Journal of the Saudi Society of Agricultural Sciences (2013) 12, 85–89

King Saud University Journal of the Saudi Society of Agricultural Sciences

www.ksu.edu.sa www.sciencedirect.com

FULL LENGTH ARTICLE A preliminary list of bee genera in the Kingdom of Saudi Arabia (Hymenoptera: Apoidea)

Michael S. Engel a,*, Abdulaziz S. Alqarni b, Mohammed A. Hannan b a Division of Entomology, Natural History Museum, and Department of Ecology & Evolutionary Biology, University of Kansas, 1501 Crestline Drive – Suite 140, Lawrence, KS 66045, USA b Department of Plant Protection, College of Food and Agriculture Sciences, King Saud University, PO Box 2460, Riyadh 11451, Kingdom of Saudi Arabia

Received 13 April 2012; accepted 25 August 2012 Available online 31 August 2012

KEYWORDS Abstract A brief overview of those bee genera and subgenera occurring in the Kingdom of Saudi Biodiversity; Arabia is provided. In total 22 genera are currently documented in the literature for the country, Apoidea; although a further 17 genera undoubtedly occur here based on extralimital records. The lists pro- Anthophila; vided here will serve as a baseline for biodiversity studies on the bee fauna of the country that are Taxonomy; presently underway. The anthidiine genus Icteranthidium Michener (Megachilidae) and the dasy- Saudi Arabia; podaine genus Promelitta Warncke (Melittidae) are newly recorded for the country. Pollination ª 2012 King Saud University. Production and hosting by Elsevier B.V. All rights reserved.

1. Introduction (e.g., Al-Ajlan, 2004), the melittological fauna remains taxo- nomically and biologically little known, and thereby the full Bees are the most important pollinators in natural and agricul- potential of the native species is not understood and underex- tural ecosystems worldwide. With a modern diversity of around ploited. Herein we attempt to provide a list of the currently 20,000 species they are also fabulously diverse. This remarkable recorded genera of bees known to occur natively in the Kingdom diversity is presently classified in six or seven families, depending of Saudi Arabia. It is hoped that this might represent an initial on the status of the Australian Stenotritinae (Engel, 2005, 2011a; baseline of biodiversity data for the melittofauna of the country, Michener, 2007). Given their importance it is not surprising that and stimulate new taxonomic and basic biological research. For the bee faunas of various regions have been investigated in terms each of these, material has been visually confirmed and identi- of the species diversity and pollination roles of sundry taxa. fied by the authors as part of on-going survey efforts. Unfortunately, while the pollination role of non-Apis bees has been given some attention in the Kingdom of Saudi Arabia 2. Preliminary lists of Saudi bee genera

Since 1970 relatively few revisionary or taxonomic works have * Corresponding author. Tel.: +1 785 864 2319. E-mail address: [email protected] (M.S. Engel). considered elements of the Saudi Arabian bee fauna. Notable Peer review under responsibility of King Saud University. examples of taxonomic works including material from Saudi Arabia since 1970 are those by Daly (1983), Ebmer (1984, 1985), Engel (2008, 2009a), Kuhlmann (2002, 2003), Moustafa (1986), Pasteels (1979, 1980, 1981), Patiny (2002), Pesenko and Production and hosting by Elsevier Pauly (2009), Tkaclu˚(1992), and van der Zanden (1987). Most 1658-077X ª 2012 King Saud University. Production and hosting by Elsevier B.V. All rights reserved. http://dx.doi.org/10.1016/j.jssas.2012.08.001 86 M.S. Engel et al.

Table 1 Hierarchical outline of short-tongued bee genera and subgenera presently recorded from Saudi Arabia. The higher classification follows those of Engel (2005) and Michener (2007). Family Colletidae Lepeletier Genus Systropha Illiger Subfamily Colletinae Lepeletier Subfamily Nomiinae Robertson Genus Colletes Latreille Genus Nomia Latreille Subfamily Hylaeinae Viereck Subgenus Crocisaspidia Ashmead Genus Hylaeus Fabricius Genus Pseudapis Kirby Subgenus Paraprosopis Popov Subgenus Pseudapis s.str. Family Andrenidae Latreille Subfamily Halictinae Thomson Subfamily Andreninae Latreille Tribe Nomioidini Bo¨rner Genus Andrena Fabricius Genus Ceylalictus Strand Subgenus Chrysandrena Hedicke Subgenus Ceylalictus s.str. Subgenus Graecandrena Warncke Genus Nomioides Schenck Subgenus Melanapis Cameron Subgenus Nomioides s.str. Subgenus Suandrena Warncke Tribe Halictini Thomson Subfamily Panurginae Leach Genus Halictus Latreille Tribe Melitturgini Newman Subgenus Seladonia Robertson Genus Borgatomelissa Patiny Subgenus Vestitohalictus Blu¨thgen Tribe Meliturgulini Engel Genus Lasioglossum Curtis Genus Meliturgula Friese Subgenus Afrodialictus Pauly Tribe Panurgini Leach Subgenus Ctenonomia Cameron Genus Panurgus Panzer Subgenus Evylaeus Robertson Subgenus Panurgus s.str. Subgenus Lasioglossum s.str. Family Melittidae Schenck Family Halictidae Thomson Subfamily Dasypodainae Bo¨rner Subfamily Rophitinae Schenck Tribe Promelittini Michener Tribe Rophitini Schenck Genus Promelitta Warncke Genus Dufourea Lepeletier Tribe Dasypodaini Bo¨rner Subgenus Dufourea s.str. Genus Dasypoda Latreille

recently, Alqarni et al. (2012a) recorded the genus Tetraloniella 3. Extralimital taxa Ashmead (Eucerini) for the country based on a species then unknown to Science, while Engel et al. (2012) provided records Given that surveys of the bee fauna of Saudi Arabia are cur- for the genus Systropha Illiger (Rophitinae). Table 1 summa- rently underway it will be interesting to see how the faunal list rizes the currently known genera and subgenera of short- continues to grow, and eventually sufficient material should tongued bees from the Kingdom of Saudi Arabia, while perhaps an enumeration of the species for the country. How- Table 2 gives those genera of the long-tongued bee families ever, it is clear from studies elsewhere in the Arabian Peninsula Megachilidae and Apidae. In total this encompasses 22 genera that various bee genera not presently recorded from Saudi (two newly recorded herein), including four genera of clepto- Arabia undoubtedly occur within the country. For example, parasitic bees. By comparison, the United Arab Emirates has among the family Colletidae the widespread and distinctive at least 47 genera recorded (Dathe, 2009: noting that Chalicod- genus Hylaeus Fabricius (Hylaeinae) is one of the most readily oma Lepeletier de Saint Fargeau is herein considered at generic encountered genera of the Colletidae, with various species in rank), and all of these are assuredly shared with Saudi Arabia the region (e.g., Dathe, 1995, 2009, 2010), and yet only one (vide infra). species has been formally recorded from Saudi Arabia, Hyla- Fortunately, new efforts toward documenting the melitto- eus (Paraprosopis) albonotatus (Walker) (Dathe, 1995). Given logical fauna of the Arabian Peninsula are bringing sufficient the close proximity of numerous other species and subgenera material to the attention of researchers (e.g., Alqarni et al., of Hylaeus, the records for this group will grow rapidly with 2012b; Baker, 2002; Dathe, 2009; Engel, 2004, 2009b, 2011b; sufficient sampling. Engel and Schwarz, 2011; Hannan et al., 2012; Kuhlmann, The tribe Allodapini is another lineage which undoubtedly 2004). While most of the aforementioned did not include mate- occurs in Saudi Arabia. A species of Braunsapis Michener has rial from Saudi Arabia, they do highlight the increased focus been described from the United Arab Emirates (Engel and on the wild bee fauna of the region. Dathe, 2011), and the genus Compsomelissa Alfken is well doc- From this baseline we will be conducting sampling through- umented in the mountains of Yemen (Michener, 1966; Engel, out Saudi Arabia to ultimately update the list, eventually pro- 2011b). Given that these same mountains extend into south- ducing a guide to the bee genera and subgenera of the western Saudi Arabia many elements of the fauna are assu- Kingdom, if not the entire Arabian Peninsula. In addition, it redly shared. Indeed, this suggests that the enigmatic and is hoped that with a few years of intensive work we might be remarkable record of a putative meganomiine from the moun- able to produce the first truly authoritative list of the species tains of Yemen (Michener, 1981) indicates the presence of this for Saudi Arabia. The brief lists of bees in Al-Ahmadi and otherwise southern African lineage in the Arabian Peninsula, Salem (1999) include various misidentifications and should including southwestern Saudi Arabia. Another melittid genus be used with caution. Moreover, their account overlooks large very likely occurring in Saudi Arabia is Eremaphanta Popov numbers of species definitely recorded from Saudi Arabia. (Dathe, 2009). A preliminary list of bee genera in the Kingdom of Saudi Arabia (Hymenoptera: Apoidea) 87

Table 2 Hierarchical outline of long-tongued bee genera and subgenera presently recorded from Saudi Arabia. The higher classification follows those of Engel (2005) and Michener (2007). Cleptoparasitic taxa are noted by an asterisk (\). Family Megachilidae Latreille Tribe Ceratinini Latreille Subfamily Megachilinae Latreille Genus Ceratina Latreille Tribe Anthidiini Ashmead Subgenus Dalyatina Terzo & al. Genus Afranthidium Michener Subgenus Euceratina Hirashima & al. Subgenus Mesanthidium Popov Subgenus Pithitis Klug Genus Anthidium Fabricius Subfamily Nomadinae Latreille* Subgenus Anthidium s.str. Tribe Ammobatini Handlirsch* Subgenus Proanthidium Friese Genus Ammobates Latreille* Genus Icteranthidium Michener Subgenus Ammobates s.str.* Genus Pachyanthidium Friese Subfamily Apinae Latreille Subgenus Trichanthidiodes Michener & Tribe Eucerini Latreille Griswold Genus Eucera Scopoli Genus Pseudoanthidium Friese Subgenus Synhalonia Patton Subgenus Pseudoanthidium s.str. Genus Tetraloniella Ashmead Tribe Osmiini Newman Subgenus Tetraloniella s.str. Genus Osmia Panzer Tribe Anthophorini Dahlbom Subgenus Helicosmia Thomson Genus Amegilla Friese Genus Pseudoheriades Peters Subgenus Amegilla s.str. Tribe Megachilini Latreille Subgenus Micramegilla Brooks Genus Chalicodoma Lepeletier Subgenus Zebramegilla Brooks Subgenus Chalicodoma s.str. Genus Anthophora Latreille Subgenus Pseudomegachile Friese Subgenus Heliophila Klug Genus Coelioxys Latreille* Subgenus Paramegilla Friese Subgenus Allocoelioxys Tkalcu˚* Subgenus Petalosternon Brooks Genus Megachile Latreille Subgenus Pyganthophora Brooks Subgenus Creightonella Cockerell Tribe Melectini Westwood* Subgenus Eutricharaea Thomsona Genus Melecta Latreille* Family Apidae Latreille Subgenus Paracrocisa Alfken* Subfamily Xylocopinae Latreille Genus Thyreus Panzer* Tribe Xylocopini Latreille Tribe Apini Latreille Genus Xylocopa Latreille Genus Apis Linnaeus Subgenus Koptortosoma Gribodo Subgenus Micrapis Ashmead Subgenus Ctenoxylocopa Michener Subgenus Apis s.str. a Includes subgenus Eurymella Pasteels.

Among additional cleptoparasitic lineages expected for genera should be present in Saudi Arabia (not including the Saudi Arabia are species of the cosmopolitan genus Sphecodes aforementioned Stelis): Hoplitis Klug (subgenera Alcidamea Latreille (Halictinae), the minute species of the nomioidine- Cresson, Annosmia Warncke, Microhoplitis Tkalcu˚, Pentaden- attacking genus Chiasmognathus Engel (Ammobatini), and tosmia Warncke), Stenosmia Michener, Heriades Spinola (sub- species of the parasitic genera Stelis Panzer (Megachilinae), genus Michenerella Krombein), Haetosmia Popov, Protosmia Nomada Scopoli (Nomadini), Parammobatodes Popov Ducke (subgenus Nanosmia Griswold), Eoanthidium Popov (Ammobatini), and Epeolus Latreille (Epeolini). Suitable hosts s.str., and the subgenera Exanthidium Pasteels and Royanthidi- for all of these genera are already documented as occurring in um Pasteels (both of Pseudoanthidium Friese). Saudi Arabia and records of each have been made from the The important message from the existing Saudi Arabian re- neighboring Arabian countries such as the United Arab Emir- cords when taken into the context of extralimital records is ates (Engel, 2009b; Dathe, 2009; Schwarz, 2010). Clearly these that the fauna of Saudi Arabia is greatly under sampled. Given genera are present in Saudi Arabia and it is only a matter of that the country represents the largest portion of the Arabian time before they are caught and recorded. Peninsula and the greatest diversity of habitats, the potential A rather large number of halictid genera are also expected total bee fauna is significant, including the possibility for the for Saudi Arabia. Aside from the aforementioned Sphecodes, discovery of many new species (e.g., Alqarni et al., 2012a, the most likely candidates are Lipotriches Gersta¨cker (Nomii- 2012b). nae) and Rophites Spinola (Rophitinae), along with the follow- ing subgenera: Halictus Latreille s.str., Leuconomia Pauly (of 4. New records Nomia Latreille), Dentirophites Warncke and Cyprirophites Warncke (of Dufourea Lepeletier de Saint Fargeau), and Meg- Herein we newly record the genera Icteranthidium Michener anomioides Pesenko (of Ceylalictus Strand). (Megachilidae) and Promelitta Warncke (Melittidae) for the Perhaps the most new generic records expected are those of Kingdom of Saudi Arabia. Icteranthidium is a genus of distinc- the Megachilinae. Indeed, based on samples from areas such tive anthidiine bees common in neighboring countries and it is as the United Arab Emirates (Engel, 2004; Dathe, 2009), Yemen therefore not surprising to find it in abundance in central Saudi (Pasteels, 1980), and Oman (Pasteels, 1980) at least the following Arabia. The record is as follows: 88 M.S. Engel et al.

Icteranthidium cinctum (Klug): 2$$,1#, labeled as, ‘‘Saudi Baker, D.B., 2002. On Palaearctic and Oriental species of the genera Arabia, Riyadh, Al Amariah, [Mazra’ah] Majra Al-Gasim Pseudapis W.F. Kirby, 1900, and Nomiapis Cockerell, 1919 [farm], 6-vi-2011 [6 June, 2011], Hannan’’. Material is (Hymenoptera, Halictidae, Nomiinae). Beitr. Entomol. 52 (1), 1–83. deposited in the King Saud University Museum of Arthro- Daly, H.V., 1983. Bees of the genus Pithitis from northeast Africa and pods (KSMA), Department of Plant Protection, College of the Arabian Peninsula (Hymenoptera: Apoidea). J. Kansas Ento- mol. Soc. 56 (4), 499–505. Food and Agriculture Sciences, Riyadh, Kingdom of Saudi Dathe, H.H., 1995. Studien zur Systematik und Taxonomie der Arabia and the Division of Entomology (Snow Entomolog- Gattung Hylaeus F. (Apidae, Colletinae). 2. Kla¨rung und Neubes- ical Collections), University of Kansas Natural History chreibung von Arten arabischer La¨nder. Beitr. Entomol. 45 (1), Museum, Lawrence, Kansas, USA. 155–174. Promelitta is a remarkably distinctive and apparently relic Dathe, H.H., 2009. Order Hymenoptera, superfamily Apoidea: Fam- genus of Dasypodainae, and is even segregated into its ilies Colletidae, Andrenidae, Halictidae, Melittidae, Megachilidae own tribe, Promelittini. While it is largely known as dis- and Apidae. Arthropod Fauna UAE 2, 335–432. junct across northern Africa (Michez et al., 2007), it was Dathe, H.H., 2010. Studien zur Systematik und Taxonomie der recently recorded for the United Arab Emirates (Dathe, Gattung Hylaeus F. (6). Arten asiatischer Hochgebirge und 2009). Herein we record a single female from central Saudi Anmerkungen zu weiteren asiatischen Arten (Hymenoptera, Anthophila, Colletidae). Linzer Biol. Beitr. 42 (1), 43–80. Arabia as follows: Ebmer, A.W., 1984. Die westpala¨arktische Arten der Gattung Dufou- Promelitta alboclypeata (Friese): 1$, labeled as, ‘‘Saudi Ara- rea Lepeletier 1841 mit illustrierten Bestimmungstabellen (Insecta: bia, Riyadh, Al Amariah, [Mazra’ah] Majra Al-Gasim Hymenoptera: Apoidea: Halictidae: Dufoureinae). Senckenberg. [farm], 3-iv-2011 [3 April, 2011], Hannan & Naser’’. Mate- Biol. 64 (4–6), 313–379. rial is deposited in the King Saud University Museum of Ebmer, A.W., 1985. Neue westpala¨arktische Halictidae V. (Hymenop- Arthropods (KSMA), Department of Plant Protection, tera, Apoidea) sowie Festlegung von Lectotypen von Morawitz College of Food and Agriculture Sciences, Riyadh, King- beschriebener, bisher ungekla¨rter Halictus-Arten. Linzer Biol. dom of Saudi Arabia. It should be noted that there are Beitr. 17 (1), 197–221. some minor discrepancies between the UAE and Saudi Ara- Engel, M.S., 2004. 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