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Hoplitis (Formicapis) excisa (Morawitz 1880) Osmia excisa Morawitz 1880: 385. Type material: Holotype ♀, ���Ordoss am Chuan-che��� (China), type depository unknown. New subgeneric placement. New records. CHINA: Jingangling, 50km W Linfen, 29.��� 30.5.1996 (leg. J. Halada). Distribution. Known so far only from the Inner Mongolia Autonomous Region and Shanxi province in northern China. Pollen hosts. The only three pollen loads available so far (from the same locality in northern China) contained Rosaceae pollen: two loads exclusively consisted of pollen of the Potentilla type (either Fragaria or Potentilla), while one load additionally contained Rosaceae pollen of unknown affiliation. Nesting biology. Unknown., Published as part of M��ller, Andreas & Mauss, Volker, 2016, Palaearctic Hoplitis bees of the subgenera Formicapis and Tkalcua (Megachilidae, Osmiini): biology, taxonomy and key to species, pp. 105-120 in Zootaxa 4127 (1) on page 110, DOI: 10.11646/zootaxa.4127.1.5, http://zenodo.org/record/259694, {"references":["Morawitz, F. (1880) Ein Beitrag zur Bienen-Fauna Mittel-Asiens. Bulletin de l'Academie Imperiale des Sciences de St. - Petersbourg, 26, 337 - 389."]}

Subgenus Formicapis Sladen 1916 Hoplitis excisa (Morawitz 1880) was treated as a representative of the subgenus Stenosmia by Mavromoustakis (1954), Wu (2006) considered it to be a member of Osmia (Osmia) and Ungricht et al. (2008) assigned it to Hoplitis subgenus incertum. The recent discovery of the hitherto unknown male clearly revealed its affiliation to the subgenus Formicapis. The Central Asian species Hoplitis latifemoralis (Wu 1985) was formerly considered to belong to Formicapis (Wu 1985, 2006; Ungricht et al. 2008). Both sexes, however, strongly differ morphologically from H. (Formicapis), e.g. in the form of the female mandible, which is narrow and three-toothed lacking the long apical edge typical of Formicapis, or in the shape of male tergum 7, which is quadrangular and medially slightly incised instead of four-lobed as in Formicapis. Furthermore, H. latifemoralis substantially differs from H. (Formicapis) species in the form of the male sterna and male genitalia. Due to these differences and the fact that H. latifemoralis possesses some exceptional characters, such as a thickened male hind femur and tibia, the presence of hooked bristles on the female proboscis and the preference for small-flowered Boraginaceae species as pollen hosts (M��ller 2016), H. latifemoralis is removed here from H. (Formicapis) and assigned to Hoplitis subgenus incertum., Published as part of M��ller, Andreas & Mauss, Volker, 2016, Palaearctic Hoplitis bees of the subgenera Formicapis and Tkalcua (Megachilidae, Osmiini): biology, taxonomy and key to species, pp. 105-120 in Zootaxa 4127 (1) on page 109, DOI: 10.11646/zootaxa.4127.1.5, http://zenodo.org/record/259694, {"references":["Sladen, F. W. L. (1916) Bees of Canada - Fam. Megachilidae. The Canadian Entomologist, 48, 269 - 272. http: // dx. doi. org / 10.4039 / Ent 48269 - 8","Morawitz, F. (1880) Ein Beitrag zur Bienen-Fauna Mittel-Asiens. Bulletin de l'Academie Imperiale des Sciences de St. - Petersbourg, 26, 337 - 389.","Mavromoustakis, G. A. (1954) New and interesting bees (Hymenoptera, Apoidea) from Israel. Bulletin of the Research Council of Israel, 4, 256 - 275.","Wu, Y. (2006) Hymenoptera: Megachilidae. Fauna Sinica, Insecta, 44, 1 - 474 [in Chinese with English summary]","Ungricht, S., Muller, A. & Dorn, S. (2008) A taxonomic catalogue of the Palaearctic bees of the tribe Osmiini (Hymenoptera: Apoidea: Megachilidae). Zootaxa, 1865, 1 - 253.","Wu, Y. (1985) The Biology of the Tianshan-Tumur Region. People Press, Wulumuqi, pp. 139 - 150 [in Chinese with English summary]","Muller, A. (2016) Palaearctic osmiine bees - systematics and biology of a fascinating group of solitary bees. ETH Zurich. Available from: http: // blogs. ethz. ch / osmiini (accessed 2 March 2016)"]}

Subgenus Tkalcua Kocak & Kemal 2010 H. (Tkalcua) species were formerly assigned to the subgenus Microhoplitis Tkalcu (Ungricht et al. 2008). As the subgenus name Microhoplitis Tkalcu is a junior homonym of the lepidopteran genus name Microhoplitis Marumo, Kocak & Kemal (2010) proposed Tkalcuaas replacement name., Published as part of M��ller, Andreas & Mauss, Volker, 2016, Palaearctic Hoplitis bees of the subgenera Formicapis and Tkalcua (Megachilidae, Osmiini): biology, taxonomy and key to species, pp. 105-120 in Zootaxa 4127 (1) on page 112, DOI: 10.11646/zootaxa.4127.1.5, http://zenodo.org/record/259694, {"references":["Kocak, A. O. & Kemal, M. (2010) Generic replacement names in the family Megachilidae (Hymenoptera). Priamus, 12, 127 - 128.","Ungricht, S., Muller, A. & Dorn, S. (2008) A taxonomic catalogue of the Palaearctic bees of the tribe Osmiini (Hymenoptera: Apoidea: Megachilidae). Zootaxa, 1865, 1 - 253."]}

Hoplitis (Tkalcua) zandeni (Teunissen & van Achterberg 1992) Osmia zandeni Teunissen & van Achterberg 1992: 313. Type material: Holotype ��, ���Puerto del Rosario, Fuerteventura, 4.��� 16.3.1984 ��� (Spain: Canary Islands), Naturalis Biodiversity Center, Leiden. Osmia (Microhoplitis) hohmanni Tkalcu 1993: 812. Type material: Holotype ��, ���Fuerteventura, Risco del Paso, 1.5.1988 ��� (Spain: Canary Islands), La Roche Collection. Type species of Microhoplitis Tkalcu. Synonymy in Zanden (1995). Literature records. SPAIN, Canary Islands: Teunissen & van Achterberg (1992), Hohmann et al. (1993) and Tkalcu (1993) list numerous records of H. zandeni from Fuerteventura and Lanzarote. These records originate mainly from coastal areas not exceeding 150m a.s.l. with the exception of two records at altitudes of 250m and 400m, respectively, and encompass a time period that lasts from the beginning of February to the beginning of May. New records. SPAIN, Canary Islands: Fuerteventura, Monta��a Pelada SW Costa Calma, 29.3.��� 5.4.2015 (leg. A. M��ller & V. Mauss); Fuerteventura, Istmo de la Pared NW Costa Calma, 4.4. 2015 (leg. A. M��ller & V. Mauss). Distribution. Endemic on the two neighbouring Canary Islands Fuerteventura and Lanzarote. Habitat. Semidesertic areas on sandy ground with scattered shrubs of Salsola divaricata (Amaranthaceae), Launaea arborescens (Asteraceae) and Zygophyllum fontanesii (Zygophyllaceae) (Figs. 9, 10). Pollen hosts. Polylectic (Tab. 1). Pollen sources as deduced from field observations and the analysis of both female pollen loads and brood cell contents at the study sites on Fuerteventura were different species of Fabaceae (e.g. Astragalus hamosus, Lotus glinoides, L. lancerottensis, Ononis natrix), Reseda lancerotae (Resedaceae), Frankenia laevis (Frankeniaceae), Heliotropium erosum (Boraginaceae), Helianthemum canariense (Cistaceae) and Kickxia sagittata (Plantaginaceae). The females collected pollen from the flowers of Reseda, Frankenia and Helianthemum by rapidly seesawing their metasomal scopa against the anthers, whereas pollen of Heliotropium was removed out of the narrow flower tubes with alternating up and down movements of the forelegs; in contrast to other bee species that collect Heliotropium pollen, the females of H. zandeni are not equipped with specialized bristles to extract pollen from the hidden anthers (see Gotlieb et al. 2014). Interestingly, flowers of Asteraceae do not seem to be exploited for pollen, although several Asteraceae species were flowering at the study sites; thus, the observation of H. zandeni on flowers of the Asteraceae species Launaea arborescens and Phagnalon sp. (Hohmann et al. 1993) probably refers to individuals that sucked nectar rather than collected pollen. Nesting biology. Nesting site: All 19 nests detected near Costa Calma on Fuerteventura from 29 March to 5 April 2015 were in snail shells of Theba geminata (Mousson). Nests were found in both juvenile and adult shells varying in maximal diameter from 11mm to 15mm. All shells selected by the female bees for nesting lay openly on the ground and were not hidden e.g. among vegetation or under stones. Their position on the ground varied with some shells laying on the umbilicus and others on the apex. As the females were never observed to move nor turn the shells during any phase of the nesting cycle, the free accessibility of the shell opening appears to be an important condition for nest site selection. A second condition pertained to the degree to which the shells were filled with sand. Due to the sandy ground and the permanent wind prevailing at the study sites, almost all shells of T. geminata were more or less filled with sand. Females searching for nesting sites were repeatedly observed to scratch sand out of the shell or even to burrow through sand masses blocking the shell entrance, probably to check whether the inner whorls of the shell are accessible for nesting. Shells with their openings being blocked with hardpacked sand or with their inner whorls being densely filled with sand were never chosen as nesting sites. Nesting cycle: After selection of a suitable snail shell for nesting, the females usually removed sand from the shell entrance to get free access to the inner shell whorls. In one case, this cleaning process lasted 8 min. The shells, however, were never completely cleaned from sand with substantial amounts of sand remaining in the entrance area (Fig. 11). Brood cell construction started with the collection of chewed leaf tissue (���leaf pulp���) on both green and yellowed leaves of Helianthemum canariense and possibly also other plant species (Fig. 17). In contrast to other snail shell nesting osmiine bees, e.g. Osmia species of the subgenera Neosmia, Allosmia or Helicosmia (M��ller 2016), H. zandeni did not glue leaf pulp onto the shell surface. Instead, the collected leaf pulp was used i) to tightly fix sand grains still present in the area of the later brood cell to the shell wall and ii) to build - under addition of many sand grains - a narrow and crescent-shaped brink of varying width along the columella of the shell (Fig. 15). The construction of such thresholds, which define the length of the brood cell in advance and are later expanded to the partition that seals the brood cell (Fig. 16), is widespread among osmiine bees (Philipps & Klostermeyer 1978; Frohlich 1983; M��ller 1994). In one case, H. zandeni needed 10 flights with nest building material to construct the threshold. The provenance of the numerous sand grains glued together with leaf pulp for the construction of thresholds and cell partitions is not fully clear. Careful examination of photos of females returning from leaf pulp collection flights revealed that sand grains adhered to the leaf pulp packets held by the females in their mandibles. This suggests that the females landed on the ground immediately after having chewed leaf pulp from the Helianthemum leaves, dipped the moist leaf pulp in the sand and transported a mixture of both leaf pulp and sand grains back to their nest. It can not be excluded, however, that part of the sand grains used for nest construction originated from sand remains in the shell entrance area itself. After the threshold had been finalized, the brood cell was provisioned with nectar and pollen. Females returning from provisioning flights, which lasted between 8 min and 20 min, entered the nest head first to regurgitate nectar before they came out, turned around at the nest entrance (Fig. 12) and entered the nest metasoma first to comb pollen out of the scopa. In larger shells, the female turned around inside the shell rather than at the shell opening. Immediately after egg deposition the threshold was expanded to a cell partition. If a second brood cell was built in front of the first, the cell partition was of moderate and uniform width of 0.75���1.25mm. If, however, no additional cell was constructed, the threshold was expanded to a thick nest plug, which had a maximal width of 2���4.5mm along the shell wall and of 1���2mm in its centre (Fig. 13). The inner side of the nest plug consisted of sand grains irregularly glued together (Fig. 14), whereas the outer side was carefully worked with sand grains and leaf pulp forming a plane wall (Fig. 16). In one case, a female needed 37 flights with nesting material to expand the threshold to a nest plug. After completion of the nest plug, the females left the nest and started searching for another shell to construct additional brood cells. In contrast to other snail shell nesting osmiine bees, e.g. Osmia species of the subgenera Allosmia, Hoplosmia or Neosmia (M��ller 2016), completed nests were neither transported to a safe place nor buried into the ground nor turned in a protected position nor covered with plant material. In one case, a female selected a shell for nesting at 1315 pm and left the completed nest containing a single brood cell at 1610 pm the following day, suggesting that the construction of a one-celled nest usually lasts about one day under favourable conditions. Nest architecture: Of the 19 nests examined, 10 contained a single brood cell (Figs. 13, 14), 6 contained two cells and 2 had an empty cell in front of the first brood cell containing moderate amounts of sand (the number of brood cells in one nest could not be assessed as the female had just started to provison her first cell). Interestingly, the nest plug in H. zandeni was never built at the shell opening as is the case in all osmiine bee species, which use snail shells as exclusive nesting sites (A. M��ller unpublished). Instead, it was built 1 / 3 to 1 whorl behind the shell opening and thus was usually not visible from the outside. The fact that the eumenine species Leptochilus fortunatus Bl��thgen, which also nested in shells of Theba geminata at the study sites, built the nest plug deep inside the shells like H. zandeni suggests that habitat conditions might underlie this unusual behaviour in either species. One possible explanation is that plug construction at the shell opening necessitates the prior removal of sand masses adhering to the shell entrance area, which might be too costly in terms of time, as this sand was often hard packed or tightly adhered to the shell wall. Alternatively, sealing the shell at its opening might not contribute to a better protection of the progeny as the everlasting wind on Fuerteventura is expected to fill the open entrances of the completed nest with sand within a short period of time. Brood parasites: At the study sites, many specimens of Chrysis hohmanni Linsenmaier searched snail shells for putative host nests. As C. hohmanni has the same size and was equally common as H. zandeni at the study sites, H. zandeni is assumed to be its specific host. It can not be excluded, however, that C. hohmanni parasitizes the nests of L. fortunatus, which, however, is larger than the chrysidid wasp and was only rarely encountered. Behaviour. Flight activity: H. zandeni was active also during windy, rather cool and cloudy weather when most other bee species had ceased their activities. H. zandeni thus seems to be well adapted to the windy conditions and often moderate temperatures prevailing on Fuerteventura during the species��� main flight period from February to April. One possible adaptation underlying this moderate sensitivity against unfavourable weather conditions might be the special flight behaviour of H. zandeni: both sexes used to fly very close to the sandy ground and often landed on the ground for shorter or longer resting periods (see below), thus probably benefitting from the heat emission of the sand. Male mating behaviour: On their search for females, males patrolled along more or less fixed routes close to the ground, which were regularly interrupted by short resting periods on the ground (Fig. 18). Mating attempts were observed both on the ground and on flowers. In one case, a male constantly patrolled flowers of Frankenia and Reseda along a fixed circular route of 1.5m diameter around a low shrub and landed once per passage at exactly the same same spot to rest for a few seconds. Females also regularly interrupted their provisioning flights by short resting periods on the ground. As the frequency and length of these resting periods in both sexes increased with decreasing temperature and increasing wind velocity, this behaviour probably served to heat up the body on the warm sandy ground. Sleeping places: Females passed the night and bad weather in their own nest, while males were observed to sleep singly in snail shells., Published as part of M��ller, Andreas & Mauss, Volker, 2016, Palaearctic Hoplitis bees of the subgenera Formicapis and Tkalcua (Megachilidae, Osmiini): biology, taxonomy and key to species, pp. 105-120 in Zootaxa 4127 (1) on pages 114-116, DOI: 10.11646/zootaxa.4127.1.5, http://zenodo.org/record/259694, {"references":["Teunissen, H. G. M. & van Achterberg, C. (1992) Osmia zandeni a new species from Fuerteventura, Canary Islands (Hymenoptera: Apidae: Megachilinae). Zoologische Mededelingen (Leiden), 66, 313 - 315.","Tkalcu, B. (1993) Neue Taxa der Bienen von den Kanarischen Inseln. Mit Bemerkungen zu einigen bereits bekannten Arten (Insecta, Hymenoptera, Apoidea). Veroffentlichungen aus dem Ubersee-Museum Bremen, 12, 791 - 858.","Rasmont, P., Ebmer, P. A., Banaszak, J. & Zanden, G. van der (1995) Hymenoptera Apoidea Gallica: Liste taxonomique des abeilles de France, de Belgique, de Suisse et du Grand-Duche de Luxembourg. Bulletin de la Societe Entomologique de France, 100 (hors serie), 1 - 98.","Hohmann, H., La Roche, F., Ortega, G. & Barquin, J. (1993) Bienen, Wespen und Ameisen der Kanarischen Inseln. Veroffentlichungen aus dem Ubersee-Museum Bremen, 12, 1 - 894.","Gotlieb, A., Pisanty, G., Rozen, J. G., Muller, A., Roder, G., Sedivy, C. & Praz, C. (2014) Nests, floral preferences, and immatures of the bee Haetosmia vechti (Hymenoptera: Megachilidae: Osmiini). American Museum Novitates, 3808, 1 - 20. http: // dx. doi. org / 10.1206 / 3808.1","Muller, A. (2016) Palaearctic osmiine bees - systematics and biology of a fascinating group of solitary bees. ETH Zurich. Available from: http: // blogs. ethz. ch / osmiini (accessed 2 March 2016)","Phillips, J. K. & Klostermeyer, E. C. (1978) Nesting behavior of Osmia lignaria propinqua Cresson (Hymenoptera: Megachilidae). Journal of the Kansas Entomological Society, 51, 91 - 108.","Frohlich, D. R. (1983) On the nesting biology of Osmia (Chenosmia) bruneri. Journal of the Kansas Entomological Society, 56, 123 - 130.","Muller, A. (1994) Die Bionomie der in leeren Schneckengehausen nistenden Biene Osmia spinulosa (Kirby, 1802) (Hymenoptera, Megachilidae). Veroffentlichungen fur Naturschutz und Landschaftspflege Baden-Wurttemberg, 68 / 69, 291 - 334."]}

Hoplitis (Tkalcua) paralias (Mavromoustakis 1954) Osmia paralias Mavromoustakis 1954: 266. Type material: Holotype ♀, ���Bat Yam, 16.4.1946 ��� (Israel), Department of Agriculture, Ministry of Agriculture, Natural Resources and Environment, Nicosia. Literature records. ISRAEL AND PALESTINE: Bat Yam, 15.4.��� 5.5.1946 (Mavromoustakis, 1954). New records. MOROCCO: 20km W Boudnib, 9.4. 1995 (leg. M. Halada); 10km N Erfoud, 10.4. 1995 (leg. M. Halada); 10km N Mhamid, 21.��� 22.4.1995 (leg. M. Halada); Tagounite, 60km S Zagora, 23.4. 1995 (leg. M. Halada); 5km S Zagora, 24.��� 25.4.1995 (leg. M. Halada); Oued Draa, 34km SE Zagora, 16.3. 1997 (leg. M. Hauser). TUNISIA: 15km W Nefta, 22.3. 2001 (leg. C. Schmid-Egger). ISRAEL AND PALESTINE, Negev: 17km NNW Elat, 1.3. 1997 (leg. J.G. Rozen & M.S. Engel); 5km E Zihor Jct., 3.3. 1997 (leg. J.G. Rozen & M.S. Engel). JORDAN: Fidan, 125km N Aqaba, 6.4. 1989 (leg. J. Gusenleitner); Wadi Rum, 10.4. 1989 (leg. J. Gusenleitner); 70km NE Aqaba, 14.4. 1989 (leg. J. Gusenleitner). Distribution. From Morocco to Tunisia along the northern border of the Sahara Desert and from the Negev Desert to central Israel; the species is expected to occur also in Libya and Egypt. Pollen hosts. Polylectic. Pollen sources known so far include Genisteae (Fabaceae), Zygophyllaceae and Brassicaceae (based on 5 pollen loads from 3 different localities in Israel and Jordan). Flower record: Retama raetam (label record). Nesting biology. Unknown., Published as part of M��ller, Andreas & Mauss, Volker, 2016, Palaearctic Hoplitis bees of the subgenera Formicapis and Tkalcua (Megachilidae, Osmiini): biology, taxonomy and key to species, pp. 105-120 in Zootaxa 4127 (1) on pages 113-114, DOI: 10.11646/zootaxa.4127.1.5, http://zenodo.org/record/259694, {"references":["Mavromoustakis, G. A. (1954) New and interesting bees (Hymenoptera, Apoidea) from Israel. Bulletin of the Research Council of Israel, 4, 256 - 275."]}

Biology of Hoplitis (Formicapis) and H. (Tkalcua) Hoplitis (Formicapis) species show a close affinity to flowers of Rosaceae as pollen sources (for details see species accounts). Among the Rosaceae, herbaceous taxa such as Fragaria or Potentilla are the most important or even the exclusive hosts. While H. maritima (Romankova 1985) and possibly also H. excisa might be strictly specialized on Rosaceae, H. robusta is polylectic with a distinct preference for Potentilla (Fig. 2, Tab. 1). Both H. maritima and H. robusta nest in insect burrows in dead wood and use chewed leaves to build cell partitions and nest plug (for details see species accounts). In the latter species, small pieces of wood are sometimes incorporated into the leaf matrix of the cell partitions. Information on the biology of Hoplitis (Tkalcua) is available only for H. zandeni (for details see species account). Recent observations on Fuerteventura (Canary Islands) revealed that this species is polylectic harvesting pollen on flowers of at least six plant families (Tab. 1), but probably not on Asteraceae. Interestingly, species of H. (Platosmia), the probable sister group of H. (Tkalcua), have a strong affinity to pollen of Fabaceae and/or Resedaceae (M��ller 2015), which are also the two most important pollen host taxa exploited by H. zandeni (Tab. 1). Whether this merely reflects the flower supply at the study sites or indicates some underlying phylogenetic pattern remains unclear. H. zandeni nests in empty snail shells and constructs 1���2 brood cells per shell (Figs. 11��� 16). Cell partitions and nest plug consist of chewed leaves densely mixed with sand grains (Figs. 13���16). Interestingly, the nest plug is not built at the shell opening as in all other snail-shell nesting osmiine bee species known so far (A. M��ller unpublished) but inside the shell about 1 / 3 to 1 whorl behind the shell aperture (Fig. 13). In contrast to other osmiine bee genera such as Osmia, Protosmia or Wainia, where nesting in empty snail shells is rather widespread (Kuhlmann et al. 2011; M��ller 2016), snail-shell nesting in the genus Hoplitis has been reported so far beside H. zandeni only for the Afrotropic Hoplitis (Anthocopa) conchophila Kuhlmann and the Palaearctic Hoplitis (Hoplitis) fertoni (P��rez) (Gess & Gess 1999, 2008; Kuhlmann et al. 2011; Sedivy et al. 2013 b), indicating three independent origins of snail-shell nesting in Hoplitis., Published as part of M��ller, Andreas & Mauss, Volker, 2016, Palaearctic Hoplitis bees of the subgenera Formicapis and Tkalcua (Megachilidae, Osmiini): biology, taxonomy and key to species, pp. 105-120 in Zootaxa 4127 (1) on page 107, DOI: 10.11646/zootaxa.4127.1.5, http://zenodo.org/record/259694, {"references":["Romankova, T. G. (1985) A new subspecies of Formicapis robusta (Hym. Megachilidae) from Primorye area. Vestnik Zoologii, 6, 66 - 68 [in Russian]","Muller, A. (2015) Palaearctic Hoplitis bees of the subgenus Platosmia (Megachilidae, Osmiini): biology, taxonomy and key to species. Zootaxa, 3936 (1), 71 - 81. http: // dx. doi. org / 10.11646 / zootaxa. 3936.1.3","Kuhlmann, M., Gess, F. G., Koch, F. & Gess, S. K. (2011) Southern African osmiine bees: taxonomic notes, two new species, a key to Wainia, and biological observations (Hymenoptera: Anthophila: Megachilidae). Zootaxa, 3108, 1 - 24.","Muller, A. (2016) Palaearctic osmiine bees - systematics and biology of a fascinating group of solitary bees. ETH Zurich. Available from: http: // blogs. ethz. ch / osmiini (accessed 2 March 2016)","Gess, F. W. & Gess, S. K. (1999) The use by wasps, bees and spiders of shells of Trigonephrus Pilsb. (Mollusca: Gasteropoda: Dorcasiidae) in desertic winter-rainfall areas in southern Africa. Journal of Arid Environments, 43, 143 - 153. http: // dx. doi. org / 10.1006 / jare. 1999.0549","Gess, S. K. & Gess, F. W. (2008) Patterns of usage of snail shells for nesting by wasps (Vespidae: Masarinae and Eumeninae) and bees (Megachilidae: Megachilinae) in Southern Africa. Journal of Hymenoptera Research, 17, 86 - 109.","Sedivy, C., Dorn, S. & Muller, A (2013 b) Evolution of nesting behaviour and kleptoparasitism in a selected group of osmiine bees (Hymenoptera: Megachilidae). Biological Journal of the Linnean Society, 108, 349 - 360. http: // dx. doi. org / 10.1111 / j. 1095 - 8312.2012.02024. x"]}

Hoplitis (Formicapis) robusta (Nylander 1848) Heriades robusta Nylander 1848: 270. Type material: Lectotype ♀, by designation of Tkalcu (1995), ���Helsingfors��� (Finland), Finnish Museum of Natural History, Helsinki. Type species of Formicapis Sladen. Osmia Rhinoceros Giraud 1861: 464. Type material: Syntypes ♀♀, ���Gastein��� (Austria), Giraud Collection. Synonymy in Schletterer (1889). Heriades trinacria Morawitz 1869: 41. Type material: ��, ���im Gdowschen Kreise��� (Russia), Zoological Institute, Russian Academy of Sciences, St. Petersburg. Synonymy in Popov (1946). Formicapis clypeata Sladen 1916: 271. Type material: Syntypes ��♀, ���Aweme, Manitoba and Waterhole, Alberta��� (Canada). Synonymy in Peters (1970). New records. SWITZERLAND, Grisons: Vals, Riefa-Prad��tsch-Riedboda, 1600���2050m, 25.7. 2008 (leg. A. M��ller); Medel, Uaul da Vergera, 1600m, 18.6. 2013 (leg. R. Neumeyer); Sedrun, Uaul Bugnei, 1700m, 5.7.��� 1.8. 2013 (leg. A. M��ller); St Gallen: Pf��fers, 1420m, 7.7.��� 31.7.2013 (leg. H. Martz); Valais: Chandolin/Siders, 2200m, 18.7. 1989 (leg. Perraudin); Visperterminen, 2000m, 12.8. 2005 (leg. A. M��ller); Montana, 15.7. 2006 (leg. K. Hirt). AUSTRIA, Carinthia: Oberschachnern SE Heiligenblut, 22.8. 1985 (leg. J. Gusenleitner); Ochsenriegel NW Weinebene, Koralpe, 1460m, 25.8. 1989 (leg. J. Gusenleitner); Tirol: Zedlach W Matrei, 20.8. 1985 (leg. J. Gusenleitner). ITALIA, South Tirol: Wolkenstein, 2.7.1921; Seiser Alp, 2.��� 14.7.1971 (leg. H. Wolf); Schnals/ Kurzras, Meran, 2100m, 7.��� 10.8.1974 (leg. H. Wolf); Pfossental, 1500���2000m, 27.8. 1984 (leg. N. Mohr). MONGOLIA: Khangaur Mountains 5km N Khunt, 20.7. 2005 (leg. J. Halada). Distribution. Boreal zone across Europe (southern half of Finland, Estonia, Latvia, Belarus, European part of Russia), Asia (Asian part of Russia, northern Mongolia, northeasternmost China) and North America (Alaska, Canada) (Gr��nwaldt 1939; Elfving 1968; Peters 1970; Romankova 1985; Tkalcu 1995; Prishchepchik 2000; Wu 2006; Ascher & Pickering 2015). In addition, H. robusta is distributed in the Alps (Italy, Switzerland, Austria) between 1200m and the timberline (Schwarz et al. 1996; Ebmer 1997, 2001; Amiet et al. 2004; Kopf 2008) and in mountainous regions of the western USA southwards to California and Colorado (Hurd & Michener 1955). According to Rasmont et al. (1985), H. robusta also occurs in Belgium and Luxembourg, which is certainly erroneous, as well as in France, which is not beyond doubt as no reliable record exists for this country (Benoist 1931; M. Aubert & G. LeGoff personal communication). Pollen hosts. Polylectic with a strong preference for Potentilla (Rosaceae), e.g. P. erecta (Tab. 1). Females collect pollen on flowers of P. erecta by rapidly seesawing their metasomal scopa against the anthers, which they embrace below the mesosoma, while turning their body axis around the flower centre (Fig. 2); simultaneously, they suck nectar from the nectaries at the base of the androecium. The importance of Potentilla as pollen source is apparent by the finding that its pollen constituted 68.1 % of the total pollen grain volume and that 18 out of 22 pollen loads analyzed contained Potentilla pollen, 13 of which were pure loads (Tab. 1). Additional pollen sources were Helianthemum (Cistaceae), Medicago (Fabaceae), Rubus (Rosaceae), Cichorioideae (Asteraceae) and Ranunculus (Ranunculaceae). The fact that the two pollen loads originating from the USA were composed of pollen of the Potentilla type and of Helianthemum, respectively, suggests that North American and European populations of H. robusta exhibit similar host plant preferences. One brood cell provision from Wyoming analysed by Clement & Rust (1975), however, was composed of putative Fabaceae pollen, suggesting that flowers of Fabaceae might locally be important pollen hosts. Flower records: Taraxacum (Hurd and Michener 1955; Elfving 1968); Anthriscus sylvestris, Geranium palustre, G. sylvaticum, Geum rivale, Leontodon autumnalis, Potentilla anserina, Satureja alpina, Sedum acre, S. aizoon, S. camtschaticum, S. cepaea, S. spurium, Silene viscaria (Elfving 1968); Silene rupestris (Peters 1970); Lotus corniculatus, Potentilla aureola, P. pulcherrima (label records). Nesting biology. Nesting sites are insect burrows and drilled borings (diameter 4���5mm) in dead wood containing 3���14 linearly arranged brood cells (Clement & Rust 1975; Frey-Gessner 1880). Some nests contain empty vestibule cells following the last brood cell. Cell partitions are made of chewed leaves, sometimes with small pieces of wood embedded in the leaf matrix. The nest plug varies in thickness from 10mm to 14mm and consists of 8���13 layers of leaf pulp (Clement & Rust 1975). In one case, a female collected leaf pulp on leaves of Potentilla erecta (A. M��ller unpublished)., Published as part of M��ller, Andreas & Mauss, Volker, 2016, Palaearctic Hoplitis bees of the subgenera Formicapis and Tkalcua (Megachilidae, Osmiini): biology, taxonomy and key to species, pp. 105-120 in Zootaxa 4127 (1) on pages 110-111, DOI: 10.11646/zootaxa.4127.1.5, http://zenodo.org/record/259694, {"references":["Nylander, W. (1848) Adnotationes in expositionem monographicam apum borealium. Notiser ur Sallskapets pro Fauna et Flora Fennica Forhandlingar, 1, 165 - 282.","Tkalcu, B. (1995) Die Bienen der Tribus Osmiini der Mongolei (Insecta: Hymenoptera: Apoidea: Megachilidae). Entomologische Abhandlungen, 57, 109 - 147.","Giraud, J. (1861) Fragments entomologiques. Verhandlungen der Zoologischen und Botanischen Gesellschaft in Wien, 11, 447 - 470.","Schletterer, A. (1889) Monographie der Bienen-Gattungen Chelostoma Latr. und Heriades Spin. Zoologisches Jahrbuch fur Systematik, 4, 591 - 691.","Morawitz, F. (1869) Die Bienen des Gouvernements von St. Petersburg. Horae Societatis Entomologicae Rossicae, 6, 27 - 71.","Popov, V. B. (1946) Notes on the nomenclature of the bees (Hymenoptera, Apoidea). Proceedings of the Royal Entomological Society of London, B, 15, 106 - 109. http: // dx. doi. org / 10.1111 / j. 1365 - 3113.1946. tb 00834. x","Sladen, F. W. L. (1916) Bees of Canada - Fam. Megachilidae. The Canadian Entomologist, 48, 269 - 272. http: // dx. doi. org / 10.4039 / Ent 48269 - 8","Peters, D. S. (1970) Die Synonymie von Osmia robusta (Nylander 1848) und Osmia forsii Alfken 1924. Senckenbergiana biologica, 51, 193 - 197.","Grunwaldt, W. (1939) Zur Apiden-Fauna Lettlands II. Die Unterfamilie Megachilinae. Korrespondenzblatt des Naturforscher- Vereins zu Riga, 63, 87 - 96.","Elfving, R. (1968) Die Bienen Finnlands. Fauna Fennica, 21 - 69.","Romankova, T. G. (1985) A new subspecies of Formicapis robusta (Hym. Megachilidae) from Primorye area. Vestnik Zoologii, 6, 66 - 68 [in Russian]","Prishchepchik, O. V. (2000) On the fauna of megachilid bees (Hymenoptera, Apoidea, Megachilidae) of Byelarus. Vestnik Vitebsk University, 2000, 88 - 93 [in Russian with English summary]","Wu, Y. (2006) Hymenoptera: Megachilidae. Fauna Sinica, Insecta, 44, 1 - 474 [in Chinese with English summary]","Ascher, J. S. & Pickering, J. (2015) Discover Life bee species guide and world checklist (Hymenoptera: Apoidea: Anthophila). Avaliable from: http: // www. discoverlife. org / mp / 20 q? guide = Apoidea _ species (accessed 30 May 2015)","Schwarz, M., Gusenleitner, F., Westrich, P. & Dathe, H. H. (1996) Katalog der Bienen Osterreichs, Deutschlands und der Schweiz (Hymenoptera, Apidae). Entomofauna (Ansfelden), Supplement, 8, 1 - 398.","Ebmer, A. W. (1997) Hymenopterologische Notizen aus Osterreich - 7 (Insecta: Hymenoptera: Apoidea). Linzer biologische Beitrage, 29, 45 - 62.","Ebmer, A. W. (2001) Hymenopterologische Notizen aus Osterreich - 14 (Insecta: Hymenoptera: Apoidea). Linzer biologische Beitrage, 33, 435 - 460","Amiet, F., Herrmann, M., Muller, A. & Neumeyer, R. (2004) Apidae 4: Anthidium, Chelostoma, Coelioxys, Dioxys, Heriades, Lithurgus, Megachile, Osmia, Stelis. Fauna Helvetica, 9, 1 - 274.","Kopf, T. (2008) Die Bienenfauna (Hymenoptera: Apidae) des Schlerngebietes (Sudtirol, Italien) mit Angaben zu den Artengemeinschaften ausgewahlter Lebensraume. Gredleriana, 8, 429 - 466.","Hurd, P. D. & Michener, C. D. (1955) The megachiline bees of California. Bulletin of the California Insect Survey, 3, 1 - 247.","Benoist, R. (1931) Les osmies de la faune francaise (Hymenopt. Apidae). Annales de la Societe Entomologique de France, 100, 23 - 60.","Clement, S. L. & Rust, R. W. (1975) The biology of Hoplitis robusta (Hymenoptera: Megachilidae). Entomological News, 86, 115 - 120.","Frey-Gessner, E. (1880) Exkursionen im Sommer 1879. Mittheilungen der Schweizerischen Entomologischen Gesellschaft, 5, 515 - 589."]}

Key to the species of Hoplitis (Tkalcua) The female of Hoplitis emiratensis is still unknown. Females 1 Clypeus apically with deep median incision of semicircular shape, from which a narrow and shallow longitudinal groove extends towards its base (Fig. 19). Antennal segments 4���12 yellowish. Mandible three-toothed (Fig. 19). Apex of inner spur of hind leg curved almost at right angles. Stigma yellowish. Terga 1���5 of unworn specimens with uninterrupted white hair bands. Body length 5���6mm. Morocco to Tunisia, Israel................................................ Hoplitis paralias - Clypeus apically without deep median incision but with weak median impression at its declivous apical third (Fig. 20). Antenna entirely black. Mandible four-toothed (Fig. 20). Apex of inner spur of hind leg only slightly curved. Stigma blackish. Terga 1���5 with broadly interrupted white hair bands. Body length 5���6mm. Canary Islands............... Hoplitis zandeni Males 1 Second segment of labial palpus distinctly less than 1.5 x as long as first segment (Fig. 22). Brightened parts of mandible yellow. Appressed white pilosity on lateral sides of tergum 1 almost reaching base of horizontal part of tergum 1 (Fig. 23). Carina separating anterior concavity of tergum 1 from its horizontal part with row of strong white hairs (Fig. 23). Disc of tergum 5 densely covered with appressed white pilosity. Head less than 0.9 x as long as wide. Body length 4���5mm. Arabian Peninsula..................................................................................... Hoplitis emiratensis - Second segment of labial palpus 1.75��� 2 x as long as first segment. Brightened parts of mandible reddish-brown or mandible entirely black. Appressed white pilosity on lateral sides of tergum 1 less extended and restricted to posterior half of horizontal part of tergum 1. Carina separating anterior concavity of tergum 1 from its horizontal part without row of strong white hairs. Disc of tergum 5 without dense appressed white pilosity. Head 0.9 x as long as wide or longer......................... 2 2 Antennal segments 6���12 about 1.25 x as long as wide. Antennal segments 4���13 yellowish. Stigma yellowish. Second segment of labial palpus about 2 x as long as first segment. Terga 1���5 of unworn specimens with uninterrupted white hair bands. Apical margins of sterna 1���2 with rather dense white hair bands. Gonoforceps of about equal width along its entire length, its apical half yellowish-brown. Body length 5���5.5mm. Morocco to Tunisia, Israel............................ Hoplitis paralias - Antennal segments 6���12 about 1.5 x as long as wide. Antenna entirely black. Stigma blackish except for its centre, which is light brown. Second segment of labial palpus about 1.75 x as long as first segment. Terga 1���5 with white hair bands, which are broadly interrupted on terga 1���4. Apical margins of sterna 1���4 sparsely ciliated with long white hairs. Apical half of gonoforceps broadened and blackish (Fig. 21). Body length 5���6mm. Canary Islands......................... Hoplitis zandeni, Published as part of M��ller, Andreas & Mauss, Volker, 2016, Palaearctic Hoplitis bees of the subgenera Formicapis and Tkalcua (Megachilidae, Osmiini): biology, taxonomy and key to species, pp. 105-120 in Zootaxa 4127 (1) on pages 117-118, DOI: 10.11646/zootaxa.4127.1.5, http://zenodo.org/record/259694

Hoplitis (Formicapis) maritima (Romankova 1985) Formicapis robusta maritima Romankova 1985: 66. Type material: Holotype ♀, ���Primorski territory, Kievka, 22.5.��� 6.7.1979 ��� (Russia), Zoological Institute, Russian Academy of Sciences, St. Petersburg. New records. MONGOLIA: 90km N Ulanbatar, Segnezer river, 1450m, 6.��� 8.7.2003 (leg. J. Halada). RUSSIA, Primorski krai, Lazovski Zapovednik: Cordon Amerika, river Perekatnaya, 43 ��02��� 32 ������N 134 ��02��� 32 ������E, 329m, 19.5. 2001 (leg. M. Quest); Ta Tshingousa, 43 ��00��� 32 ������N 134 ��07��� 34 ������E, 14.6. 2001 (leg. M. Quest). Distribution. From northern Mongolia to Primorski krai in southernmost Russian Far East. Pollen hosts. Oligolectic on Rosaceae (e.g. Fragaria, Potentilla) (Romankova 1985; Proshchalykin in Quest 2009). Flower records: Fragaria orientalis, Potentilla amurensis, P. fragarioides (Romankova 1985; label records). Nesting biology. Nests were found in cerambycid beetle burrows (width 3.5���6.5mm, height 2.7���3.5mm) in 4��� 5cm thick branches of conifers lying on the ground containing 2���6 linearly arranged brood cells (Romankova 1985). Cell partitions and nest plug are made of chewed leaves. The females clean the burrows from wood debris with their mandibles., Published as part of M��ller, Andreas & Mauss, Volker, 2016, Palaearctic Hoplitis bees of the subgenera Formicapis and Tkalcua (Megachilidae, Osmiini): biology, taxonomy and key to species, pp. 105-120 in Zootaxa 4127 (1) on page 110, DOI: 10.11646/zootaxa.4127.1.5, http://zenodo.org/record/259694, {"references":["Romankova, T. G. (1985) A new subspecies of Formicapis robusta (Hym. Megachilidae) from Primorye area. Vestnik Zoologii, 6, 66 - 68 [in Russian]","Quest, M. (2009) Artbestand, Okologie und Habitatwahl von Bienen ausgewahlter Offenlebensraume im Lazovski Zapovednik (Ferner Osten Russland). Entomofauna, Supplement 15, 1 - 357."]}

Mauss, Volker, Mauss, Henrik (2016): Field observations of the behaviour of the pollen wasp Quartinia tenerifina RICHARDS, 1969 (Hymenoptera, Vespidae, Masarinae) at flowers in Tenerife (Canary Islands, Spain). Linzer biologische Beiträge 48 (2): 1327-1332, DOI: http://doi.org/10.5281/zenodo.5355480, {"references":["GESS S.K. & F.W. GESS (2010): Pollen wasps and flowers in southern Africa. - SANBI Biodiversity Series 18: 1-147. South African National Biodiversity Institute, Pretoria.","GOBIERNO DE CANARIAS (2004): Reserva Natural Especial del Malpais de Guimar, Plan Director. - http://www.todotenerife.es/assets/downloads/e6e94e0f78.pdf 108 pp. (called up 2016/08/29).","GUSENLEITNER J. (1990): Kurze Mitteilung uber Faltenwespen der Insel Lanzarote (Canarische Inseln) (Hymenoptera, Vespoidea, Eumenidae und Masaridae). - Linzer biologische Beitrage 22 (2): 561-564.","HOHENESTER A. & W. WELSS (1993): Exkursionsflora fur die Kanarischen Inseln: mit Ausblicken auf ganz Makronesien. - E. Ulmer, Stuttgart, 374 pp.","HOHMANN H., LA ROCHE F., ORTEGA G. & J. BARQUIN (1993): Bienen, Wespen und Ameisen der Kanarischen Inseln (Insecta: Hymenoptera: Aculeata). - Veroffentlichungen aus dem Ubersee-Museum Bremen Naturwissenschaften (vormals Reihe A) 12: 14-712.","MAUSS V. (2007): Evolution verschiedener Lebensformtypen innerhalb basaler Teilgruppen der Faltenwespen (Hymenoptera, Vespidae). - Denisia 20: 701-722.","MAUSS V. & A. MULLER (2016): Contribution to the bionomics of the pollen wasp Quartinia canariensis BLUTHGEN, 1958 (Hymenoptera, Vespidae, Masarinae) in Fuerteventura (Canary Islands, Spain). - Journal of Hymenoptera Research 50: 1-24. doi: 10.3897/JHR.50.6870","MULLER A. & M. KUHLMANN (2008): Pollen hosts of western Palaearctic bees of the genus Colletes (Colletidae) - the Asteraceae paradox. - Biological Journal of the Linnean Society 95: 719-733.","RICHARDS O.W. (1969): Records of Masarid wasps with descriptions of two new species of Quartinia Ed. ANDRE (Hymenoptera). - Journal of Natural History 3: 79-83."]}