other orientation uncertain. 7. Entire body of last larval instar, lateral view, with enlarged view of abdominal apex showing protruding anal region. 8, 9. Head of last larval instar, front and lateral views, respectively, with close-up of antenna in maximum profile. 10, 11. Right mandible of same, dorsal view, and outer view of apex in maximum profile, respectively. to species of subgenus Hoplitis, which are 1954); H. idalia (Mavromoustakis) (Le Goff, generally surface or cavity nesters but are more personal commun.); H. jakovlevi (Radosz- closely related to H. monstrabilis. Nesting data kowski) (as serrilabris (Morawitz) (Banaszak for thirteen species of ground-nesting Hoplitis and Romasenko, 2001; Le Goff, personal (Anthocopa) are available: H. bisulca Gerst- commun.); H. longispina (Pe´rez) (Cros, 1937); aecker (as Osmia lanosa Pe´rez) (Ferton, 1895); H. mocsaryi (Friese) (Westrich, 1989); H. Le Goff, personal commun.); H. cristatula papaveris (Latreille) (Ferton, 1896 [as O. Zanden (as O. cristata Fonscolombe) (Ferton, papaveris Latreille]; Mu¨ller, 1907; Friese, 1893); Le Goff, personal comm.); H. cypriaca 1923); H. perezi Ferton (as O. perezi Ferton) cypriaca (Mavromoustakis) (Mavromoustakis, (Ferton 1895; Le Goff, personal commmun.); 6 AMERICAN MUSEUM NOVITATES NO. 3645

H. saundersi Vachal (as O. annulata Latreille and some Nearctic Hoplitis (Proteriades)nest and O. saundersi Vachal) (Ferton, 1891, 1911); in holes in the ground (Michener, 2007). H. similis (Friese) (Brauns, 1926; Michener, Hoplitis monstrabilis is the first report of a 1968, as H. anthodemnion Michener); H. Hoplitis (Hoplitis) nesting in the ground (A. singularis (Morawitz) (Marikovskaya, 1995); Mu¨ller, C. J. Praz, and G. Le Goff, review of H. villosa (Schenck) (Friese, 1923; Mu¨ller, Osmiini nesting biology in prep.), but this 1907; Petit, 1970, 1977; Westrich, 1989).5 behavior may be more widespread in the Our limited sample of nests of Hoplitis subgenus; the nesting biology of many species, monstrabilis was on an essentially horizontal including the two other species in the lapidaria surface of hard soil. However, Michener species group, remain unknown. (1968) found numerous nest entrances of H. Evidence is clear that the two females of (Anthocopa) similis in vertical clay banks, Hoplitis monstrabilis excavated their own nests whereas Brauns (1926) apparently retrieved since there were no remnants such as old nest them from a horizontal nesting site. This tunnels or cells at the site. However, we also suggests that inclination of the nesting surface know that within the Megachilinae, some taxa is not important for that species, and accord- nest in preexisting cavities including aban- ing to Le Goff (personal commun.) that is true doned nests from previous generations of both for most Anthocopa species that he studied. other taxa and conspecific taxa. Our sample Furthermore, degree of compactness of nest- of two nests was too limited to ascertain if ing soil may be of little importance since H. this species might also use pre-existing cavi- papaveris nests in both situations (Friese, ties. Among the other species of ground- 1923). By contrast, known species of subgenus nesting Hoplitis, many species probably Hoplitis, which are more closely related to H. avail themselves to preexisting cavities, but monstrabilis than the ground-nesting species if none is available, females perform their discussed above, are not ground nesting own excavations. Hoplitis anthocopoides but instead make mortar and pebble nests partly reuse previous years mortar and pebble adhering to the exposed surfaces of rocks nests after cleaning them of debris (Eickwort, or within stems or various cavities (Westrich, 1973). 1989). Nesting data are available for many A consistent feature of known nests of species in Westrich (1989, and references ground-nesting Hoplitis, including those of cited therein) including: Hoplitis adunca Hoplitis monstrabilis and all species of subge- (Panzer) (Westrich, 1989); H. anthocopoides nus Anthocopa, is their shallow depth, whether (Schenck) (see in particular Eickwort, 1973), the nesting surface is horizontal or vertical. H. ravouxi (Pe´rez) (Westrich, 1989). Among This seems a remarkable feature since such known species of subgenus Hoplitis, the surfaces lose moisture readily and are subject- majority, including the well known H. antho- ed to excessive heat and cold; bees hibernating copoides, make mortar and pebble nests and/or aestivating under these conditions for adhering to exposed rock surfaces, and initi- many months would seem to be at risk from ated in a rock crevice, whereas others are more desiccation or extreme temperatures. How- flexible, building nests within various cavities ever, this is not a feature unique to Hoplitis: such as holes in wood or stems or in the soil as many groups of megachilid bees cope well does H. adunca. Hoplitis fertoni (Pe´rez) is under these circumstances, for example, exceptional as it nests in snail shells (Le Goff, Megachile in trap nests (Rozen and Kamel, 2003). Additional Palearctic subgenera of 2008: figs. 1–3), and Hoplitis (Hoplitis) in mud Hoplitis including Annosmia and Pentadent- nests adhering to rocks. osmia (involving numerous desert-inhabiting In common with other species of Hoplitis species) are ground nesting (Praz et al., 2008), (Hoplitis), but in contrast to most known species of subgenus Anthocopa, H. monstrabilis 5 In addition, Marikovskaya (1995) has published do not line its brood cells and nest tunnels with biological information on Hoplitis (Anthocopa) singularis blossom petals or other vegetative tissue (Morawitz), the type species of the formerly recognized subgenus Glossosmia, but we have been unable the secure (Westrich, 1989). Cells of H. monstrabilis need a copy of the paper. further study to determine the nature of the 2009 ROZEN ET AL.: BIOLOGY OF THE BEE HOPLITIS (ANTHOCOPA)7

Figs. 12, 13. Photographs of spiracle on cleared larva of Hoplitis monstrabilis. 12. Side view, showing elevation of rim and globular shape. 13. Oblique lateral view, showing peritreme and faint atrial ridges. thin cell wall of soil: is it a special clay coating of H. monstrabilis were almost horizontal, transported into the cell and applied by the those of H. (Anthocopa) similis ranged in female, or is it merely the result of the female inclination from 20u–60u from horizontal smoothing the inner surface of the newly (Michener, 1968). The illustrations of Ferton excavated cell? A chance recovery of a cell in (1895: figs. 7, 12, 13) and photographs of Le construction might provide the answer. Goff (personal commun.) would seem to Both nests of Hoplitis monstrabilis were suggest that within the subgenus cell inclination actively being provisioned when found and and arrangement relative to one another may excavated, so we can offer no information be significantly variable. concerning its nest closure. With respect to In studies of solitary bees, we tend to ground-nesting Anthocopa species, Westrich consider whether some species mate at the (1989) states that nest entrances of H. flowers where females gather pollen and mocsaryi are left open after the female departs. nectar and/or at the nesting site where females Nests of H. papaveris are closed with sand are emerging and returning. In the case of (Friese, 1923); the nest tunnel of H. similis is Hoplitis monstrabilis we saw no males at the filled with soil to the ground surface nesting site, and collected only a single male (Michener, 1968), which is apparently also in the general area. This hints that mating the case with H. papaveris (Mu¨ller, 1907). Nest may take place earlier in the season in the closures and cell partitions of H. singularis are vicinity of flowers. Hoplitis (Hoplitis) species formed from masticated green leaves such as anthocopoides patrol and defend (Marikovskaya, 1995). territories consisting of blooming host plants Although some Hoplitis nests consist of only and rock or bare soil resting perches; females single cells, other species including H. mon- mate a single time when visiting flowers strabilis construct nests of more than one cell. (Eickwort, 1977). The nest of H. monstrabilis with eight cells Although we found no cleptoparasites asso- might have become larger had we not excavat- ciated with Hoplitis monstrabilis, other species ed it and collected the female. (Eickwort, 1973, of Hoplitis are reportedly parasitized by species cited a mean number of cells in new H. of Coelioxys, Stelis,andDioxys (Westrich, anthocopoides nests of 7.2, SE = 0.86, n = 29). 1989; Banaszak and Romasenko, 2001). Small nests imply that many species of ground- Because our nest sample was so small, we nesting Hoplitis (Anthocopa) normally make would not be surprised if future studies were to more than one nest, as was also concluded by reveal that cleptoparasitic bees are associated Le Goff (personal comm.). Although the cells with H. monstrabilis. 8 AMERICAN MUSEUM NOVITATES NO. 3645

DESCRIPTION OF LAST LARVAL articulation), internal head ridges at articula- INSTAR tion with mandibles, articulating arms of stipites, points of articulation of cardines with Figures 7–19 stipites, dorsal surface of premental sclerite between attachment of articulating arms of The following description, the first for any stipites; other areas scarcely pigmented; fine ground-nesting Hoplitis (Hoplitis) and for the spiculation apparently restricted to dorsal lapidaria species group, is based on a single surface of maxilla at its base and to lateral last-stage larva that was still feeding on lobes of hypopharynx. Area immediately provisions when preserved. Because last in- above hypostomal ridge and just behind stars gradually develop head pigmentation posterior mandibular articulation not pro- and grow robust as they ingest provisions, duced as downward-directed tubercle as pres- neither the pigmentation nor body form ent in many Coelioxys (Rozen and Kamel, described here can reliably predict head 2007: fig. 47). Hypostomal ridge giving rise to pigmentation and body shape of a postdefe- dorsal ramus that extends posteriorly from cating larva. This specimen agrees with the middle of ridge nearly to postoccipital ridge preliminary description of mature larvae of (fig. 9) in association with inflection of pari- the Megachilini in Rozen and Kamel (2007: etal at deeply set posterior tentorial pit; 19) unless stated otherwise. See that descrip- anterior tentorial pit distinctly closer to tion for details not presented below. anterior mandibular articulation than distance In studying the specimens, we first drew it between it and basal ring of antenna; episto- for the larval diagrams and then cleared the mal ridge extending dorsally for short distance specimens in a hot aqueous solution of sodium mesad of anterior tentorial pit. Parietal bands hydroxide. After lightly staining the cleared clearly evident. Diameter of basal ring of specimen with Chlorazol Black E and trans- antenna small, somewhat less than twice ferring it to a glycerin-filled well slide, we distance from ring to center of anterior added details to the illustrations. We then tentorial pit; antennal papilla (figs. 9, 14, 15) prepared the head capsule for SEM examina- small, slender, gradually, evenly tapering tion by critical-point drying and coating it apically, about three times as long as basal with gold-palladium alloy. diameter, bearing three sensilla. Apical margin DIAGNOSIS: Although the apically attenu- of clypeus strongly angled upward at midline ate mandibles (figs. 11, 17) of this specimen (fig. 16), so that at midpoint margin at to appear distinctive from those of most other slightly above level of anterior tentorial pits.6 described megachilids, this condition might be Labral sclerite unpigmented, with lower mar- ephemeral, disappearing as the mandibles gin poorly defined, but sclerite obviously wear. In other respects, this larva seems transverse (fig. 8); labrum unpigmented, with- almost indistinguishable from other known out pigmented median spot extending from nonparasitic members of the Megachilinae, labral sclerite to apical labral margin as in including previously described species of Coelioxys (Rozen and Kamel: 2007: figs. 44, Hoplitis (Hoplitis) (Eickwort, 1973, and refs. 45); apical labral margin broad, weakly therein), and emphasizes the overall homoge- concave (fig. 16). neity of last larval instars of the subfamily Mandible (figs. 10, 11, 17) moderately Megachilinae. elongate, apically attenuate, curved, and DESCRIPTION: Head (figs. 8, 9, 14–19): slender; apex bidentate with ventral tooth Setae long; those of parietals widely and considerably larger and longer than dorsal sparsely scattered, erect, more or less curved, tooth; both teeth acutely pointed; dorsal and with large, slightly elevated alveoli at base (figs. 18, 19); those of maxillary and labial 6 The apparent clypeal/labral discrepancy between figs. 8 apices straight, forward projecting and those and 16 is presumably due to the retraction of the labrum of labral apex decumbent, downward directed. resulting from critical-point drying for SEM examination. The soft conjunctive connecting the lower end of the Following areas moderately to faintly pig- clypeus to the labrum folded in, causing the retreat of the mented: mandibles especially at apices, inter- labrum. Thus the strong angling of the lower clypeal nal mandibular ridges (especially at points of margin is revealed in fig. 16. 2009 ROZEN ET AL.: BIOLOGY OF THE BEE HOPLITIS (ANTHOCOPA)9

Figs. 14–19. SEM micrographs of mature larva of Hoplitis monstrabilis. 14. Right side of head showing antenna, anterior tentorial pit, part of labrum, and cranial setae, frontal view; rectangle identifying seta enlarged in fig. 18. 15. Close-up of antenna, showing three apical sensilla. 16. Apex of clypeus and labrum, frontal view. 17. Mouthparts, including salivary opening, frontal view. 18, 19. Two long cranial setae, showing large, elevated alveoli identified by arrows; fig. 18, from side of parietal at level of lower edge of parietal band, identified by rectangle in fig. 14; fig. 19 from upper area of parietal band. 10 AMERICAN MUSEUM NOVITATES NO. 3645 apical edge of dorsal tooth finely, irregularly erately narrow, so that atrial opening distinct- serrate when viewed with compound micro- ly wider than peritreme width; atrial inner scope (not clearly visible with stereomicro- surface with faint rows (fig. 13) concentric scope); ventral apical edge of lower tooth less with primary tracheal opening; these rows distinctly serrate; inner apical surface shallow- with fine ridges which may have occasional ly concave; cuspal area not developed; outer sharp, fine projections (alternatively, fine surface with single seta not borne on tubercle. ridges may occasionally trap and hold fine Cardo and stipes well-developed sclerites but debris); primary tracheal opening with collar; not pigmented except for articulating arm of subatrium moderately robust; subatrium mod- stipes; maxillary palpus small, about as long as erately short, with 7–10 chambers. Male sex surrounding setae, perhaps slightly larger than characters unknown; female with paired, antennal papilla. Apex of labium normally ventral, paramedian integumental scars on wide (figs. 8, 17); premental sclerite not abdominal segments 7–9, with those of seg- evident laterad of attachments of articulating ment 7 farthest apart and those of segment 9 arms of stipites because of lack of pigmenta- closest together. tion, but probably well represented on post- MATERIAL EXAMINED: One fifth instar, defecating form since labium clearly divided Turkey: Erzurum, VII-11-2007, J.G. Rozen, into prementum and postmentum; maxillary H. O¨ zbek, M.G. Rightmyer, J.S. Ascher. and labial palpi subequal in length. Salivary REMARKS: The moderately elongate, apical- lips projecting, broadly transverse, width ly curved and attenuate mandibles of this about equal to distance between bases of species suggest that they might serve as a labial palpi; inner surface of lips, visible only defensive weapon against larval cleptoparasites, after specimen subjected to critical-point although no cleptoparasite has yet been associ- drying process, with numerous parallel, raised ated with this species. However, their length ridges extending outward. Hypopharynx con- may wear down with use, as seems to be the case sisting of widely separated lateral lobes that with last instars of other megachilid taxa. are spiculate; area between them irregular but nonspiculate. ACKNOWLEDGMENTS Body (fig. 7): Body setae rather short and inconspicuous compared with last larval in- J.S.A. and J.G.R.’s participation in the stars of many other megachilids, but still quite fieldtrip that resulted in this paper was visible, and with distinct alveoli compared supported by the Robert G. Goelet Bee with last larval instars of other families; Fieldtrip Fund through the American pleural area of abdominal segment 8 with Museum of Natural History. M.G.R.’s field approximately 5 setae; integument with scat- support came from the World Wildlife Fund. tered patches of fine spicules. Body form We thank Suzanne Rab-Green, Nadine robust; intersegmental line weakly incised Dupe´rre´, Eileen Westwig, and Boris Zakharov, because of bloated body shape; intrasegmental American Museum of Natural History, for lines not evident but possibly visible on kindly translating into English passages of postdefecating form; paired body tubercles books and papers written in German, French, absent; middorsal body tubercles more or less and Russian, thereby enabling comparisons evident on abdominal segments 2 and 3 and with many of the previous studies on Hoplitis. possibly elsewhere; pleural swellings poorly Susanne Rab-Green also assisted in the analysis developed in early fifth instar; abdominal of resulting data. Steve Thurston, scientific segment 10 attached to approximate middle assistant, American Museum of Natural of segment 9; anus apparently positioned History, expertly prepared the illustrations for toward top of segment 10 on projection, publications. Maximilian Schwarz, Linz, which may be eversible, as perhaps character- Austria, provided essential literature. istic of many cocoon-spinning bees. Spiracles The manuscript has been improved by the well sclerotized, unpigmented, subequal in careful reviews of two anonymous outside size; globular atrium projecting well above readers, one of whom (Ge´rard Le Goff) shared body wall, with rim (fig. 12); peritreme mod- with us extensive information from his field 2009 ROZEN ET AL.: BIOLOGY OF THE BEE HOPLITIS (ANTHOCOPA)11 notes concerning the nesting biology of Hoplitis. series. Annales de la Socie´te´ Entomologique de We are extremely grateful to Andreas Mu¨ller France 77: 535–586. Ferton, C. 1911 [1912]. Notes de´tache´s sur l’instinct and Cristophe J. Praz for identifying Hoplitis e monstrabilis and for generously sharing unpub- des Hyme´nopte`res mellife`res et ravisseurs, 7 lished information from their ongoing review of series. Annales de la Socie´te´ Entomologique de France 80: 351–412. the comparative biology of Osmiini (with G. Le Friese, H. 1923. Die europa¨ischen Bienen. Berlin: Goff) and studies of osmiine phylogeny. Walter de Gruyter, 456 pp. Iwata, K., and S.F. Sakagami. 1966. Gigantism and dwarfism in bee eggs in relation to the mode of REFERENCES life, with notes on the number of ovarioles. Japanese Journal of Ecology 16: 4–16. Banaszak, J., and L. Romasenko. 2001. Megachilid Le Goff, G. 2003. Note sur la nidification d’Hoplitis bees of Europe. 2nd ed. Poland: Bydgoszez (Hoplitis) fertoni Pe´rez dans la province University Press, 239 pp. d’Alicante (Espagne) (Hymenoptera: Apoidea; Brauns, H. 1926. V. Nachtrag zu ‘‘Friese, Bienen Megachilidae; Osmiini). Cette abeille est un Afrikas.’’ Zoologische Jahrbu¨cher Abteilung nouvel ho´te pour Chrysis hybrida Lepeletier fu¨r Systematik O¨ kologie und Geographie der (Hymenoptera; Chrysidoidea; Chrysididae; Tiere 52: 187–230. Chrysidini. 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