Monotypic no more – a new species of the unusual genus Schwarzia (Hymenoptera, Apidae, Biastini)
expand article infoSilas Bossert
‡ Cornell University, Ithaca, United States of America
Open Access


Schwarzia elizabethae Bossert, sp. n., a previously unknown species of the enigmatic cleptoparasitic genus Schwarzia Eardley, 2009 is described. Both sexes are illustrated and compared to the type species of the genus, Schwarzia emmae Eardley, 2009. The male habitus of S. emmae is illustrated and potential hosts of Schwarzia are discussed. Unusual morphological features of Schwarzia are examined in light of the presumably close phylogenetic relationship to other Biastini. The new species represents the second species of Biastini outside the Holarctic region.


Bees, cleptoparasitism, East Africa


Biastini (Apidae, Nomadinae) is a small tribe of cleptoparasitic bees with just 13 described species in four genera (Ascher and Pickering 2019). Their distribution is primarily Holarctic. Biastes Panzer, 1806 is Palearctic and has five described species. Four species were treated by Warncke (1982), and B. popovi Proshchalykin & Lelej, 2004 has been described since. Neopasites Ashmead, 1898 has five described species restricted to the Nearctic; they are morphologically similar to Biastes and the genera were synonymized by certain authors (e.g., Warncke 1982). Currently, Neopasites is regarded as a separate genus (Michener 2007). The distinguishing morphological characters are rather unobtrusive, but unambiguous and stable over their entire distribution. The only other Nearctic biastine genus is Rhopalolemma Roig-Alsina, 1991, with two described species and a morphology that is somewhat different than the remaining Biastini (Roig-Alsina 1991, Rozen et al. 1997). For example, Rhopalolemma lacks the preapical mandibular tooth that all other Biastini have (Michener 2007; but see discussion below) and the number of submarginal cells may be more variable. While all other Biastini have two submarginal cells (of varying length), individuals of Rhopalolemma were documented with 2 or 3 cells, sometimes in the same individual (Rozen et al. 1997). However, intraspecific variation in the number of submarginal cells is not uncommon in bees, and bees that have principally two submarginal cells are more likely to gain a third cell rather than losing the second cell (Scarpulla 2018).

The fourth genus and the most recently described biastine bee was Schwarzia emmae Eardley, 2009. It is an enigmatic bee with bizarre morphological characters that required the description of a new genus. Eardley (2009) assigned Schwarzia to Biastini based on shared morphological features with Biastes, i.e., the presence of a pre-apical mandibular tooth, the number of two submarginal cells (of different lengths), and the elevated median area of the metanotum (Eardley 2009). He further highlighted the striking morphological differences. Most notably, the disc of the fifth tergum of the female is strongly modified with a pre-apical pseudopygidial plate (Eardley 2009), which contrasts the surrounding area with its smooth surface, ivory-colored integument, and anterior brush of thick setae (see below). Addtitionally, tergum 5 extends posteriorly with two arched projections, forming an opening which is somewhat hourglass-shaped. The sixth sternum (illustrated in Eardley 2009, van Noort 2019) is spinose, slightly hooked posteriorly, and protrudes through the opening of the fifth tergum. Such modifications are unique among bees, and an adaptive function is not understood. However, modifications of terminal metasomal segments are very common in Nomadinae, particularly of the sixth sternum (Roig-Alsina 1991, Michener 2007). They presumably serve specific functions during oviposition of the parasites’ eggs into the host bees’ brood cells (Roig-Alsina 1991, Torchio 1986). Rare detail is provided by Torchio and Burdick (1988), who describe how the cleptoparasite Epeolus compactus Cresson, 1878 uses the spinose sixth sternum to slice openings into the lining of the host’s brood cell. Unfortunately, no observational data which could elucidate the significance of the modified terminal structures of female Schwarzia is available, and potential host(s) are unknown.

Up to now, Schwarzia was only known from the type species and from very few records. All localities are in Eastern Africa, thereby rendering the genus the only Biastini outside the Holarctic (Eardley 2009). During a survey of Malaise trap material from Kenya, donated to the Smithsonian National Museum of Natural History by Robert S. Copeland, a second undescribed species was found, along with new records of S. emmae. The aim of this study is to describe the new species and provide new insights on the classification and morphology of the genus. I further discuss potential hosts in light of other bees collected during the same collecting events.

Material and methods

Specimens were collected in Kenya during four collecting events in 1998 and 1999. The type locality of the newly described species is in Tsavo East National Park in eastern Kenya, which is dominated by xeric plains and savanna habitats. As all specimens were collected in Malaise traps, no information on flower visitation is available. Photographs were taken with a Macropod Pro imaging system, consisting of a Canon EOS 6D camera and a Stackshot 3× Cognisys device. Z-stacking was carried out with Zerene Stacker Pro stacking software v1.04. The distribution map was generated with SimpleMappr (Shorthouse 2010).

The holotype is deposited in the type collection of the National Museums of Kenya (NMK). All but one paratype are deposited in the collection of the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM). A male paratype is kept in the Packer Collection at York University, Canada.

The following abbreviations are used in the description: tergum (T), sternum (S), flagellomere (FL). For the description of the male, morphometric measures are given as a range from the smallest to the largest specimen examined.


Schwarzia elizabethae Bossert, sp. n.

Figs 1, 2, 3, 4

Material examined

KENYA: Eastern Province, at Athi River, Tsavo East National Park, 2°38.51'S, 38°21.98'E, 22.-29.XI.1999, leg. R. Copeland, Malaise trap, 5♂, 1♀. Eastern Province, at Athi River, Tsavo East National Park, 2°38.51'S, 38°21.98'E, 29.III-1.IV.1999, leg. R. Copeland, Malaise trap, 1♂.


Forewing with two submarginal cells, first of which at least twice as long as second (Fig. 1A). Omaulus lamellate. Labrum broader than long. Body size smaller compared to S. emmae (S. elizabethae: 5.1–5.9 mm, S. emmae: 6.3–8.8 mm). Protrusions on supra-antennal area absent (present in S. emmae), area evenly punctate (Figs 2C, 4C). Margin of compound eye entire, not indented as in S. emmae (Figs 2C, 4C). Punctation on T1–T4 dense and distinctly coarser compared to S. emmae (Figs 2A–B, 4A–B). Integument color of tegulae tawny (compared to dark brown in S. emmae; Fig. 2G–H). Distinct pre-apical mandible tooth absent. Integument color of legs largely brown. Male: FL2–FL4 at least two times broader than long (Fig. 2E). Female: S5 in lateral view evenly rounded, median area shiny, without punctation (S5 in lateral view more angled and with punctation in S. emmae; Fig. 4E–F).

Figure 1. 

Schwarzia elizabethae Bossert, sp. n., holotype, male A habitus, dorsal view B head, frontal view C habitus, lateral view D head, dorsal view E tergum 7 with pygidial plate and genital capsule F label information as deposited with the holotype. Scale bars: 0.5 mm.

Figure 2. 

Comparison of selected diagnostic characteristics of males of Schwarzia elizabethae Bossert, sp. n. (left column: A, C, E, G) vs. S. emmae Eardley, 2009 (right column: B, D, F, H) A–B discs of T1 and T2 C–D head characteristics in lateral view. Arrows point to finely pitted protrusions on frontal area of S. emmae and to the intended inner margin of the compound eye E–F antennae G–H tegulae.


Male (Fig. 1). Head: Head (Fig. 1B) wider than long (1.6–1.9 mm width to 1.2–1.4 mm length). Integument black with thick short setae around antennal sockets, clypeus, labrum, genal area, vertex. Antennal sockets in lower half of frons. Clypeus protruding, much shorter than broad (0.6–0.8 mm width to 0.2–0.3 mm length). Punctation on clypeus deep and coarse. Interspaces shiny, averaging less than half diameter of individual pits. Mandible dark, densely hairy, pre-apical teeth absent or just very minutely produced, outer ridge strongly developed. Labrum broad, about 1.3× times broader than long. Proboscis very long (~2.5–2.8 mm, Fig. 1A–C). Maxillary palps with 2 palpomeres, labial palps with 3. Width of compound eye exceeds twice genal width in lateral view (Figs 1C, 3C). Deep, pit-like punctation more or less evenly distributed on supraclypeal area, supra-antennal area, frons, vertex (Fig. 1B, 1D). Interspaces shiny. Flagellum with 10 flagellomeres, FL2 (0.14–0.18 mm width to 0.07–0.08 mm length) and FL3 (0.16–0.19 mm width to 0.07–0.08 mm length) at least twice as broader as long (Fig. 2E). Coloration black to brown. Diameter of lateral ocellus 0.13–0.14 mm, median ocellus 0.14–0.15 mm. Ocellocular distance 0.32–0.35 mm. Mesosoma: Integument black, except legs and pronotal lobe tawny-colored (Fig. 1A–C). Short-thick setae primarily present on margins of segments, i.e., along episternal groove, scrobal groove, around pronotal lobe, posterior margin of scutum, around scutellum, along midline of scutum, on metanotum and metapostnotum. Intertegular distance 1.2–1.5 mm. Axillae weakly produced. Size and density of punctation on scutum similar to punctation on vertex. Interspaces shiny, up to 3 times puncture diameter. Punctation on scutellum slightly larger and denser. Integument of tegulae largely light-colored (Fig. 2G). Wing venation brown. Meso- and metatibial spurs with two ventrally oriented rows of teeth, most strongly produced on metatibia. Metasoma: Integument predominantly black. Junctions of terga brown. T1 with basal patch of thick white setae. T1–T5 with basal bands of setae, which may be covered by apical margin of preceding tergum. Punctation on T1–T5 coarse, interspaces smaller than pit diameters (Fig. 2A). T6 covered with setae. T7 with distinctly pitted pygidial plate, shown together with genital capsule in Fig. 1E. S5 with single patch of white setae medially. Remaining visible sterna unmodified.

Figure 3. 

Schwarzia elizabethae Bossert, sp. n., paratype, female A habitus, dorsal view B head, frontal view C habitus, lateral view D head, dorsal view E tergum 5 F label information as deposited with the only female paratype. Scale bars: 0.5 mm.

Figure 4. 

Comparison of selected diagnostic characteristics of females of Schwarzia elizabethae Bossert, sp. n. (left column: A, C, E) vs. S. emmae Eardley, 2009 (right column: B, D, F) A–B discs of T1 and T2 C–D head characteristics in lateral view. Arrows point to finely pitted protrusions on frontal area of S. emmae and to the intended inner margin of the compound eye E–F details of T5. Arrows point to the differently curved median area of the disc.

Female (Fig. 3). Overall very similar to male except terminal terga. Head: Head (Fig. 3B) wider than long (1.8 mm width to 1.4 mm length). Integument black with thick short setae around antennal sockets, clypeus, labrum, genal area, vertex. Antennal sockets in lower half of face. Clypeus protruding, much shorter than broad (0.8 mm width to 0.3 mm length). Punctation on clypeus deep, more irregular than on supra-antennal area and frons. Interspaces shiny, averaging less than half diameter of individual pits. Mandible mostly brown, tips darkened. Mandible densely hairy, pre-apical teeth absent, outer ridge strongly developed. Labrum broad, about 1.3× times broader than long. Maxillary palps with 2 palpomeres, labial palps with 3. Width of compound eye exceeds twice genal width in lateral view (Figs 3C, 4C). Deep, pit-like punctation evenly distributed on supraclypeal area, supra-antennal area, frons, vertex (Fig. 3B, 3D). Protrusions on frontal area entirely absent. Flagellum with 10 flagellomeres, FL2 (0.14–0.18 mm width to 0.07–0.08 mm length) and FL3 more than twice as broad as long. Coloration black to brown. Diameter of lateral ocellus 0.14 mm, median ocellus 0.16 mm. Ocellocular distance 0.36 mm. Mesosoma: As in male except intertegular distance 1.4 mm. Metasoma: Integument predominantly black. T3–T4 laterally brown. T5 laterally and apically brown. T1 with basal patch of thick white setae. Punctation on T1–T4 coarse, interspaces generally smaller than pit diameters (Fig. 4A). Disc of T5 (Fig. 3E) bears distinct teardrop-shaped, ivory-colored pseudopygidial plate with anterior brush of golden setae. Posterior T5 with two hook-shaped projections medio-laterally, forming opening through which bifid S6 protrudes. Punctation on basal areas of S2–S5 fine and granulose, distinctly different from coarser punctation medially and posteriorly. S1–S5 without median patches of setae.


Males of both described species of Schwarzia have only 10 flagellomeres. For nearly all species of bees, male individuals have 11 flagellomeres and females have 10 (Michener 2007, Schmiedeknecht 1930). As both sexes of Schwarzia have this character, which is usually associated with females, it seems appropriate to designate female first names as species names for the genus. The first described species of the genus, Schwarzia emmae, was named after Emma Schwarz, the late wife of Austrian melittologist Maximilian Schwarz (Eardley 2009). The new species is named after American hymenopterist Elizabeth A. Murray.

Additional records

KENYA: Eastern Province, base of Ukasi Hill, 0°49.262'S, 38°32.666'E, 21.XI.-05.XII.2011, leg. R. Copeland, Malaise trap, Acacia/Commiphora savanna, 1♀. Eastern Province, base of Ukasi Hill, 0°49.201'S, 38°32.627'E, 5.-12.V.2018, leg. R. Copeland, Malaise trap, Acacia/Commiphora savanna, 1♂. Eastern Province, Mulu Musingila farm, 2°6.847'S, 38°14.393'E, 13.-27.XII.2016, leg. R. Copeland, Malaise trap, farmland near small seasonal wet area, 1♂. All additional records are deposited in the ICIPE collection, Nairobi, Kenya.


The species is known from four different localities, all of which are in the former Eastern Province of Kenya (Fig. 5). All sites are dominated by savanna type habitats.

Figure 5. 

Distribution map of Schwarzia, including the reported localities from Eardley (2009). Shown are selected towns near the collection localities.

Schwarzia emmae Eardley, 2009

Figs 2, 4, 6, 7

Material examined

KENYA: Rift Valley Province, Mathews Range, Sarara Camp, 1.1°N, 37.387°E, 10.-13.VI.1998, leg. R. Copeland, Malaise trap, 1♂, 1♀, dep. in USNM. Eastern Province, near Athi River, Tsavo East National Park, 2°37'S, 38°22'E, 10.VI.1998, leg. R. Copeland, Malaise trap, 1♂, 1♀, dep. in USNM.


With two newly reported collecting sites, the species is now known from a total of five different localities in Kenya and Tanzania (Fig. 5).


Classification and morphological features

Schwarzia are unusual bees that can easily be recognized and distinguished from other cleptoparasitic bees in the Afrotropical region. Nonetheless, the discovery of a new and distinctly different species allows us to refine the diagnosis of the genus and re-evaluate characters that were previously thought to be diagnostic. First, the pre-apical tooth described as diagnostic for the genus (Eardley 2009) is lacking in S. elizabethae. The mandible is unidentate in both sexes; there may only be a very slight elevation on the inner margin of the mandible. The structure is different from the truly bidentate mandibles of presumably closely related species of Biastini, such as B. brevicornis (Panzer, 1798) or B. emarginatus (Schenck, 1853). Eardley (2009) further noted that the first recurrent vein meets the first submarginal crossvein at the posterior margin of the submarginal cells in the forewing (e.g., Fig. 6C). While this character holds up for S. emmae, it is more variable in S. elizabethae, with some individuals having this character, while in others the first recurrent vein merges distal to the submarginal crossvein. However, examination of both species of Schwarzia revealed two additional diagnostic features. First, all individuals have conspicuously broadened tibial spurs which possess two ventral rows of minute teeth. This character is most obvious for the large inner metatibial spur and is somewhat similar to the modified tibial spur of Austrosystropha Patiny & Michez, 2006, a subgenus of Systropha Illiger, 1806 (Patiny et al. 2013, Patiny and Michez 2006), and certain species of Neopasites (e.g., N. cressoni Crawford, 1916; N. fulviventris (Cresson, 1878)). Second, males of Schwarzia only have 10 flagellomeres. As described in the etymology section, males of most known bee species have 11 flagellomeres (= 13 antennal segments, including scape and pedicel) and females have 10 (= 12 antennal segments).

Figure 6. 

Schwarzia emmae Eardley, 2009, male A habitus, dorsal view B head, frontal view. Dotted line highlights the paired protrusions on the frontal area, which are noticeably more finely pitted than the surround areas C habitus, lateral view D head, dorsal view. Scale bars: 1 mm.

The reduced number of antennal segments in Schwarzia males is indicative of a close relationship to other Biastini. Reduced flagellomeres of males are generally rare in bees but occur more frequently in certain tribes of cleptoparasitic Apidae. In Ammobatini, males of Pasites Jurine, 1807, Melanempis Saussure, 1890, Parammobatodes Popov, 1932 and Chiasmognathus Engel, 2006 have reduced numbers of flagellomeres (Eardley and Brothers 1996, Engel and Packer 2013, Michener 2007, Pauly et al. 2001), and so do Neopasites (e.g., N. cressoni, N. fulviventris), and certain Biastes (e.g., B. brevicornis, B. emarginatus) in Biastini. Some species of Holcopasites Ashmead, 1899 (Ammobatoidini), such as H. heliopsis (Robertson, 1897) or H. haematurus Cockerell & Hicks, 1926, have reduced male antennae as well. Further, all these bees have usually just two submarginal cells, both characteristics that could indicate a closer relationship to Schwarzia. However, the labrum of Schwarzia is clearly broader than long, a decisive characteristic which delineates the Biastini from all the aforementioned bees. I therefore follow Eardley’s (2009) understanding that Schwarzia is most likely a lineage of Biastini. Furthermore, I argue that even though the morphology of the females’ fifth tergum is clearly derived, the large projections could actually be homologous to the much smaller posterolateral projections on the fifth tergum of certain species Biastes or Neopasites, such as a B. brevicornis, or N. fulviventris (Fig. 7).

Figure 7. 

Comparison of modified apical terga of different female Biastini A Biastes brevicornis B Neopasites fulviventris C Schwarzia emmae. Scale bars: 1 mm.

An interesting and prominent morphological difference between S. emmae and S. elizabethae are the finely punctate supraclypeal protrusions of S. emmae, which are entirely absent in S. elizabethae (Figs 2C-D, 4C-D). Very similar morphological structures occur in several not closely related bees, such as diverse species of Epeolus Latreille, 1802 (e.g., E. bifasciatus Cresson, 1864, E. boliviensis Friese, 1908; Onuferko 2017, Onuferko 2018), Holcopasites (H. bigibbosus Hurd & Linsley, 1972) or Chilicola Spinola, 1851 (e.g., Chilicola obesifrons Packer, 2007; Packer and Genaro 2007). Certain species of the Coelioxys Latreille, 1809 subgenera Synocoelioxys Mitchell, 1973 (e.g., Coelioxys apacheorum Cockerell, 1900, Coelioxys texana Cresson, 1872) and Glyptocoelioxys Mitchell, 1973 bear granulose swellings around the median ocellus, which are similar in surface structure to protrusions of aforementioned taxa. As all these taxa are quite distantly related, it can be assumed that these features evolved independently and that they are not necessarily similar in function. However, while the exact function of any of these protrusions is unknown, it seems plausible that they may be involved in sensory perception or processing. Interestingly, all bees but Chilicola are cleptoparasites, which indicates that the evolution of such protrusions may at least be favored in bees that do not collect pollen, but search and parasitize nests of host bees.

Potential hosts of Schwarzia

Due to the lack of field observational data for Schwarzia, the biology of both species is essentially unknown. All presently available material was collected with traps and therefore no information on flower visitation or potential host(s) is available. Nonetheless, the morphology and presumably close phylogenetic relationship to Biastes leaves no doubt that Schwarzia has a cleptoparasitic lifestyle. All Biastini are cleptoparasites of species of Rophitinae (Michener 2007), and Eardley (2009) singles out Systropha as the only rophitine genus in Eastern Africa. Systropha is common in Eastern Africa (Eardley 2009) and at least five species are known from Kenya or directly adjacent countries (Ascher and Pickering 2019, Bossert and Patiny 2017, Patiny et al. 2013, Patiny and Michez 2006), with likely more species to be documented.

Schwarzia and the potential host Systropha even share certain morphological features: both sexes have an unusually long glossa, which is rather uncommon for cleptoparasitic bees. The long mouthparts suggest that Schwarzia bees collect nectar at flowers with a deep corolla, and therefore may even visit the same flowers as their potential host Systropha, which are specialists on flowers of the morning glory family (Convolvulaceae; Michener 2007, Patiny et al. 2008). Nonetheless, no specimen of Systropha was collected during the same trap collecting events coincident with the specimens of Schwarzia examined in this study. Bees of the following genera were collected with Schwarzia:

Collecting event 1 (near the Sarara Camp, 10.-13.VI.1998; collection site of S. emmae): Afranthidium Michener, 1949, Amegilla Friese, 1897, Braunsapis Michener, 1969, Ceratina Latreille, 1802, Macrogalea Cockerell, 1930, Macronomia Cockerell, 1917, Maynenomia Pauly, 1984, Nubenomia Pauly, 1980, Seladonia Robertson, 1918, Thrinchostoma de Saussure, 1890.

Collecting event 2 (in Tsavo East National Park, 10.VI.1998; collection site of S. emmae): Acunomia Cockerell, 1930, Afromelecta Lieftinck, 1972, Amegilla, Anthidium Fabricius, 1804, Ceratina, Coelioxys, Lasioglossum s.l. Curtis, 1833, Lipotriches Gerstaecker, 1858 subgenus Patellotriches Pauly, 2014, Lipotriches subgenus Stellotriches Pauly, 2014, Macronomia, Megachile Latreille, 1802, Meganomia Cockerell, 1909, Mermiglossa Friese, 1912, Nubenomia, Pseudapis (s. str.) Kirby, 1900, Seladonia, Sphecodes Latreille, 1804, Thyreus Panzer, 1806.

Collecting event 3 (in Tsavo East National Park, 22.-29.XI.1999; collection site of S. elizabethae): Amegilla, Braunsapis, Coelioxys, Lipotriches subgenus Stellotriches, Macrogalea, Maynenomia, Megachile, Pasites Jurine, 1807, Pseudapis (s. str.).

Collecting event 4 (in Tsavo East National Park, 29.III-1.IV.1999; collection site of S. elizabethae): Pasites.

Non-parasitic bees were only collected in collecting events 1, 2, and 3. The only bee genus that was collected during all these three events is Amegilla, an apid bee genus in the subfamily Anthophorinae (Bossert et al. 2019). However, since Amegilla is diverse, common, and widespread throughout Africa (Eardley et al. 2010), it is quite likely that these findings are simply a representation of the abundance of Amegilla at those sites, and do not necessarily reflect a host-parasite relationship. Most bees of the genus Amegilla are also much larger than the species of Schwarzia, and all collected specimens of Amegilla significantly exceeded the size of Schwarzia. In the future, field observational efforts in Eastern Africa are required to shed light on the potential host of Schwarzia, in order to better understand the biology of these intriguing bees.


The species examined in this study are part of a donation of insects made by Robert Copeland to the U.S. National Museum of Natural History. I thank him for his support to develop this study. I further thank Matthew Buffington (USDA-ARS) for granting access to the imaging system, and Laurence Packer, Michael Orr, and Michael Kuhlmann for discussion on the manuscript. Lastly, I thank Simon van Noort (Iziko South African Museum) for correspondence concerning S. emmae. This work was supported by a U.S. National Science Foundation (NSF) grant to Bryan N. Danforth, Seán G. Brady, James P. Pitts, and Robert M. Ross (DEB-1555905), and a Peter Buck predoctoral fellowship at the Smithsonian Institution to the author.


  • Bossert S, Murray EA, Almeida EAB, Brady SG, Blaimer BB, Danforth BN (2019) Combining transcriptomes and ultraconserved elements to illuminate the phylogeny of Apidae. Molecular Phylogenetics and Evolution 130: 121–131.
  • Bossert S, Patiny S (2017) Description of the previously unknown male of Systropha (Austrosystropha) macronasuta (Hymenoptera: Halictidae: Rophitinae) from Kenya. Journal of Melittology 67: 1–9.
  • Eardley C (2009) First record of the tribe Biastini from the Afrotropical Region (Hymenoptera: Apidae). Zootaxa 2264: 65–68.
  • Eardley CD, Brothers DJ (1996) Phylogeny of the Ammobatini and revision of the Afrotropical genera (Hymenoptera: Apidae: Nomadinae). Journal of Hymenoptera Research 6: 353–418.
  • Eardley CD, Kuhlmann M, Pauly A (2010) The bee genera and subgenera of Sub-Saharan Africa. Abc Taxa 7: 1–139.
  • Michener CD (2007) The Bees of the World. The Johns Hopkins University Press, Baltimore, 953 pp.
  • Onuferko TM (2017) Cleptoparasitic bees of the genus Epeolus Latreille (Hymenoptera: Apidae) in Canada. Canadian Journal of Arthropod Identification: 1–62.
  • Patiny S, Baldock D, Michez D (2013) Systematics of the bee subgenus Systropha (Austrosystropha) (Hymenoptera: Halictidae): Description of a new species and proposal of a new sex association. Zootaxa 3647: 577–584.
  • Patiny S, Michez D (2006) Phylogenetic analysis of the Systropha Illiger 1806 (Hymenoptera: Apoidea: Halictidae) and description of a new subgenus. Annales de la Société entomologique de France (NS) 42: 27–44.
  • Pauly A, Brooks R, Nilsson L, Pesenko Y, Eardley C, Terzo M, Griswold T, Schwarz M, Patiny S, Munzinger J (2001) Hymenoptera Apoidea de Madagascar et des îles voisines. Annales du Musée royal de l’Afrique centrale (Sciences Zoologiques) 286: 1–406.
  • Proshchalykin MY, Lelej AS (2004) New and little known bees (Hymenoptera: Colletidae, Apidae) from the Russian Far East. Far Eastern Entomologist 136: 1–10.
  • Roig-Alsina A (1991) Cladistic analysis of the Nomadinae s. str. with description of a new genus (Hymenoptera: Anthophoridae). Journal of the Kansas Entomological Society 64: 23–37.
  • Rozen JG, Roig-Alsina A, Alexander BA (1997) The cleptoparasitic bee genus Rhopalolemma, with reference to other Nomadinae (Apidae), and biology of its host Protodufourea (Halictidae, Rophitinae). American Museum Novitates 3194: 1–28.
  • Scarpulla EJ (2018) Four submarginal cells on a forewing of Melitoma taurea (Say) (Hymenoptera: Apidae), and a summary of known records of atypical and variable numbers of submarginal cells. Insecta Mundi 0667: 1–28.
  • Schmiedeknecht O (1930) Die Hymenopteren Nord- und Mitteleuropas. G. Fischer, Jena, 1062 pp.
  • Shorthouse DP (2010) SimpleMappr, an online tool to produce publication-quality point maps. [accessed 9 January 2018]
  • Torchio PF (1986) Late embryogenesis and egg eclosion in Triepeolus and Anthophora with a prospectus of Nomadine classification (Hymenoptera: Anthophoridae). Annals of the Entomological Society of America 79: 588–596.
  • Torchio PF, Burdick D (1988) Comparative notes on the biology and development of Epeolus compactus Cresson, a cleptoparasite of Colletes kincaidii Cockerell (Hymenoptera: Anthophoridae, Colletidae). Annals of the Entomological Society of America 81: 626–636.
  • van Noort S (2019) WaspWeb: Hymenoptera of the Afrotropical region. [accessed 9 January 2019]
  • Warncke K (1982) Zur Systematik der Bienen – Die Unterfamilie Nomadinae (Hymenoptera, Apidae). Entomofauna 3: 97–126.