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Supplement 18, 144 Seiten ISSN 0250-4413 / ISBN 978-3-925064-71-8 Ansfelden,24.März 2015 The Cuckoo Bees of the Genus Panzer, 1806 in Europe, North Africa and the Middle East A Review and Identification Guide Table of Contents Introduction ............................................................................................................ 4 General Part ............................................................................................................ 7 – Description of the Genus ........................................................................... 7 – Number of Species Described ................................................................... 7 – Species Diversity on Country Level .......................................................... 8 – Abundance of Stelis ................................................................................... 9 – Host Associations ...................................................................................... 9 – Flower Preferences ................................................................................... 15 – Flight Season ............................................................................................ 19 – Sexual Dimorphism .................................................................................. 19 – Geographic Variation ............................................................................... 19 – Taxonomy: The Subgenera of Stelis ........................................................ 20 Coverage and Methodology ................................................................................. 23 – Geographic and Taxonomic Coverage ..................................................... 23 – Terminology and Abbreviations ............................................................... 23 – Depositories ............................................................................................. 24 – Maps ......................................................................................................... 24 – Photographs .............................................................................................. 25 – Acknowledgements .................................................................................. 25 Key to the West Palaearctic and Middle Eastern Species of Stelis ................... 26 – Females .................................................................................................... 26 – Males ........................................................................................................ 32 Species Accounts ................................................................................................... 39 References ........................................................................................................... 135 3 Introduction Stelis bees are relatively small-sized bees known as cuckoo bees: they lay their eggs in the nests of other bees, reminiscent of the behaviour of cuckoo birds. Their para- sitic larva destroys the host’s egg or kills the young larvae of its host. The scientific name Stelis already points to this parasitic character: Stelis is the Greek word for “mistletoe”, a parasitic plant. As the Stelis larva consumes the stored food of its host, this form of parasitism is called “cleptoparasitism”. Hosts of Stelis are exclusively members of the Megachilinae, i.e. the same subfamily of bees to which the parasite itself belongs. The appearance of some species of Stelis is so similar to Megachil- inae, especially to the members of the genus Anthidium (s.l.), that they are some- times difficult to distinguish; some Stelis species have originally been described in the genus Anthidium. Stelis species are of equal size or smaller than their hosts. Females can easily be distinguished from other Anthidiini bees as they lack a pollen-collecting structures (the scopa) on the underside of the abdomen. Being parases, Stelis females do not construct their own nests. The genus name Stelis also refers to a genus of plants: Stelis Olof Peter Swartz, 1799 is the name of a genus of leach orchids, which comprises approximately 500 species. These mainly epiphytic plants are found predominantly in tropical North and Central America. In a few cases, the usage of the name Stelis for both an animal and a plant genus has caused some confusion. There is even a plant species and a bee species which bear the same name: Stelis gigantea Friese, 1921 is a Middle Eastern bee species, while Stelis gigantea Pridgeon & M. W. Chase, 2002 is an orchid, which occurs in the tropical Central America. The genus Stelis comprises approximately 105 species worldwide, with about 20 to 25% of them occurring in the Western Palaearctic and the Middle East. In one of the first reviews, Nylander (1852) recognised four species of Stelis for Central Europe. The first comprehensive account of Stelis has been given by Friese (1895) within his synopsis of the bees of Europe. His work is a compilation of the hitherto known information and includes the reproduction of the original species descriptions in the original language (mostly Latin, but also French and German). Friese’s work comprises 14 species of Stelis, and is still a very useful source today. Almost at the same time, de Dalla Torre (1896) published his world list of Hyme- noptera. Although this work did not include descriptions or similar information, the list of synonyms and literature references constituted an important compilation which helps to shed light on the sometimes confusing diversity of names. The worldwide list of de Dalla Torre covered 35 species of Stelis. Schmiedeknecht (1907) , in his work “Die Hymenopteren Mitteleuropas”, covered nine Central Euro- pean species of Stelis and provided an identification key for these species. 4 The genus Stelis was intensively studied by the Russian zoologist Vladimir B. Popov (1933, 1935, 1939, 1941, 1944, 1956). While none of the West Palaearctic Stelis taxa described by him is still regarded as valid, one of Popov’s main achieve- ments clearly was to introduce the morphology of male genitalia for species identifi- cation. For most species, his drawings are still the only ones available. Also the German entomologist Johann Dietrich Alfken (1862–1945) needs to be mentioned here among those who contributed much to our knowledge of Palaearctic Stelis bees. He published over 200 works on bees, several of which included infor- mation on Stelis. The Polish entomologist Jan Noskiewicz (1890–1963) described several new taxa, three of which are included here as valid species: S. hungarica, S. iugae, and S. odontopyga. Some of Noskiewicz’s illustrations were used in this identification guide. The Greek entomologist Georgios A. Mavromoustakis (b. 1901) made significant contributions to the studies of the bee fauna, and in a series of publications which appeared between 1939 and 1968, he dealt with the bees of Cyprus and Greece, as well as Israel, Lebanon, Syria and other Mediterranean coun- tries. Mavromoustakis is the author of two species, Stelis pentelica and S. rhodia. The Belgian entomologist Jean J. Pasteels, whose work was mainly focused on Afri- can bees, made in two works, both published in 1969, significant contributions mainly to the higher taxonomy of Anthidiini. The subgeneric classification of the genus Stelis has been treated by Charles D. Michener and Terry L. Griswold (1994), and this system has principally been ap- plied by Michener in his epochal work “The Bees of the World” (2000, 2007). Klaus Warncke (1937–1993) was the first after quite some time who intensively researched the genus Stelis again. His work resulted in a comprehensive list of the West Palaearctic species and was based on rich new, own material mainly from Turkey and other Middle East countries, as well as extensive museum studies (Warncke 1992). He described two new species, Stelis maroccana and S. orientalis, and synonymized several names. His descriptions and explanations are usually very brief and not accompanied by drawings; some of his conclusions still need to be fully verified. Klaus Warncke’s identification key to the West Palaearctic species is still the most comprehensive account and was used as the principal basis for this work. While quite a lot of new information has become known since Warncke’s (1992) account, his work still remains the most comprehensive work on Palaearctic Stelis species. More recent publications include those of Donald B. Baker, who published in 1999 a “Provisional list of species-group names in old-world Stelis” and described two new species; Jérôme G. Rozen and Soliman M. Kamel (2009), who published a review of Stelis biology; Maximilian Schwarz and Fritz Gusenleitner (2010), who described Stelis ortizi and re-examined the status of S. hispanica; and Concepción 5 Ornosa, F. Torres and F. Javier Ortiz Sánchez (2009), who provided a key in Span- ish to the 10 species occurring on the Iberian Peninsula. Many others contributed to our knowledge mainly in the frame of faunistic work – too many to be listed here individually. Our understanding of this group of bees is now much better than a few decades ago, but a comprehensive review of this information was so far not availa- ble. The only comprehensive identification keys for Stelis bees are, therefore, those by Friese (1895), Schmiedeknecht (1907) and Warncke (1992). None of them have illustrations; and Warncke’s key is very brief and concise, which makes species identification often a difficult task especially for

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