Content uploaded by Jakub Straka
Author content
All content in this area was uploaded by Jakub Straka
Content may be subject to copyright.
Accepted by C. Rasmussen: 20 Jan. 2012; published: 14 May 2012
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2012 · Magnolia Press
Zootaxa 3311: 1–41 (2012)
www.mapress.com/zootaxa/Article
1
Review and identification of the cuckoo bees of central Europe
(Hymenoptera: Halictidae: Sphecodes)
PETR BOGUSCH1 & JAKUB STRAKA2
1University of Hradec Králové, Department of Biology, Rokitanského 62, CZ-500 03 Hradec Králové, Czech Republic.
E-mail: bogusch.petr@gmail.com
2Charles University in Prague, Faculty of Science, Department of Zoology, Vini
č
ná 7, CZ-128 44 Praha 2, Czech Republic.
E-mail: straka.jakub.1@gmail.com
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
Identification key of Sphecodes of central Europe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
List of species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Sphecodes albilabris (Fabricius, 1793) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
Sphecodes alternatus Smith, 1853 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Sphecodes crassanus Warncke, 1992 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Sphecodes crassus Thomson, 1870. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Sphecodes cristatus Hagens, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Sphecodes croaticus Meyer, 1922 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9
Sphecodes dusmeti Blüthgen, 1924 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10
Sphecodes ephippius (Linné, 1767) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Sphecodes ferruginatus Hagens, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10
Sphecodes geoffrellus (Kirby, 1802) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10
Sphecodes gibbus (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Sphecodes hyalinatus Hagens, 1882. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Sphecodes intermedius Blüthgen, 1923 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Sphecodes longulus Hagens, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Sphecodes majalis Pérez, 1903. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Sphecodes marginatus Hagens, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .13
Sphecodes miniatus Hagens, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .13
Sphecodes monilicornis (Kirby, 1802) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Sphecodes niger Hagens, 1874 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Sphecodes nomioidis Pesenko, 1979. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Sphecodes olivieri Lepeletier, 1825 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Sphecodes pellucidus Smith, 1845 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Sphecodes pinguiculus Pérez, 1903 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .15
Sphecodes pseudofasciatus Blüthgen, 1925 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Sphecodes puncticeps Thomson, 1870 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Sphecodes reticulatus Thomson, 1870 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16
Sphecodes rubicundus Hagens, 1875 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Sphecodes ruficrus (Erichson, 1835) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Sphecodes rufiventris (Panzer, 1798) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .17
Sphecodes scabricollis Wesmael, 1835. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Sphecodes schenckii Hagens, 1882 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .18
Sphecodes spinulosus Hagens, 1875 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .18
Sphecodes zangherii Noskiewicz, 1931 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Acknowledgement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .38
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .39
BOGUSCH & STRAKA
2 · Zootaxa 3311 © 2012
Magnolia Press
Abstract
We reviewed nomenclature, biology, hosts, geographical distribution and compiled an identification key for all 33 Sphe-
codes Latreille, 1804 species known from central Europe. The identification key is separated for females and males and
include 204 figures (photographs) of identification characters as well as male genitalia of all species. Taxonomically dif-
ficult groups within the genus were critically studied and new characters, as well as corrected geographical distribution,
are presented, i.e., the S. reticulatus group (S. alternatus Smith, 1853, S. crassanus Warncke, 1992 and S. reticulatus
Thomson, 1870), S. croaticus group (S. croaticus Meyer, 1922, S. pseudofasciatus Blüthgen, 1925 and S. zangherii Nosk-
iewicz, 1931) and S. miniatus group (S. marginatus Hagens, 1882, S. miniatus Hagens, 1882 and S. nomioidis Pesenko,
1979). The name S. nomioidis is used because it is the only available name for the taxon formerly identified as S. margi-
natus in Eastern Europe. Sphecodes capverdensis Pesenko & La Roche, 2002 is considered to be a junior synonym of S.
pinguiculus Pérez, 1903 (syn. nov.). In addition we summarized all known host records of Sphecodes, including a discus-
sion of the likelihood of published data and presentation of new host data.
Key words: identification key, taxonomy, ecology, hosts
Introduction
The bees (Hymenoptera: Apoidea: Apiformes) represent one of the richest groups of aculeate Hymenoptera in
terms of species. Yet, even as of recently several taxonomicaly complicated groups remain poorly studied and
rarely reviewed. Such typical groups are within the family Halictidae where the only European cuckoo bee genus,
Sphecodes Latreille, 1804, has been studied several times, but remains one of the most difficult European bee gen-
era to identify at the species level. Knowledge on the taxonomy of the genus was summarized by Hagens (1874,
1875, 1882), who also described the majority of all currently valid species. His descriptions were for the most part
the source of information for compilation of later identification keys. Additional species were described by Meyer
(1919, 1922) and Noskiewicz (1931). Recently, eight new species from Italy were described by Campadelli &
Nobile (2000) and Nobile & Turrisi (2004). However, the validity of these species is doubtful (Schwarz & Gusen-
leitner 2012) and thus, we do not include them in this article. Burger & Reum (2004) and Burger et al. (2006)
examined difficult groups of this genus, in particular S. croaticus Meyer, 1922 and S. zangherii Noskiewicz, 1931.
European Sphecodes have been keyed four times. Blüthgen (1923a) compiled the first key, followed by Šustera
(1959). Although the nomenclature used in those papers is old, the authors found many previously unknown char-
acters. A useful key was published by Warncke (1992). This publication contains good drawings of male genitalia
and distribution maps of all European species. However, most of the key couplets are based on a single and some-
times a very variable character. Thus the key is good but some species are hardly identifiable. The latest key, pub-
lished by Amiet et al. (1999), is in comparison to Warncke’s key with fewer species but rely on better identification
characters.
The biology of Sphecodes was studied by few authors. Brief references are found in monographs on the bees of
Europe (Friese 1898, Stoeckhert 1930, Westrich 1989, Macek et al. 2010), where host species of Sphecodes are
mentioned occasionally. Blüthgen (1923b, 1934) provided the first review on the biology of Sphecodes, including
the hosts. These data were cited by Westrich (1989), Celary (1991) and Amiet et al. (1999). Hosts of Sphecodes
were also studied by Vegter (1985, 1993). Sick et al. (1994) studied hosts of selected Sphecodes species experimen-
tally and reported new information. Except some ecological studies reporting Sphecodes hosts and mentioned in
the text under the appropriate species, the following species have been studied in detail: S. cristatus Hagens, 1882
in Sweden by Svensson (1982), S. majalis Pérez, 1903 by Herrmann et al. (2003), and S. ruficrus Erichson, 1852
by Herrmann (2006). Host specificity of two common species, S. ephippius (Linnaeus, 1767) and S. monilicornis
(Kirby, 1802) was studied by Bogusch et al. (2006). Data on the visited flowers were published by Westrich (1989)
and Celary (1991), demonstrating polylectic behavior of all species. Sphecodes bees are nest cleptoparasites, they
only forage on nectar on flowers and do not collect pollen. Thus the visited flowers are not discussed in this publi-
cation.
Here, we would like to present a newly compiled identification key based on the study of the material of all
included species, and emphasizing the advantages and eliminating the disadvantages of previously published keys.
The publication also reviews the synonymy, taxonomy, distribution, and biology of all species included. Host spec-
trum of all species is critically analyzed. The key includes all species known from central Europe and closely allied
areas.
Zootaxa 3311 © 2012 Magnolia Press · 3
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
Methods
The nomenclature was adopted from the following sources: Warncke (1992), Schwarz et al. (1996) and Bogusch et
al. (2007). The material studied is from the following collections: BMNH—The Natural History Museum, London,
UK, HNHM—Magyar Természettudományi Múzeum Allattara, Budapest, Hungary, JSPC—Jakub Straka, private
collection, Prague, Czech Republic, MSAC—Maximilian Schwarz, private collection, Ansfelden, Austria,
NMPC—National Museum, Prague, Czech Republic, PBHK—Petr Bogusch, private collection, Hradec Králové,
Czech Republic. Frank Burger (Weimar, Germany), Mike Herrmann (Konstanz, Germany), Toshko Ljubomirov
(Sofia, Bulgaria), Denis Michez (Mons, Belgium) and Guido Pagliano (Torino, Italy) sent us gift material of Sphe-
codes. In the key, the following abbreviations are used: T—tergite, S—sternite (T4 means the fourth tergite). Other
morphological terms are used in English and Latin languages. The characters in all parts are sorted from the most
important (best seen and least variable). Most of the characters were photographed and are shown under appropri-
ate number within the captions. Numbers of figures are put in square brackets to the text of the key. The key also
contains photos of male genitalia of all species from dorsal and lateral view. The list of species includes all species
mentioned in the key, with the following information: name, synonymy, distribution and ecology (with a focus to
host species and biotope preferences). Only names and pages within the publication with description of all taxa are
mentioned, whole citations of the descriptions are in the reference list.
Identification key of Sphecodes of central Europe
Females (
♀♀
)
1 a. Clypeus about three times wider than long, with furrow in the middle and teeth on sides, shorter than supraclypeus [fig. 1];
scutum finely and densely punctate, 6–8 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. majalis Pérez
1 b. Clypeus less than three times (usually 2x) wider than long, without teeth on sides [fig. 32]; scutum coarsely punctate . . . . . . 2
2 a. Vertex behind the ocelli with sharp transverse ridge [fig. 2]; flagellomeres one and half longer than wide, so the antenna looks
longer than other species; 1st flagellomere distinctly wider than 2nd [fig. 3], 8–11 mm . . . . . . . . . . . . . . . . . spinulosus Hagens
2 b. Vertex without ridge; flagellomeres maximally as long as wide, equal in width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 a. Hind wing with 7–14 hamuli [fig. 4]; angle between cubital and anal veins on hind wing usually acute [fig. 6], if the angle
looks right, then head square and large; vertex behind the ocelli usually gibbous. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
3 b. Hind wing with 5–6 hamuli [fig. 5] (only in some specimens of S. pellucidus, S. rubicundus and S. ruficrus up to 8 hamuli but
they all have right angle between veins on hindwing); usually right angle between cubital and anal veins on hind wing [fig. 7];
vertex behind the ocelli not gibbous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14
4 a. Gena on backside with sharp ridge [fig. 8], sharp or weak (S. scabricollis) elongate ridge behind ocelli . . . . . . . . . . . . . . . . .5
4 b. Gena without ridge, ridge behind ocelli absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
5 a. Face (clypeus and paraocular area), pronotum and scutellum with dense felt-like white pubescence; base of T1 very coarsely
punctate; thorax often reddish [fig. 10], 8–11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . olivieri Lepeletier
5 b. Face, pronotum and scutellum without felt-like pubescence; base of T1 unpunctate; thorax black. . . . . . . . . . . . . . . . . . . . . . 6
6 a. Vertex behind the ocelli with elongate ridge [fig. 11]; scutum coarsely but sparsely punctate; tibiae and tarsi reddish; smaller
and slenderer species, 6–8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cristatus Hagens
6 b. Vertex behind the ocelli only with weak elongate ridge, which is in some cases very poor; scutum very densely and coarsely
punctate [fig. 9]; legs dark; larger, robust species, 9–12 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . scabricollis Wesmael
7 a. Genae behind the eyes wide, vertex gibbous; genal width similar as the width of compound eye; head square-shaped [fig. 12];
angle between cubital and anal veins on hind wing acute but nearly right (about 80°), 7–10 mm . . . . . . monilicornis (Kirby)
7 b. Genae narrow, narrower than the compound eye; head looks flat and smaller; angle between cubital and anal veins on hind
wing always acute . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8
8 a. Mesopleura and propodeal sides regularly and finely ridged, ridges reach to hypoepimeral area and scutellum [fig. 15]; legs. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . rufiventris (Panzer)
8 b. Mesopleurae grained without elongate ridges [fig. 14] (except ventral part of mesopleurae and propodeal sides with coarser
ridges in S. schenckii); legs with pale or dark hair . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9 a. Scutum coarsely and densely punctate, interspaces smaller than punctures [fig. 13]; tergites also coarsely and densely punctate;
wings dark; large and conspicuous species with red abdomen with dark apex (only last tergite completely black), 11–15 mm .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . albilabris (Fabricius)
9 b. Scutum coarsely but sparsely punctate with shiny and large interspaces among punctures; tergites usually conspicuously
punctate, but not as in S. albilabris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10
10 a. Basis of T4 densely punctate [fig. 18]; vertex behind ocelli not elevated [fig. 16]; genae very narrow, getting narrower just
behind the compound eyes; pygidium shiny and as wide as first flagellomere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
10 b. Basis of T4 sparsely punctate [fig. 19]; vertex behind ocelli strongly elevated [fig. 17]; genae wider; pygidium dull, distinctly
narrower than first flagellomere . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .13
BOGUSCH & STRAKA
4 · Zootaxa 3311 © 2012
Magnolia Press
11 a. Apical depression of T4 without punctures, but finely wrinkled [fig. 18]; T1 finely and sparsely punctate; face coarsely and
densely punctate, dull, 7–10 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. reticulatus Thomson
11 b. Apical depression of T4 punctate without wrinkles; T1 coarsely punctate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
12 a. Tergites densely punctate but with distinct interspaces (well recognized on T4); end of T4 only with few fine punctures [fig.
20]; frons with distinct shiny interspaces among punctures [fig. 22], 8–11 mm . . . . . . . . . . . . . . . . . . . . . . . alternatus Smith
12 b. Tergites coarsely and densely punctate; T4 with riddle-like punctures without distinct interspaces [fig. 21]; also the end of T4
finely punctate; frons dull, very densely punctate [fig. 23], 8–11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . crassanus Warncke
13 a. Scutum only with few coarse and deep punctures [fig. 24]; abdomen nearly unpunctate (but very variable punctures); hair on
scapus as long as width of scapus [fig. 26]; face without dense white pubescence, 7–13 mm . . . . . . . . . . . gibbus (Linnaeus)
13 b. Scutum more densely punctate in the middle than on sides [fig. 25]; hair on scapus twice longer than width of scapus [fig. 27];
face usually densely whitish haired, 8–11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . schenckii Hagens
14 a. Tergites (also T1) densely and finely punctate [fig. 28]; wings hyaline, pterostigma yellow; face above antennal socket with
dense whitish pubescence [fig. 31] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
14 b. Tergites only sparsely punctate, T1 in many cases only with few punctures; wings not hyaline, usually slightly infumate; face
above antennal socket with longer sparse hair . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16
15 a. Scutum finely and sparsely punctate [fig. 29]; propodeum with fine sculpture; completely red to dark with only few red marks,
5–7 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinguiculus Pérez
15 b. Scutum in the middle coarsely but sparsely punctate [fig. 30]; propodeum with grainy sculpture; dark with red abdomen, 5–8
mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . intermedius Blüthgen
16 a. Mandibles without lateral tooth [fig. 32]; small species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .17
16 b. Mandibles with lateral tooth [fig. 33]; small to big species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .18
17 a. Head distinctly wider than long [fig. 32]; genae narrowing just behind the compound eyes; T3 densely and coarsely punctate
[fig. 37], 5–7 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . puncticeps Thomson
17 b. Head only a little wider than long [fig. 34]; genae wider; T3 only sparsely punctate [fig. 38], 4–5 mm. . . . . longulus Hagens
18 a. Hypoepimeral area smooth and shiny [fig. 35]; head nearly square-shaped, genae nearly as wide as compound eyes;
propodeum rugous only anteriorly [fig. 36]; all sterna reddish to brown, 4.5–5.5 mm . . . . . . . . . . . . . . . . . . . . . niger Hagens
18 b. Hypoepimeral area and the whole propodeum rugous; at least posterior sterna black; small to large species. . . . . . . . . . . . . .19
19 a. Pygidium wide and dull [fig. 39]; clypeus gibbous with medial furrow, all coarsely punctate [fig. 42]; scutum coarsely
punctate; medium size to big species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20
19 b. Pygidium narrow and shiny, without sculpture [fig. 40]; clypeus only sparsely punctate; scutum finely punctate or with only
few coarse punctures [fig. 41]; small species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .23
20 a. Face and thorax with long black hair; tibiae and tarsi reddish; abdomen red only with black end (last tergite) [fig. 43], 6–9 mm
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ruficrus Erichson
20 b. Face and thorax with short and pale hair, if with longer dark hair, then legs dark and/or abdomen with at least last 2 tergites
black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21
21 a. T4 red on sides and dark in the middle; apical part of T1 delimited by two rows of punctures [fig. 45]; angle between the
cubital and anal veins on hind wing 90° or slightly more [fig. 44], 8–12 mm . . . . . . . . . . . . . . . . . . . . . . . rubicundus Hagens
21 b. T4 completely dark; apical part of T1 delimited by a row of punctures [fig. 46]; angle between cubital and anal veins on hind
wing less than 90° . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22
22 a. Head wider than long; clypeus with very long, pale hair; hair on scapus twice longer than width of scapu [fig. 47]s; inner side
of front tibiae always dark, 7–11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pellucidus Thomson
22 b. Head only a little wider than long; clypeus with shorter and sparse pubescence; hair on scapus as long as the width of scapus;
inner side of front tibiae usually (but not always!!) with reddish or yellowish spot, 6–9 mm . . . . . . . . . ephippius (Linnaeus)
23 a. Pronotal projections round, not angulated [fig. 48]; dorsal part of propodeum round on sides, without sharp ledge [fig. 50]; face
very finely and densely punctate; clypeus flat without central furrow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24
23 b. Pronotal projections with sharp angle [fig. 49]; dorsal part of propodeum on sides with sharp ledge; face more coarsely but
sparsely punctate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .25
24 a. Face around antennal sulcus dull, frons densely punctate [fig. 53]; ventral part of thorax ridged [fig. 51], 6–9 mm . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ferruginatus Hagens
24 b. Face around antennal sulcus with shiny interspaces among punctures [fig. 54]; ventral part of thorax only finely sculptured
[fig. 52], 5–7 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . hyalinatus Hagens
25 a. Face square-shaped (like in S. monilicornis), outer margin of eye straight; inner margins of compound eyes parallel (see from
above) [fig. 55]; genae strongly developed behind eyes [fig. 56] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .26
25 b. Face oval, outer margin of eyes largely rounded; inner margin of compound eye (in S. croaticus only a little) convergent [fig.
62] (see from above); genae narrower (except some S. croaticus). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .27
26 a. Eyes distinctly hairy (in old individuals can be worn); lower gena densely and uniformly hairy; interocellar distance smaller
than ocelloocular distance; occiput elevated above ocelli (distinct in frontal view); interocellar area with large interspaces
among punctures [fig. 57, 58]; posterolateral part of mesopleurae dull, at most with fine punctures; T1 finely and in second half
also densely punctate, 5–8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . zangherii Noskiewicz
26 b. Eyes at most with few short hair; lower gena sparsely hairy; ocelli on the top of occiput (distinct in frontal view); interocellar
distance as long as ocelloocular distance; interocellar area densely punctate [fig. 55, 56]; posterolateral part of mesopleurae
with distinct bright shiny area with well-developed punctures; T1 finely and sparsely punctate, 5–7 mm. . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pseudofasciatus Blüthgen
27 a. Furrow in the middle of frons extending at most to the middle of frons [fig. 59, 60]; T1 with diffuse punctures; T2 and T3
Zootaxa 3311 © 2012 Magnolia Press · 5
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
densely punctate in front half [fig. 61], 5–7 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . croaticus Meyer
27 b. Furrow in the middle of frons almost touching front ocellus [fig. 74, 75]; tergites usually finely and sparsely punctate or with
diffuse punctures .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .28
28 a. Scutum densely punctate; legs dark; supraclypeus very densely punctate [fig. 62], interspaces among punctures
microsculptured, dull, 4–6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . dusmeti Blüthgen
28 b. Scutum sparsely punctate; legs dark or pale; supraclypeus sparsely punctate, shiny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .29
29 a. Flagellomere 3 longer than 1 and about as long as 4 [fig. 69]; T2 finely punctate or smooth; pygidium narrower [fig. 40], 4.5–
7 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .30
29 b. Flagellomeres 1, 2 and 3 similarly long, all distinctly shorter than long, no of them as long as flagellomere 4 [fig. 71]; at least
basal third of T2 and basal half of T3 with distinct punctures; pygidium wider [fig. 84], 4–6 mm. . . . . .(miniatus group) ...31
30 a. Head distinctly wider than long [fig. 63]; flagellomere 3 longer than 2 [fig. 69]; scapus sparsely punctate ventromedially and
shiny; hypoepimeral area rugous [fig. 65]; T2 finely punctate [fig. 67], 5–7 mm . . . . . . . . . . . . . . . . . . . . . . crassus Thomson
30 b. Head only slightly wider than long [fig. 64]; flagellomere 2 quadrate, wider and longer than flagellomere 3 [fig. 70]
(sometimes looks as subequal; measuring recommended); scapus densely punctate ventromedially; hypoepimeral area finely
striated [fig. 66]; T2 with a few punctures [fig. 68], 4.5–6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . geoffrellus (Kirby)
31. Three species very hard to distinguish in females. No character fits in 100% of individuals:
31 a. Pygidial plate very wide, surface with distinct elevated longitudinal furrow in the middle [fig. 83, 84]; frons in front of the
oceli only slightly convex, densely punctate, distances among punctures variable and often form chains and elevated lines
along, punctures variable in size, some well-defined and round, some with irregular margins [fig. 74]; vertex distinct, from
lateral view slightly decreasing behind hind oceli [fig. 72]; T2 very finely punctate, punctures ill-defined; in the middle of T2,
punctated area does not reach apical depression, puncures on the basal margin of apical depression distinctly isolated from
punctures of T2 base [fig. 80] (sometimes with more isolated punctures in this area, but punctures becoming posteriorly more
shallow and finer) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . miniatus Hagens
31 b. Pygidial plate usually narrowing posteriorly, or all narrower, surface flat, or indistinctly elevated in the middle [fig. 85, 86]
(when pygidium wider, than still indistinctly elevated in the middle); frons in front of the oceli strongly convex; frons sparsely,
or densely, but very uniformly punctate, interspaces among punctures flat, most punctures uniform in size, well defined–round
[fig. 76]; vertex short, from lateral view strongly decreasing immediately behind hind oceli [fig. 73]; T2 distinctly punctate,
punctures small, but well-defined; in some specimens in the middle of T2 [fig. 81], punctation almost reach apical depression
or fully joining punctures on base of depression .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .32
32 a. All frons uniformly densely or sparsely punctate; punctures on scutum uniformely distributed, seldom some interspaces among
punctures larger than others [fig. 76]; T2 with fine, but deep and well distinct punctures, punctated area usually does not reach
apical depression [fig. 82]; paraocular area above clypeus densely punctate, interspaces among large punctures small, filled by
small punctures; gena narrow, but just behind eyes converging just slightly, in dorsal view seems to be angle between posterior
eye margin and lateral margin of gena more than 45°; pygidial plate longer, in most specimens parallel-sided, apical margin ill-
defined; glabrous base of T6 slightly wider than pygidium [fig. 86] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . nomioidis Pesenko
32 b. Frons in front of the oceli sparsely punctate, lateral parts of frons in some specimens densely punctate [fig. 75]; punctures on
scutum uniformely distributed, except around paranotal ridges, where are distances often larger (like missing punctures); T2
with well developed and deep punctures, punctated area usually reach apical depression in the middle of T2 and joining
punctures on its margin [fig. 81]; paraocular area above clypeus more sparsely punctate, than in the rest of paraocular area, this
area looks shiny almost like clypeus; gena narrow, just behind eyes strongly converging, in dorsal view seems to be angle
between posterior eye margin and lateral margin of gena less than 45°; pygidial plate shorter, in most specimens narrowing
apically, apical margin thick; glabrous base of T6 distinctly wider than pygidium [fig. 85] . . . . . . . . . . . . marginatus Hagens
Males
♂♂
1 a. Gonocoxite without impression; small to big species [fig. 139]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
1 b. Gonocoxite with impression; smaller species [fig. 159]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .22
2 a. Upper side of hind tibiae with red spines in pale pubescence [fig. 87]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
2 b. Upper side of hind tibiae only with pale pubescence, without any spine [fig. 88]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
3 a. Occiput with transverse ridge [fig. cf. 2]; T4 and T5 finely and densely punctate; large species, gonostylus [figs. 197, 198], 9–
11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . spinulosus Hagens
3 b. Occiput without transverse ridge; T4 and T5 finely, very sparsely punctate with fine sculpture among punctures; medium size
species, gonostylus [figs. 163, 164], 6–8 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . majalis Pérez
4 a. Felt-like pubescence covers larger part of flagellomere, flagellomeres only with small circular glabrous areas; flagellomere 2
short, maximally one and half longer than wide [fig. 90]; lower parts of mesopleurae with elongate ridges [fig. 89]; genital
usually pale brown with dark apex, gonostylus [figs. 191, 192], 6–8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . rufiventris (Panzer)
4 b. Felt-like pubescence different; flagellomere 2 twice or more longer than wide; mesopleurae with grainy sculpture . . . . . . 10
5 a. Hind wing with 7–14 hamuli on the anterior part [fig. cf. 4]; upper and lower halves of flagellomeres conspicuously separated
by a ledge [fig. 95]; angle between cubital and anal veins on hind wing acute [fig. cf. 6] . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
5 b. Hind wing with 5–6 hamuli [fig. cf. 5] (only at some specimens of S. pellucidus, S. rubicundus and S. ruficrus up to 8 hamuli);
flagellomeres undivided [fig. 121]; angle between cubital and anal veins on hind wing acute to right. . . . . . . . . . . . . . . . . . 15
6 a. Sharp, crested ridge on gena [fig. cf. 8] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
BOGUSCH & STRAKA
6 · Zootaxa 3311 © 2012
Magnolia Press
6 b. Without any ridge on gena . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
7 a. Scutum densely and coarsely punctate with indistinct interspaces among punctures [fig. 93]; top of vertex with indistinct
longitudinal crista, gonostylus [figs. 193, 194], 9–12 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . scabricollis Wesmael
7 b. Interspaces among punctures on scutum well developed; top of vertex with distinct longitudinal crista . . . . . . . . . . . . . . . . . .8
8 a. Scutum sparsely, finely punctate with large interspaces [fig. 91]; T1 finely and sparsely punctate; felt-like pubescence on
flagellomeres 5–10 distinct, covers about 1/3 of the length of flagellomere [fig. 94]; sparse hair on head and thorax, gonostylus
[fig. 149, 150], 6–8 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cristatus Hagens
8 b. Scutum coarsely punctate [fig. 92]; T1 uniformely densely and coarsely punctate; felt-like pubescence on flagellomeres very
narrow, less than 1/4 of the length [fig. 95]; dense white hair on head and thorax, gonostylus [fig. 175, 176], 7–11 mm . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . olivieri Lepeletier
9 a. Scutum and tergites coarsely, densely punctate with minute interspaces among punctures [fig. 96]; the whole body looks dull,
gonostylus [figs. 139, 140], 11–15 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . albilabris (Fabricius)
9 b. Scutum and tergites sparsely punctate with large interspaces among punctures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10
10 a. Head about as long as wide [fig. 97]; apical parts of T1 to T3 very sparsely punctate [fig. 99]; gonostylus short and without
long hair [figs. 169, 170], 7–10 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . monilicornis (Kirby)
10 b. Head wider than long [fig. 98]; apical parts of T1 and T2 more densely punctate; gonostylus longer and hairy . . . . . . . . . . . 11
11 a. Vertex gibbous, behind ocelli strongly elevated, coarsely punctate with shiny interspaces [fig. 100]; apical depression of T2 to
T4 well defined . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12
11 b. Vertex behind ocelli slightly elevated, finely punctate and ridged [fig. 101]; apical depression of T2 to T4 ill defined . . . . . 13
12 a. Flagellomeres twice longer than wide; felt-like pubescence on flagellomeres 5 to 10 overall base of flagellomeres [fig. 103];
apical depression of T4 smooth, unpunctate; gonostylus with elongated hairy projection [figs. 157, 158], 7–13 mm . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . gibbus (Linnaeus)
12 b. Flagellomeres less than twice as long as wide; felt-like pubescence on flagellomeres 5 to 10 short [fig. 102]; apical depression
of T4 with sparse coarse punctures; gonostylus projection not elongated [figs. 195, 196], 8–11 mm . . . . . . . schenckii Hagens
13 a. T4 finely punctate with microsculptured apical depression [fig. 104], gonostylus [fig. 185, 186], 7–10 mm. . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . reticulatus Thomson
13 b. T4 coarsely punctate with smooth, shiny apical depression. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14
14 a. Flagellomere 10 (forelast flagellomere) a little longer than wide [fig. 107]; T4 with well visible interspaces among punctures
[fig. 105], gonostylus [figs. 141, 142], 8–11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . alternatus Smith
14 b. Flagellomere 10 about 1.5x longer than wide [fig. 108]; T4 riddle-like punctate with indistinct interspaces [fig. 106],
gonostylus [figs. 143, 144], 8–11 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . crassanus Warncke
15 a. The whole body (including legs and abdomen) black; hypoepimeral area smooth and shiny [fig. cf. 35]; head behind eyes
nearly as wide as compound eye, gonostylus [figs. 203, 204], 4.5–5.5 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . niger Hagens
15 b. The body always black and red, if only black, then tarsi reddish/yellow and hypoepimeral area punctate or sculptured . . . . 16
16 a. T2 in basal half densely and coarsely punctate; apical part of T1 coarsely punctate [fig. 109]; gonostylus narrow, loaflike [figs.
187, 188] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
16 b. T2 sparsely punctate; apical part of T1 shiny with only few punctures [fig. 110]; frons with distinct interspaces among
punctures; gonostylus of another shape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
17 a. Right angle between cubital and anal veins on hindwing [fig. cf. 44]; T4 dark in the middle, T5 completely dark, gonostylus
[figs. 187, 188], 8–12 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . rubicundus Hagens
17 b. Acute angle between the cubital and anal veins on hindwing; T4 all, T5 reddish on sides; T1 often with basal dark spot [fig.
111], gonostylus [figs. 189, 190], 6–9 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ruficrus Erichson
18 a. Felt-like pubescence on flagellomeres 8 to 10 short, up to 1/3 of the length of flagellomere [fig. 115]; T2 coarsely and densely
punctate [fig. 112]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
18 b. Felt-like pubescence on flagellomeres 8 to 10 long, at least 1/2 of the length of flagellomere [fig. 114]; T2 sparsely punctate
[fig. 113] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .20
19 a. Flagellomere 1 as wide as long [fig. 116]; gonostylus elongated and prominent [figs. 201, 202], 5–6 mm. ..dusmeti Blüthgen
19 b. Flagellomere 1 distinctly wider than long [fig. 115]; gonostylus short [figs. 183, 184], 4.5–7 mm . . . . . puncticeps Thomson
20 a. Head distinctly wider than thorax, with whitish long pubescence; metatarsus dark; felt-like pubescence on flagellomeres more
than ½ of the flagellomere [fig. 114]; wings hyaline, gonostylus [figs. 177, 178], 7–11 mm . . . . . . . . . . pellucidus Thomson
20 b. Head not distincly wider than thorax and without long hair; metatarsus reddish; felt-like pubescence on flagellomeres about ½
of the flagellomere; wings slightly infumate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .21
21 a. Frons and vertex coarsely punctate, interspaces dull; scutellum densely punctate, interspaces smaller than punctures [fig. 117];
head distinctly wider than long, gonostylus [figs. 151, 152], 6–9 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . ephippius (Linnaeus)
21 b. Frons and vertex sparsely and finely punctate, interspaces shiny; scutellum with few large punctures and large interspaces
among them [fig. 118]; head about as long as wide, gonostylus [figs. 171, 172], 3.5–5 mm. . . . . . . . . . . . . . longulus Hagens
22 a. Pronotal projections round [fig. cf. 48]; abdomen as wide as thorax or wider; face finely rugose, dull.. . . . . . . . . . . . . . . . . .23
22 b. Pronotal projections sharp [fig. cf. 49] (less distinct in some geoffrellus); abdomen narrower than thorax; face punctate, shiny
or dull . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24
23 a. Tibiae and tarsi reddish; felt-like pubescence about or more than half of the length of the flagellomere [fig. 119]; end of
gonostylus curved [figs. 159, 160] , 5–7 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . hyalinatus Hagens
23 b. Tibiae and tarsi dark or black; felt-like pubescence as ¼ of the maximal length of flagellomere [fig. 120]; end of gonostylus
straight, triangular [figs. 153, 154], 6–9 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ferruginatus Hagens
24 a. Felt-like pubescence of middle flagellomeres not longer than half of flagellomere [fig. 121]. . . . . . . . . . . . . . . . . . . . . . . . . 25
Zootaxa 3311 © 2012 Magnolia Press · 7
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
24 b. Felt-like pubescence of middle flagellomeres distinctly longer than half of flagellomere, in some species cover all length of
flagellomeres 4 to 11[figs. 122, 123] . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .27
25 a. Head a little longer than wide [fig. 124]; scutum with coarse but shallow punctures [fig. 129]; T1 and T2 with very fine, but
numerous punctures, gonostylus [figs. 181, 182], 5–8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pseudofasciatus Blüthgen
25 b. Head round or wider than long [fig. 125]; T1 and T2 with fine and strong punctures mixed . . . . . . . . . . . . . . . . . . . . . . . . . .26
26 a. T1 with a few variable punctures [fig. 127]; gonostylus with large membranous part; apex of gonobase on ventral side with
patch of long hair [figs. 145, 146], 5–7 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . crassus Thomson
26 b. T1 with numerous well-developed punctures [fig. 128]; gonostylus with small triangular membranous part; apex of gonobase
on ventral side with indistinct short hair [figs. 147, 148], 6–8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . croaticus Meyer
27 a. Frons with apressed white pubescence bellow as well as above antennal sockets; flagellomeres 2 to 11 long, more than 1.5x
longer than wide, covered by felt-like pubescence to the end of each segment [fig. 132] . . . . . . . . . . . . . . . . . . . . . . . . . . . .28
27 b. Frons with apressed white pubescence bellow antennal sockets only; at least flagellomere 2 different . . . . . . . . . . . . . . . . . .29
28 a. Scutum densely punctate [fig. 131]; first abdominal tergites in most cases red, gonostylus [figs. 161, 162], 5–8 mm . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . intermedius Blüthgen
28 b. Scutum sparsely punctate [fig. 130]; abdomen usually dark to black, gonostylus [figs. 179, 180], 5–7 mm . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . pinguiculus Pérez
29 a. T1 dorsally and T2 and T3 basally finely and very sparsely punctate, punctures nearly indistinct [fig. 133]; gonostylus short
[figs. 155, 156], 4–6 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . geoffrellus (Kirby)
29 b. T2 and T3 basally densely punctate, punctures well-defined [fig. 134]; gonostylus short or long . . . . . . . . . . . . . . . . . . . . . 30
30 a. Interspaces among punctures on area between eye and lateral ocellus, as well as in front of the middle ocellus densely
microsculptured, dull [fig. 135]; T1 densely punctate [fig. 136]; scutum densely punctate, punctures deep and coarse [fig. 138].
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .31
30 b. Interspaces among punctures on area between eye and lateral ocellus as well as in front of the middle ocellus shiny [fig. 137];
T1 sparsely punctate; scutum coarsely, but sparsely punctate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .32
31 a. Lower gena densely hairy, hair uniform in length, surface distinctly ridged and microsculptured, dull; vertex wide and long,
upper gena behind eyes narrowing slowly, distance between hind ocellus and posterior margin of vertex almost as long as
distance between hind ocelli, vertex behind oceli flat; front ocellus always larger than hind ocelli; area of felt like pubescence
on flagellomeres strongly impressed [fig. 135], gonostylus [figs. 199, 200], 5–7 mm . . . . . . . . . . . . . . . zangherii Noskiewicz
31 b. Lower gena sparsely hairy, with intermixed long and short hair, surface distinctly microsculptured, but shiny; vertex narower
and shorter, upper gena behind eyes strongly narrowing, distance between hind ocellus and posterior margin of vertex
distinctly shorter than distance between hind ocelli, vertex behind ocelli decreasing; felt like pubescence on flagelomers very
variable, gonostylus [figs. 147, 148], 6–8 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . croaticus Meyer
32 a. Membranous part of gonostylus enormous, trapezoidal; apex of gonobase on ventral side angulated, with patch of long hair
[figs. 167, 168]; T1 very finely and sparsely punctate [fig. 134]; frons densely punctate, interspaces among punctures very
small [fig. 126], 4–6 mm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . miniatus Hagens
32 b. Membranous part of gonostylus smaller; apex of gonobase on dorsal side with much shorter hair; T1 distinctly and densely
punctate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .33
33 a. Membranous part of gonostylus large, slightly, but distinctly emarginated on its inner margin; apex of gonobase on ventral side
angulated, with patch of short, but distinct erected hair [fig. 173, 174]; head wider than long (distinct only in large specimens);
felt-like pubescence on flagellomeres allways very long, from flagellomere 4 to 11 covers all length of flagellomere [fig. 123],
3–6 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . nomioidis Pesenko
33 b. Membranous part of gonostylus small, triangular, on its inner margin straight; apex of gonobase on ventral side straight,
without distinct erected hair [figs. 165, 166]; head round (distinct only in large specimens); felt-like pubescence of
flagellomeres 3 to 6 very variable, but usually does not reach end of flagellomere, separated by elevated curled edge on distal
margin in most central-European specimens [fig. 122], 3–5 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . marginatus Hagens
List of species
Sphecodes albilabris (Fabricius, 1793)
Described as: Nomada albilabris Fabricius, 1793: 349.
Synonyms: Dichroa fuscipennis Germar, 1819: Tab. 18.
Sphecodes latreillii Wesmael, 1835: 285.
Sphecodes rubripes Spinola, 1838: 512.
Sphecodes africanus Lepeletier, 1841: 541.
Sphecodes nigripes Lepeletier, 1841: 542.
Sphecodes rugosus Smith, 1848: 2209–2210.
Sphecodes nodicornis Gistel, 1857: 554.
Sphecodes fuscipennis var. basalis Dalla Torre, 1877: 185.
Sabulicola cirsii Verhoeff, 1890: 329–331.
Sphecodes grandis Meyer, 1922: 173.
BOGUSCH & STRAKA
8 · Zootaxa 3311 © 2012
Magnolia Press
Sphecodes rufipennis Cockerell, 1931: 348.
Sphecodes atrescens Cockerell, 1931: 350.
Distribution. South and central Europe, in the north reaching Denmark, Estonia, southern Finland and Sweden,
absent from Norway and on the British Isles; the distribution area extends to Asia (Westrich 1989, Warncke 1992,
Lönnell & Cederberg 2007).
Biology. Species found in sandy sites, sand dunes, river banks and semideserts. Usually, it occurs in warmer
regions and is locally highly abundant. Colletes cunicularius (Linnaeus) is the main host of this specialized cuckoo
bee (e.g. Blüthgen 1934, Westrich 1989). This cuckoo bee has only one generation during the year. It is highly
probable that females survive for long time and fly in early summer, when Colletes is still not available. In this sit-
uation S. albilabris can accept another (secondary) host. Old worn females were observed entering nests of Melit-
turga clavicornis (Latreille) and larvae of S. albilabris were excavated and described from its nest by Rozen
(1965). Blüthgen (1934) also stated Halictus quadricinctus (Fabricius) as unconfirmed host. We observed females
of S. albilabris invading nests of this species at the exactly same situation as was Rozen’s (1965) observation of
this species parasitizing Melliturga. Our record was made at Stroupe
č
Natural Monument in the Czech Republic in
July 2011. However, suggested parasitic activity of the second generation, e.g. in nests of Dasypoda hirtipes (Fabr-
icius), seems to be highly unlikely (A. P
ř
idal, pers. comm.). Adult females and males emerge in July, mate, females
find shelter and overwinter. There is no place for parasitic activity before overwintering.
Taxonomic note. Warncke (1992) regarded S. rubripes Spinola, 1838 (with synonyms S. africanus Lepeletier,
1841, Sphecodes rufipennis Cockerell, 1931, Sphecodes atrescens Cockerell, 1931) as subspecies of S. albilabris.
However, the difference among S. albilabris, S. rubripes and S. africanus is major (Blüthgen, 1924) and not only in
coloration as Warncke (1992) suggested. They also differ in phenology: males of S. rubripes and S. africanus were
collected in spring (April) and males of S. albilabris in summer. These forms can be valid species, but it requires
further taxonomic study.
Sphecodes alternatus Smith, 1853
Descibed as: Sphecodes alternatus Smith, 1853: 36.
Synonyms: Sphecodes gibbus var. similis subvar. scariosus Sichel, 1865: 444.
Sphecodes punctiventris Hagens, 1882: 219.
Sphecodes gracilior Morawitz, 1894: 78.
Sphecodes antigae Tournier, 1901: 258–260.
Sphecodes reticulatus var. algeriensis Alfken, 1914: 195.
Sphecodes alternatus lindbergi Pittioni, 1950: 61–62.
Subspecies: S. a. gracilior Morawitz, 1894: 78.
S. a. algeriensis Alfken, 1914: 195.
Distribution. South of Europe, reaching central Europe to Switzerland, Slovakia, Hungary, and Ukraine, in the
east through Turkey to Turkestan, present also in North Africa (Warncke 1992, Amiet et al. 1999).
Biology. Species of warm sites, especially sand dunes; quite common in various warm biotopes in southern
Europe. In central Europe it is rare. Abundant in extensive sandy sites such as military training areas or larger sand
quarries. Hosts unconfirmed. It was observed in association with Halictus compressus (Walckener) in Hungary
(own observations); H. langobardicus Blüthgen is another likely host. Blüthgen (1934) recorded H. patellatus F.
Morawitz as the likely host.
Sphecodes crassanus Warncke, 1992
Described as: Sphecodes crassanus Warncke, 1992: 26–27.
No synonyms.
Distribution. South Europe—Spain, France (Warncke 1992), Switzerland (Amiet et al. 1999), Algeria.
Biology. Poorly known species of open warm habitats with unknown host. The biology is probably similar to
S. alternatus and hosts can be expected within the medium sized species of Halictus.
Zootaxa 3311 © 2012 Magnolia Press · 9
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
Sphecodes crassus Thomson, 1870
Described as: Sphecodes crassus Thomson, 1870: 100.
Synonyms: Sphecodes variegatus Hagens, 1874: 40–41.
Sphecodes divisus Hagens, 1882 (nec Kirby, 1802): 223.
Sphecodes valesianus Frey-Gessner, 1903: 100.
Distribution. Europe north to 64° N, Sweden, Finland and Norway, present on British Isles; Turkey, Iran, and
North Africa (Warncke 1992).
Biology. Common species that usually occurs in semi-open biotopes with shrubs, forest-steppes, steppes, and
forest margins. In central Europe widespread, although except for warm sites it is never numerous. This species is a
nest cleptoparasite of smaller species of Halictidae. Sick et al. (1994) experimentally proved Stoeckhert’s (1933)
finding of Lasioglossum pauxillum (Schenck) as a host, Westrich (1989) and Vegter (1993) mentioned also L.
punctatissimum (Schenck) as a confirmed host. Other similar species are likely to be hosts as well: L.
quadrinotatulum (Schenck) recorded as likely host by Alfken (1912), L. nitidiusculum (Kirby) by Stoeckhert
(1933) and L. prasinum (Smith) by Vegter (1993). The hosts can be different in parts of the distributional area, as L.
prasinum is rare in central but quite common in north-western Europe.
Sphecodes cristatus Hagens, 1882
Described as: Sphecodes cristatus Hagens, 1882: 218.
No synonyms.
Distribution. Most of Europe north to 54°N, present in Sweden, absent on British Isles and Balkan Peninsula.
Known from Turkey, Tajikistan and Mongolia (Westrich 1989, Warncke 1992).
Biology. Very rare species of sand dunes and larger sandy sites, in most countries known only from few
records. Blüthgen (1934) mentioned Lasioglossum nigripes (Lepeletier) as the likely host, but this species differs
slightly in ecology. Westrich (1989) mentioned Halictus confusus Smith and H. subauratus (Rossi) as likely hosts,
without any evidence. In our observations, the most likely host is H. leucaheneus Ebmer and probably also H.
seladonius (Fabricius). We collected few S. cristatus at places with common occurence of H. leucaheneus in
Slovakia, Hungary and Mongolia and the association seems to be possible. However, the most likely host and the
cleptoparasite are both rare in central Europe, and to prove the host association is difficult.
Sphecodes croaticus Meyer, 1922
Described as: Sphecodes croaticus Meyer, 1922: 171.
No synonyms.
Distribution. South Europe, reaching central Europe, known also from Cyprus, Russia, Turkey and Morocco
(Warncke 1992). Due the confusion with S. pseudofasciatus and S. zangherii, the distribution can be incorrect. We
examined material from: Austria, Bulgaria, Croatia, Czech Republic, France, Germany, Greece, Hungary, Italy,
Portugal, Slovakia, Spain and Turkey.
Biology. Rare species of warm open habitats, usually steppes and warm sites, either on loess walls, or sandy
dunes; very rare and only in the warmest parts of central Europe. Lasioglossum interruptum (Panzer) is the only
published host (Blüthgen 1934) and was also observed by us and M. Herrmann (pers. comm.).
Note. Burger & Reum (2004) and Burger et al. (2006) stated that specimens from central Europe are not S.
croaticus but instead S. zangherii. Examination of specimens from various parts of Europe and also specimens
identified by F. Burger showed that these specimens are in fact S. croaticus. Thus, S. croaticus is thermophilous and
widespread in central Europe, contrary to the much rarer S. zangherii.
BOGUSCH & STRAKA
10 · Zootaxa 3311 © 2012
Magnolia Press
Sphecodes dusmeti Blüthgen, 1924
Described as: Sphecodes dusmeti Blüthgen, 1924: 470–472.
No synonyms.
Distribution. Southern Europe (Spain, France, Greece, Switzerland), North Africa (Morocco, Algeria) and Turkey
(Warncke 1992).
Biology. Unknown.
Sphecodes ephippius (Linné, 1767)
Described as: Sphex ephippia Linné, 1767: 944.
Synonyms: Apis minimus Harris, 1776 (nec Poda, 1761): Taf. 39, Fig. 21.
Apis obscura Geoffroy, 1785 (nec Linné, 1774, nec Müller, 1776): 447–448.
Apis rufescens Geoffroy, 1785 (nec Gmelin, 1790): 447.
Apis rufescens Gmelin, 1790 (nec Geoffroy, 1785): 2790.
Apis labiata Fabricius, 1793: 342.
Melitta divisa Kirby, 1802: 49–50.
Andrena minuta Fabricius, 1804: 327.
Sphecodes similis Wesmael, 1835: 283.
Sphecodes zablocki Blüthgen, 1923c: 188.
Distribution. Europe to 62°N, present in Britain and Scandinavia, North Africa (Morocco to Egypt), Turkey, the
distribution area continues through Asia to Japan (Warncke 1992).
Biology. Dominant species in central and common in south Europe, where it is not as common probably due to
competition with other species of the genus (S. gibbus, S. nomioidis). No biotope specialization; occurs nearly
everywhere. Generalist with 18 known hosts, Bogusch et al. (2006) recorded the following species as confirmed
hosts with complete references: Halictus tumulorum (Linnaeus), Lasioglossum laticeps (Schenck), L. leucozonium
(Schrank), L. malachurum (Kirby), L. pauxillum and L. quadrinotatulum. In addition Andrena barbilabris (Kirby),
A. flavipes Panzer, A. chrysopyga Schenck, A. labialis (Kirby), A. minutula (Kirby), A. wilkella (Kirby), Halictus
maculatus Smith, H. rubicundus (Christ), Lasioglossum fratellum (Péréz), L. lativentre (Schenck), and L. prasinum
were published as likely, unconfirmed hosts. The same authors also proved the individual specialization of the
females of this species. It could be one of the reasons why is this species so numerous and widespread. Addition-
ally, Andrena argentata Smith was observed as a host of S. ephippius.
Sphecodes ferruginatus Hagens, 1882
Described as: Sphecodes ferruginatus Hagens, 1882: 221.
Synonym: Sphecodes rufescens var. alpestris Frey-Gessner, 1903: 107.
Distribution. Europe to 66°N, present in Scandinavia and Britain, Turkey, absent from North Africa (Warncke
1992).
Biology. Species of midlands, usually bound to grasslands or sunny slopes, but not in warm areas. In central
Europe widespread but rare and uncommon. Lasioglossum fulvicorne (Kirby) is the only known documented host
(Stoeckhert, 1933). Similar species such as L. laticeps and L. pauxillum have been published as likely hosts of this
species (Stoeckhert 1933, Westrich 1989). Surveys on localities with these species present did not support this as
the females of S. ferruginatus were invading only nests of L. fulvicorne.
Sphecodes geoffrellus (Kirby, 1802)
Described as: Melitta geoffrella Kirby, 1802: 45–46.
Synonyms: Sphecodes affinis Hagens, 1882: 224.
Zootaxa 3311 © 2012 Magnolia Press · 11
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
Sphecodes fasciatus Hagens, 1882: 224.
Sphecodes rimalis Pérez, 1903: CCXXI.
Sphecodes impunctatus Meyer, 1922: 171–172.
Subspecies: Sphecodes geoffrellus atlasa Warncke, 1992: 36.
Sphecodes geoffrellus hakkariensis Warncke, 1992: 36.
Distribution. Most of Europe, present in Britain and Scandinavia, North Africa (Morocco, Tunisia), Turkey, Near
East (Warncke 1992). Very numerous in southern Europe.
Biology. Common species both in warmer and cooler regions, with no strict biotope preferences. In central
Europe it is widespread and numerous, although recently sparser, probably due to the spread of similar species
S. miniatus and S. nomioidis. Generalist parasitizing several hosts: Westrich (1989) has reported the following
hosts as confirmed Lasioglossum leucopus (Kirby), L. morio (Fabricius) and L. nitidiusculum, Bogusch (2003) L.
pauxillum. Likely hosts are the following: L. fratellum published by Field (1996), L. rufitarse (Zetterstedt) by
Neumeyer & Obrist (2000), L. sexstrigatum (Schenck) by Vegter (1993) and L. marginellum (Schenck) by Westrich
(2006). It is possible that this species invades nests of various smaller Lasioglossum species, and the females are
individually specialized, as Bogusch et al. (2006) have confirmed in S. ephippius and S. monilicornis.
Sphecodes gibbus (Linnaeus, 1758)
Described as: Sphex gibba Linnaeus, 1758: 571.
Synonyms: Apis glabra Füessly, 1775: 51.
Andrena ferruginea Olivier, 1789, nomen novum for Nomada gibba Fabricius, 1775 (nec Linnaeus, 1758): 139.
Apis gibbosa Christ, 1791, nomen novum for Nomada gibba Fabricius, 1775 (nec Linnaeus, 1758): 177.
Melitta picea Kirby, 1802: 48–49.
Melitta sphecoides Kirby, 1802: 46–47.
Andrena austriaca Fabricius, 1804: 325.
Dichroa analis Illiger, 1806, nomen novum for Nomada gibba Fabricius, 1775 (nec Linnaeus, 1758): 48.
Sphecodes apicatus Smith, 1853: 36–37.
Sphecodes nigripennis Morawitz, 1876: 257.
Sphecodes sutor Nurse, 1903: 538.
Sphecodes gibbus var. rufispinosus Meyer, 1920: 113.
Sphecodes gibbus var. turcestanicus Meyer, 1920: 113.
Sphecodes nippon Meyer, 1922: 171.
Sphecodes castilianus Blüthgen, 1924: 473–475.
Sphecodes lustrans Cockerell, 1931: 411.
Sphecodes pergibbus Blüthgen, 1938: 50.
Distribution. Nearly all of Europe (north to 63° N), the distribution area goes far into Asian mainland; also present
in North Africa (Warncke 1992).
Biology. Species of warm open or bushy habitats, more common in southern Europe (one of the most
numerous Sphecodes species). In central Europe it shows affinity to warm biotopes, and is one of the most common
species of Sphecodes. This species is very variable in appearance and also in host spectrum. Main hosts are larger
or medium sized species of the genus Halictus. Westrich (1989) recorded H. quadricinctus, H. rubicundus, and H.
sexcinctus (Fabricius), Bogusch (2003) confirmed H. simplex Blüthgen as a host and presented Lasioglossum
malachurum as a likely host. Other records of likely hosts are difficult to accept: Andrena vaga Panzer published
by Möschler (1938) and Colletes cunicularius published by Riemann (1987). We observed a nesting site of A. vaga
for several years, but no attempt to enter the nest by this cuckoo bee was observed, even though the activity of S.
gibbus was observed few times. Westrich (1989) also put Halictus maculatus as likely host, which can be accepted.
The list of hosts is probably larger than recently known and individual specialization to various hosts across the
distribution area is likely.
Sphecodes hyalinatus Hagens, 1882
Described as: Sphecodes hyalinatus Hagens, 1882: 222.
No synonyms.
BOGUSCH & STRAKA
12 · Zootaxa 3311 © 2012
Magnolia Press
Distribution. Most of the Europe to 68°N, rare in south Europe; absent from Turkey and North Africa (Westrich
1989, Warncke 1992).
Biology. Species of sunny sites in cooler regions, usually in rocky steppes on non-basic substrate. In central
Europe local and usually rare. Two known hosts are Lasioglossum fratellum confirmed by Field (1996) and L. ful-
vicorne first mentioned by Stoeckhert (1933) as the likely host and since confirmed by Westrich (1989). The biol-
ogy of L. fratellum and S. hyalinatus was studied by Heide (1992) and Field (1996). The abundance of this species
is usually bound to the abundance of hosts. We have observed association of S. hyalinatus with both of the men-
tioned hosts.
Sphecodes intermedius Blüthgen, 1923
Described as: Sphecodes intermedius Blüthgen, 1923a: 500–502.
Synonym: Sphecodes lactipennis Meyer, 1925: 7–8.
Distribution. Very rare species, known from the Czech Republic, Slovakia, Hungary, Spain, Russia, and Turkey,
Caucasus, Kazakhstan, Turkmenistan, Tajikistan, and Algeria (Warncke 1992). In all countries only one to several
records are known, and most of them are very old. Recently, the species was recorded at Brat
č
ice in south Moravia
(Czech Republic) in 2011.
Biology. Biology is poorly known; probably a species of warm and dry biotopes. Hosts unknown, in our opin-
ion Halictus kessleri Bramson can be the host of this species. This bee is present at sand walls in Brat
č
ice, where
we found two specimens in 2011.
Sphecodes longulus Hagens, 1882
Described as: Sphecodes longulus Hagens, 1882: 226.
Synonyms: Sphecodes longulus var. epidus Hagens, 1882: 226.
Sphecodes nitidulus Hagens, 1882: 226.
Distribution. Europe, north to south Finland, Sweden and Denmark, south England, eastwards to Tajikistan
(Westrich 1989, Warncke 1992, Sörensson et al. 2009, Madsen & Calabuig 2011).
Biology. Rare species of sandy sites, uncommon in the localities in most of European countries. This species is
nest cleptoparasite of small species of Lasioglossum; L. minutissimum (Kirby) is a confirmed host (Alfken 1912).
Vegter (1993) brought evidence of two additional hosts: L. leucopus and L. zonulum (Smith), which is possible.
Other likely hosts are the following: L. lucidulum (Schenck) and L. sexstrigatum published by Vegter (1993) and L.
morio mentioned by Westrich (1989). During our observations nests of L. punctatissimum and L. semilucens
(Alfken) were also invaded by females of this species.
Sphecodes majalis Pérez, 1903
Described as: Sphecodes majalis Pérez, 1903: CCXIX.
Synonyms: Sphecodes gracilior Pérez, 1903: CCXIX.
Sphecodes problematicus Schulz, 1906: 235.
Sphecodes barbarus Blüthgen, 1923a: 497.
Distribution. Spain, France, Belgium, Germany, Switzerland, Austria, Italy, Slovenia, Czech Republic, Slovakia,
Hungary, Russia, Algeria (Warncke 1992).
Biology. Very rare species of steppes and sunny sites, bound to its host Lasioglossum pallens (Brullé). Her-
rmann et al. (2003) described the biology both of the host and the parasite. The cleptoparasite as its host occur in
spring (both males and females), and are recordable only during few days or weeks from March to May. Thus it
seems to be rarer than it is.
Zootaxa 3311 © 2012 Magnolia Press · 13
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
Sphecodes marginatus Hagens, 1882
Described as: Sphecodes marginatus Hagens, 1882: 223.
Synonyms: Sphecodes atratus Hagens, 1882: 224.
Sphecodes nigritulus Hagens, 1882: 225.
Sphecodes biskrensis Pérez, 1903: CCXXI.
Subspecies: S. m. larochei Warncke, 1992: 34.
S. m. creticus Warncke, 1992: 35.
Distribution. Unclear because of past confusion with S. nomioidis and reported by Warncke (1992) as S. marginatus
biskrensis. We examined wide material of the S. marginatus group and this species seems to be atlanto-mediterranean
(absent from eastern Europe, Scandinavia and British Isles). Occurrence of S. marginatus was confirmed in the
following countries: Belgium, France, Germany (up to north), Italy (Sicilia), Morocco, Portugal, Spain, Tunisia and
as subspecies in Canary Islands and Crete, also published for Denmark (Madsen & Calabuig 2011).
Biology. Species of warm open biotopes, usually sandy sites. In central Europe bound to heath and sandpits
and other open sandy biotopes. Hosts unknown, Vegter (1993) mentioned Dufourea minuta Lepeletier as likely
host. However, the biology of both species is quite different, so we cannot accept it. In our opinion, small species of
Lasioglossum occurring in sandy localities will be the hosts, eg. L. lucidulum, L. punctatissimum, L. semilucens, L.
sexstrigatum or L. sabulosum, but this needs more study.
Sphecodes miniatus Hagens, 1882
Described as: Sphecodes miniatus Hagens, 1882: 223.
Synonyms: Sphecodes dimidiatus Hagens, 1882: 224.
Sphecodes murithianus Frey-Gessner, 1903: 100.
Sphecodes pilicornis Meyer, 1922: 170.
Distribution. Most of Europe (north to south Sweden), present in southern England; rare in southern Europe
(confirmed only in Pelopponnese), absent from North Africa (Warncke 1992).
Biology. Common species especially of warmer and sandy sites. Westrich (1989) mentioned Lasioglossum
nitidiusculum as confirmed host and Lasioglossum morio as likely host, Vegter (1993) L. sexstrigatum and
Bogusch (2003) L. politum Schenck. We also observed females of this species in association of L. pauxillum and L.
punctatissimum. However, females of S. miniatus are similar to S. marginatus, and other small Sphecodes, so study
of the specialization in the field is difficult.
Sphecodes monilicornis (Kirby, 1802)
Described as: Melitta monilicornis Kirby, 1802: 47–48.
Synonyms: Sphecodes maculatus Lepeletier, 1841: 545.
Sphecodes subquadratus Smith, 1845: 1014.
Sphecodes gibbus var. subquadratus subvar. incertus Sichel, 1865: 420.
Sphecodes gibbus var. ephippium subvar. nigrescens Sichel, 1865: 427.
Sphecodes gibbus var. ephippium subvar. testaceipes Sichel, 1865: 428.
Sphecodes gibbus var. ephippium subvar. rufipes Sichel, 1865: 428.
Sphecodes gibbus var. subquadratus subvar. dubius Sichel, 1865: 419.
Sphecodes ruficrus Dalla Torre, 1896 (nec Erichson, 1835), nomen novum for S. rufipes Sichel, 1865: 9.
Sphecodes hanuman Nurse, 1903: 538–539.
Sphecodes smyrnaensis Meyer, 1920: 116.
Sphecodes caucasicus Meyer, 1920: 124.
Sphecodes quadratus Meyer, 1920: 129.
Sphecodes cephalotes Meyer, 1920: 129.
Sphecodes monilicornis var. nigerrima Blüthgen, 1927: 41.
Sphecodes quadratus cephalotiformes Pittioni, 1950: 62.
Subspecies: Sphecodes monilicornis quadratus Meyer, 1920: 129.
BOGUSCH & STRAKA
14 · Zootaxa 3311 © 2012
Magnolia Press
Sphecodes monilicornis cephalotes Meyer, 1920: 129.
Sphecodes monilicornis berberus Warncke, 1992: 22.
Distribution. Most of Europe, north to 64°N, in the east through Asia to Japan, present in North Africa (Westrich
1989, Warncke 1992).
Biology. Common species, especially in warm habitats. Occurs mainly on steppes and sandy sites. It seems to
be generalist with many hosts. Bogusch et al. (2006) summarized all known hosts of this species. Confirmed hosts
are the following: Halictus rubicundus, Lasioglossum albipes (Fabricius), L. calceatum (Scopoli), L. leucozonium,
L. quadrinotatulum and L. zonulum. L. malachurum is another confirmed host by our research. Vegter (1993)
brought data of possible parasitation in nests of L. prasinum, Bogusch (2003) of Andrena flavipes, Halictus
maculatus, H. tumulorum, Lasioglossum laticeps, L. pauxillum and L. villosulum (Kirby). Thus the number of hosts
of this species is high and the females are individually specialized. Most of the hosts are primitively eusocial and
females of S. monilicornis lay eggs either into their nests within the solitary phase in spring or summer worker
phase. Females of S. monilicornis can kill the workers of a nest and than lay eggs in all cells including those sealed
and those not fully supplied (A. P
ř
idal, pers. comm.).
Sphecodes niger Hagens, 1874
Described as: Sphecodes niger Hagens, 1874: 43.
Synonym: Sphecodes niger var. holomelaena Blüthgen, 1949: 79.
Distribution. Europe from north-east Spain to Ukraine, Turkey, absent from Scandinavia and North Africa
(Westrich 1989, Warncke 1992).
Biology. Species of warm biotopes, where can be common. Usually, it is not very abundant but widespread.
The only confirmed host is Lasioglossum morio; Alfken (1912) also thought L. lucidulum to be a host. However,
this species occurs mainly in sandy sites, where S. niger is uncommon.
Sphecodes nomioidis Pesenko, 1979
Described as: Sphecodes nomioidis Pesenko, 1979: 136.
No synonyms.
Distribution. Unclear due to confusion with S. marginatus. We examined wide material of S. marginatus group
and S. nomioidis seems to be ponto-mediterranean (absent from west Europe and North Africa) species. The occur-
rence of S. nomioidis was confirmed in the following countries: Austria, Bulgaria, Czech Republic, Greece, Hun-
gary, Jordan, Romania, Slovakia and Turkey.
Biology. Poorly known, because of confusion with S. marginatus. It occurs on warm sandy and loess biotopes
in central Europe. The species is expanding northwards recently and getting more common in central Europe.
Taxonomic note. Sphecodes marginatus biskrensis Pérez, 1903 sensu Warncke (1992) was characterized by
strongly elongated felt-like pubescence on antennomeres. This character seems to be very variable and present
commonly in S. marginatus as well as in S. nomioidis. The type of S. biskrensis comes from North Africa, where S.
nomioidis does not occur. For this reason, S. nomioidis remains the only available name for species from eastern
Europe. The application of the name needs to be confirmed by study of the type material as well as examination of
a wider material from the Ukraine and other eastern European localities.
Sphecodes olivieri Lepeletier, 1825
Described as: S. olivieri Lepeletier, 1825: 448.
Synonyms: Sphecodes collaris Spinola, 1843: 137–139.
Sphecodes hispanicus var. abyssinicus Sichel, 1865: 447.
Sphecodes ruficornis Sichel, 1865: 440.
Zootaxa 3311 © 2012 Magnolia Press · 15
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
Sphecodes punctulatus Sichel, 1865: 443–444.
Sphecodes subpunctulatus Sichel, 1865: 445–446.
Sphecodes rufithorax Morawitz, 1876: 255–256.
Sphecodes verticalis Hagens, 1882: 219.
Sphecodes desertus Nurse, 1903: 540–541.
Sphecodes chionospilus Cockerell, 1911: 217–218.
Sphecodes chionospilus var. sanguinatus Cockerell, 1911: 217.
Sphecodes tenuis Meyer, 1920: 121–122.
Sphecodes olivieri var. niveatus Meyer, 1925: 4.
Distribution: Southern Europe from Spain to Turkey, North Africa (Morocco to Egypt), Israel, Caucasus
(Warncke 1992) and Iran.
Biology. Southern species occurring in warm semidesert habitats. The hosts of this rare species are not well
known. Blüthgen (1934) mentioned Lasioglossum aegyptiellum (Strand) and L. vagans (Smith) as likely hosts.
Sphecodes pellucidus Smith, 1845
Described as: Sphecodes pellucidus Smith, 1845: 1014.
Synonyms: Sphecodes pilifrons Thomson, 1870: 99.
Sphecodes brevicornis Hagens, 1874: 39–40.
Sphecodes volatilis Smith, 1879: 26.
Sphecodes pellucidus var. algirus Alfken, 1914: 195.
Sphecodes pellucidus var. hypridus Blüthgen, 1925: 516.
Sphecodes pellucidus var. niveipennis Meyer, 1925: 7.
Distribution. Europe north to 66°N, present in southern Sweden and Finland, east parts of Asia, North Africa
(Westrich 1989, Warncke 1992).
Biology. Species of sandy sites, sand dunes and semidesert biotopes. Locally common. Alfken (1913) con-
firmed host Andrena barbilabris. Witt (1992) described the biology of this host and the cleptoparasite. Schönitzer
& Klinksik (1990) confirmed also A. nycthemera Imhoff; Sick et al. (1994) Lasioglossum leucozonium as hosts of
S. pellucidus. Other bee species of sandy biotopes were usually published as likely hosts: Andrena argentata Smith,
A. bicolor Fabricius, A. humilis Imhoff, A. ventralis Imhoff and A. wilkella; Vegter (1993) mentioned also Lasio-
glossum prasinum as likely host. This species was probably common in the localities investigated by this author,
because it was considered a likely host of many Sphecodes species. In our survey, S. pellucidus was usually
observed in association with A. barbilabris, and S. ephippius from the same localities invaded nests of A. argen-
tata. In our opinion, S. pellucidus is specialized and other hosts are unlikely. To the contrary, we could not reject
similar behaviour in S. albilabris, where females try to attack nests of other species.
Sphecodes pinguiculus Pérez, 1903
Described as: Sphecodes pinguiculus Pérez, 1903: CCXX.
Synonyms: Sphecodes sareptensis Meyer, 1922: 170–171.
Sphecodes excellens Meyer, 1922: 170.
Sphecodes punctatissimus Meyer, 1922: 172.
Sphecodes hungaricus Blüthgen, 1923a: 498–499.
Sphecodes coelebs Blüthgen, 1923a: 505–506.
Sphecodes consobrinus Blüthgen, 1923a: 507–508.
Sphecodes persicus Blüthgen, 1925: 509–511.
Sphecodes capverdensis Pesenko & La Roche, 2002: 72, 203 in Pauly et al. (2002), syn. nov.
Taxonomy. We studied material of this species from Mongolia, Hungary, the Mediterranean region and Cape Verde
islands. Variablity of this species is not large and for this reason we propose synonymization of S. capverdensis
with S. pinguiculus.
BOGUSCH & STRAKA
16 · Zootaxa 3311 © 2012
Magnolia Press
Distribution. Spain, Hungary, Slovakia, Italy (Sicilia), Russia, North Africa, Turkey, Mongolia (Warncke
1992), Cape Verde.
Biology. Species of warm, usually sandy habitats. This species is widely distributed in desert and semidesert
regions, where it can be quite abundant. This species is almost surely a parasite of Halictus lucidipennis Smith of
Cape Verde Islands. It is the only available common bee at the sites with occurence of S. pinguiculus and of course,
close association of both species has been observed in Cape Verde. Possibly, H. lucidipennis is main host in most
areas of its occurrence, because this Halictus is common species in desert and semidesert areas from Mongolia,
through Mediterranean region to Cape Verde. We suggest H. smaragdulus Vachal to be possible host for central
European region, where similar H. lucidipennis does not occur. Nobile & Turrisi (2004) described seven new spe-
cies related to S. pinguiculus. However, the characters are poor and probably relate to the way of preservation or
collecting rather than true structural differences. Recently, the type material was studied (Schwarz & Gusenleitner
2012) and the described species are not related to S. pinguiculus but S. miniatus group.
Sphecodes pseudofasciatus Blüthgen, 1925
Described as: Sphecodes pseudofasciatus Blüthgen, 1925: 473.
No synonyms.
Distribution. Distribution is poorly known, because of former incorrect synonymization of this species under S.
croaticus by Warncke (1992). We examined material from: Austria, Czech Republic, France, Hungary, Italy, Portu-
gal, Romania, Russia, Slovakia, Spain, Switzerland, Morocco, Turkey and Ukraine.
Biology. This species is collected in low numbers and only few records are known from each country. Host is
unknown.
Sphecodes puncticeps Thomson, 1870
Described as: Sphecodes puncticeps Thomson, 1870: 99–100.
Synonyms: Sphecodes bituberculatus Pérez, 1903: CCXX–CCXXI.
Sphecodes opacifrons Pérez, 1903: CCXIX–CCXX.
Sphecodes puncticeps var. cretanus Strand, 1921: 305.
Distribution. Europe north to south Finland and Sweden, Asia, North Africa to Egypt, Israel (Westrich 1989,
Warncke 1992), Iran.
Biology. Species of warm sandy biotopes, in south Europe quite common, in central Europe local but in sandy
localities usually abundant. The only confirmed host is Lasioglossum villosulum, published by Alfken (1913). Bis-
choff (1927) put L. brevicorne (Schenck) as likely host. We confirm both hosts and have observed S. puncticeps
also in an association with L. politum and L. sabulosum (Warncke), but with no evidence.
Sphecodes reticulatus Thomson, 1870
Described as: Sphecodes reticulatus Thomson, 1870: 98.
Synonym: Sphecodes distinguendus Hagens, 1874: 38–39.
Distribution. Europe north to 62°N, present in south Sweden and Finland, British Isles, east to Turkestan. In the
Mediterranean very rare, known from Bulgaria, Greece and Turkey (Westrich 1989, Warncke 1992).
Biology. Species of various biotopes, usually in sandy sites, heathlands and sandpits. Present also in other open
or semi-open habitats, as steppes, shrubby sites, etc. Rare in the whole distribution area. Andrena barbilabris was
mentioned as confirmed host by Stoeckhert (1933) and Blüthgen (1934); A. argentata, A. wilkella, Lasioglossum
prasinum and L. leucozonium cedri Ebmer as likely hosts (Blüthgen 1934, Vegter 1993, M. Schwarz, pers. comm.).
Zootaxa 3311 © 2012 Magnolia Press · 17
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
Sphecodes rubicundus Hagens, 1875
Described as: Sphecodes rubicundus Hagens, 1875: 318.
Synonym: Sphecodes rubicundus altisilesiacus Torka, 1926: 126–127.
Distribution. Europe north to 56° N, England, eastern to Ural and Armenia. Present in Turkey (Westrich 1989,
Warncke 1992).
Biology. Very rare species. Older findings are from open habitats: usually sandy sites, sanddunes and steppes.
Sowa & Mostowska (1978) described the biology of Andrena labialis (Kirby) and confirmed it as a host of this spe-
cies. Blüthgen (1934) also put A. labialis as confirmed and A. nigroaenea (Kirby) as likely host of S. rubicundus.
Torka (1925) brought data on possible parasitization in nests of A. agilissima (Scopoli). Currently, the mentioned
hosts occur mostly in different biotopes from the cleptoparasite.
Sphecodes ruficrus (Erichson, 1835)
Described as: Dichroa ruficrus Erichson, 1835: 101–102.
Synonyms: Sphecodes hispanicus Wesmael, 1835: 285–286.
Sphecodes rufipes Smith, 1853: 37.
Sphecodes gibbus var. tunetanus Gribodo, 1894: 293–294.
Sphecodes atrohirtus Pérez, 1903 (nomen novum for S. hispanicus Wesmael, 1836 sensu Hagens, 1882): CCXIX.
Distribution. South and western Europe: Spain, France, Switzerland, Italy, Germany, Hungary, Russia. This spe-
cies extends its distribution area to north-east (Westrich 1989, Warncke 1992).
Biology. Species of sandy sites, in southern Europe common in such localities. Herrmann (2006) described the
biology and hosts. The main host is Andrena humilis, also related species A. livens Pérez and A. taraxaci Giraud are
possible hosts of this species. We have studied this species in Spain in association with A. livens. Westrich (1989)
reported Andrena decipiens Schenck as a host, but with no further details.
Sphecodes rufiventris (Panzer, 1798)
Described as: Tiphia rufiventris Panzer, 1798: 4.
Synonyms: Sphecodes subovalis Schenck, 1853: 223–224.
Sphecodes brevis Hagens, 1875: 317–318.
Sphecodes singularis Meyer, 1920: 130.
Sphecodes combinatus Blüthgen, 1927: 37–39.
Sphecodes tadschicus Blüthgen, 1935: 366.
Sphecodes subovalis austrinus Erlandsson, 1979: 123.
Sphecodes subovalis austrinus var. balcanicus Erlandsson, 1979: 123.
Sphecodes rufiventris hethiticus Warncke, 1992: 28.
Distribution. Europe up to 57°N, absent from Scandinavia and Britain, in the east the distribution area goes to Ural
River, present in North Africa: Morocco, Algeria (Warncke 1992).
Biology. Species of warmer open or shrubby habitats (forest steppes), common on loess walls. In central
Europe quite rare, more abundant only at xerothermes. The only known host is Halictus maculatus, recorded by
Stoeckhert (1933) and Blüthgen (1934). We have only observed S. rufiventris in association with this species.
Sphecodes scabricollis Wesmael, 1835
Described as: Sphecodes scabricollis Wesmael, 1835: 287.
Synonym: Sphecodes perversus Ritsema, 1879: 56–57.
BOGUSCH & STRAKA
18 · Zootaxa 3311 © 2012
Magnolia Press
Distribution. Europe north to southern Finland, present in England, southward absent from the Balkan Peninsula
and Turkey (Westrich 1989, Warncke 1992).
Biology. Very rare species of sandy wetlands, in most countries endangered or extinct. The distribution is much
weaker than in the past. The host is Lasioglossum zonulum, recorded in the literature (Blüthgen 1934, Yates 2002),
the first author also recorded Halictus compressus, H. quadricinctus and Lasioglossum prasinum as likely hosts.
More recently, parasitation only in nests of L. zonulum was observed (M. Herrmann, pers. comm.).
Sphecodes schenckii Hagens, 1882
Described as: Sphecodes schenckii Hagens, 1882: 217–218.
Synonyms: Sphecodes sulcicollis Pérez, 1903: CCVIII.
Subspecies: Sphecodes schenckii caspicus Meyer, 1920: 113–114.
Distribution. North-east Spain, France, Switzerland, Italy, Hungary, Austria, Slovakia, Germany, Russia, Turkey
and Iran (Warncke 1992).
Biology. Rare species of warm localities, in most of the countries only few records. Lasioglossum discum
(Smith) is supposed to be a host due to the similar distribution area, size and appearance (Blüthgen 1934). Groz-
dani
ć
(1971) confirmed this species as host of S. schenckii. In northern Switzerland and in Germany L. discum is
missing, so there has to be one or more other hosts, probably Halictus simplex (M. Herrmann, pers. comm. and
pers. obs.).
Sphecodes spinulosus Hagens, 1875
Described as: Sphecodes spinulosus Hagens, 1875: 317.
No synonyms.
Distribution. Europe north to 56°N, present in England, Turkey, south-east Russia, and the distribution area
extends probably east to Asia. Present in North Africa (Warncke 1992, Edwards & Telfer 2001).
Biology. Rare species of warm biotopes, usually steppes. In central Europe only few records in recent years.
The only known host is Lasioglossum xanthopus (Kirby) recorded by Stoeckhert (1933) and Blüthgen (1934).
Sphecodes zangherii Noskiewicz, 1931
Described as: Sphecodes zangherii Noskiewicz, 1931: 139.
No synonyms.
Distribution. The distribution is poorly known due the taxonomical problems with this species in the past.
Warncke (1992) suggested wide distribution in south and central Europe, from France to Turkey and north to Ger-
many and Czech Republic. However, we examined large part of Warncke’s material and found only few specimens
from the Alps in France and Switzerland, north and central Italy and mountains in Greece and Turkey. All other
specimens of S. zangherii determined by Warncke were missidentified S. croaticus.
Biology. Rare species of warmer localities in mountains, but sometimes also in low altitudes. Details of its
biology are unknown.
Zootaxa 3311 © 2012 Magnolia Press · 19
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
TABLE 1. List of hosts of Sphecodes Latreille species of central Europe. Likely hosts are marked by asterisk.
Species Hosts
Sphecodes albilabris (Fabricius, 1793) Colletes cunicularius (Linnaeus)
Halictus quadricinctus (Fabricius)
Melitturga clavicornis (Latreille)
*Dasypoda hirtipes (Fabricius)
Sphecodes alternatus Smith, 1853 *Halictus compressus (Walckener)
*Halictus langobardicus Blüthgen
*Halictus patellatus F. Morawitz
Sphecodes crassanus Warncke, 1992 hosts unknown
Sphecodes crassus Thomson, 1870 Lasioglossum pauxillum (Schenck)
Lasioglossum punctatissimum (Schenck)
*Lasioglossum quadrinotatulum (Schenck)
*Lasioglossum nitidiusculum (Kirby)
*Lasioglossum prasinum (Smith)
Sphecodes cristatus Hagens, 1882 *Halictus confusus Smith
*Halictus leucaheneus Ebmer
*Halictus seladonius (Fabricius)
*Halictus subauratus (Rossi)
*Lasioglossum nigripes (Lepeletier)
Sphecodes croaticus Meyer, 1922 *Lasioglossum interruptum (Panzer)
Sphecodes dusmeti Blüthgen, 1924 hosts unknown
Sphecodes ephippius (Linné, 1767) Andrena argentata Smith
Halictus tumulorum (Linnaeus)
Lasioglossum laticeps (Schenck)
Lasioglossum leucozonium (Schrank)
Lasioglossum malachurum (Kirby)
Lasioglossum pauxillum Schenck
Lasioglossum quadrinotatulum (Schenck)
*Andrena barbilabris (Kirby)
*Andrena flavipes Panzer
*Andrena chrysopyga Schenck
*Andrena labialis (Kirby)
*Andrena minutula (Kirby)
*Andrena wilkella (Kirby)
*Halictus maculatus Smith
*Halictus rubicundus (Christ)
continued next page
BOGUSCH & STRAKA
20 · Zootaxa 3311 © 2012
Magnolia Press
TABLE 1. (continued)
Species Hosts
Sphecodes ephippius (Linné, 1767) *Lasioglossum fratellum (Péréz)
*Lasioglossum lativentre (Schenck)
*Lasioglossum prasinum (Smith)
Sphecodes ferruginatus Hagens, 1882 Lasioglossum fulvicorne (Kirby)
*Lasioglossum laticeps (Schenck)
*Lasioglossum pauxillum Schenck
Sphecodes geoffrellus (Kirby, 1802) Lasioglossum leucopus (Kirby)
Lasioglossum morio (Fabricius)
Lasioglossum nitidiusculum (Kirby)
Lasioglossum pauxillum Schenck
*Lasioglossum fratellum (Péréz)
*Lasioglossum marginellum (Schenck)
*Lasioglossum rufitarse (Zetterstedt)
*Lasioglossum sexstrigatum (Schenck)
Sphecodes gibbus (Linnaeus, 1758) Halictus quadricinctus (Fabricius)
Halictus rubicundus (Christ)
Halictus sexcinctus (Fabricius)
Halictus simplex Blüthgen
*Andrena vaga Panzer
*Colletes cunicularius (Linnaeus)
*Halictus maculatus Smith
*Lasioglossum malachurum (Kirby)
Sphecodes hyalinatus Hagens, 1882 Lasioglossum fratellum (Péréz)
Lasioglossum fulvicorne (Kirby)
Sphecodes intermedius Blüthgen, 1923 *Halictus kessleri Bramson
Sphecodes longulus Hagens, 1882 Lasioglossum leucopus (Kirby)
Lasioglossum minutissimum (Kirby)
Lasioglossum punctatissimum (Schenck)
Lasioglossum semilucens (Alfken)
Lasioglossum zonulum (Smith)
*Lasioglossum lucidulum (Schenck)
*Lasioglossum sexstrigatum
Sphecodes majalis Pérez, 1903 Lasioglossum pallens (Brullé)
Sphecodes marginatus Hagens, 1882 *Dufourea minuta Lepeletier
continued next page
Zootaxa 3311 © 2012 Magnolia Press · 21
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
TABLE 1. (continued)
Species Hosts
Sphecodes marginatus Hagens, 1882 * Lasioglossum lucidulum (Schenck)
*Lasioglossum punctatissimum (Schenck)
*Lasioglossum sabulosum (Warncke)
*Lasioglossum semilucens (Alfken)
*Lasioglossum sexstrigatum (Schenck)
Sphecodes miniatus Hagens, 1882 Lasioglossum nitidiusculum (Kirby)
*Lasioglossum morio (Fabricius)
*Lasioglossum pauxillum Schenck
*Lasioglossum politum Schenck
*Lasioglossum punctatissimum (Schenck)
*Lasioglossum sexstrigatum (Schenck)
Sphecodes monilicornis (Kirby, 1802) Halictus rubicundus (Christ)
Lasioglossum albipes (Fabricius)
Lasioglossum calceatum (Scopoli)
Lasioglossum leucozonium (Schrank)
Lasioglossum malachurum (Kirby)
Lasioglossum quadrinotatulum (Schenck)
Lasioglossum zonulum (Smith)
*Andrena flavipes Panzer
*Halictus maculatus Smith
*Halictus tumulorum (Linnaeus)
*Lasioglossum laticeps (Schenck)
*Lasioglossum pauxillum Schenck
*Lasioglossum prasinum (Smith)
*Lasioglossum villosulum (Kirby)
Sphecodes niger Hagens, 1874 Lasioglossum morio (Fabricius)
*Lasioglossum lucidulum (Schenck)
Sphecodes nomioidis Pesenko, 1979 hosts unknown
Sphecodes olivieri Lepeletier, 1825 *Lasioglossum aegyptiellum (Strand)
*Lasioglossum vagans (Smith)
Sphecodes pellucidus Smith, 1845 Andrena barbilabris (Kirby)
Andrena nycthemera Imhoff
Lasioglossum leucozonium (Schrank)
*Andrena argentata Smith
*Andrena bicolor Fabricius
*Andrena humilis Imhoff
continued next page
BOGUSCH & STRAKA
22 · Zootaxa 3311 © 2012
Magnolia Press
TABLE 1. (continued)
Species Hosts
Sphecodes pellucidus Smith, 1845 *Andrena ventralis Imhoff
*Andrena wilkella (Kirby)
*Lasioglossum prasinum (Smith)
Sphecodes pinguiculus Pérez, 1903 Halictus lucidipennis Smith
*Halictus smaragdulus Vachal
Sphecodes pseudofasciatus Blüthgen, 1925 hosts unknown
Sphecodes puncticeps Thomson, 1870 Lasioglossum villosulum (Kirby)
*Lasioglossum brevicorne (Schenck
*Lasioglossum politum Schenck
*Lasioglossum sabulosum (Warncke)
Sphecodes reticulatus Thomson, 1870 Andrena barbilabris (Kirby)
*Andrena argentata Smith
*Andrena wilkella (Kirby)
*Lasioglossum leucozonium cedri Ebmer
*Lasioglossum prasinum (Smith)
Sphecodes rubicundus Hagens, 1875 Andrena labialis (Kirby)
*Andrena agilissima (Scopoli)
*Andrena nigroaenea (Kirby)
Sphecodes ruficrus (Erichson, 1835) Andrena humilis Imhoff
Andrena livens Pérez
*Andrena decipiens Schenck
*Andrena taraxaci Giraud
Sphecodes rufiventris (Panzer, 1798) Halictus maculatus Smith
Sphecodes scabricollis Wesmael, 1835 Lasioglossum zonulum (Smith)
*Halictus compressus (Walckener)
*Halictus quadricinctus (Fabricius)
*Lasioglossum prasinum (Smith)
Sphecodes schenckii Hagens, 1882 Lasioglossum discum (Smith)
*Halictus simplex Blüthgen
Sphecodes spinulosus Hagens, 1875 Lasioglossum xanthopus (Kirby)
Sphecodes zangherii Noskiewicz, 1931 hosts unknown
Zootaxa 3311 © 2012 Magnolia Press · 23
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
FIGURES 1–9. Females. 1—Sphecodes majalis, head frontal view, clypeus; 2—S. spinulosus, vertex with ridge; 3—S. spinu-
losus, antenna; 4—S. schenckii, eight wing hammuli; 5—S. puncticeps, five wing hammuli; 6—S. crassanus, hindwing vena-
tion; 7—S. hyalinatus, hindwing venation; 8—S. scabricolis, occiput with ridge; 9—S. scabricolis, head and scutum, dorsal
view.
BOGUSCH & STRAKA
24 · Zootaxa 3311 © 2012
Magnolia Press
FIGURES 10–17. Females. 10—Sphecodes olivieri, dorsal view; 11—S. cristatus, vertex with longitudinal ridge; 12—S.
monilicornis, head, frontal view; 13—S. albilabris, scutum; 14—S. monilicornis, mesopleura; 15—S. rufiventris, mesopleura;
16—S. reticulatus, head, lateral view; 17—S. gibbus, head, lateral view.
Zootaxa 3311 © 2012 Magnolia Press · 25
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
FIGURES 18–27. Females. 18—Sphecodes reticulatus, tergum 4; 19—S. gibbus, tergum 4; 20—S. alternatus, tergum 4; 21—
S. crassanus, tergum 4; 22—S. alternatus, head, frontal view; 23—S. crassanus, head, frontal view; 24—S. gibbus, scutum;
25—S. schenckii, scutum; 26—S. gibbus, scapus; 27—S. schenckii, scapus.
BOGUSCH & STRAKA
26 · Zootaxa 3311 © 2012
Magnolia Press
FIGURES 28–38. Females. 28—Sphecodes pinguiculus, metasoma, sculpture of tergites 1–3; 29—S. pinguiculus, scutum;
30—S. intermedius, scutum; 31—S. pinguiculus, head, frontal view; 32—S. puncticeps, head, frontal view; 33—S. crassus,
head, frontal view; 34—S. longulus, head, frontal view; 35—S. niger, mesopleura; 36—S. niger, propodeum, dorsal view; 37—
S. puncticeps, tergum 3; 38—S. longulus, tergum 3.
Zootaxa 3311 © 2012 Magnolia Press · 27
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
FIGURES 39–47. Females. 39—Sphecodes pellucidus, pygidium; 40—S. crassus, pygidium; 41—S. crassus, sculpture of
clypeus; 42—S. ephippius, sculpture of clypeus; 43—S. ruficrus, all body, dorsal view; 44—S. rubicundus, hind wing venation;
45—S. rubicundus, apical margin of tergum 1; 46—S. ephippius, apical margin of tergum 1; 47—S. pellucidus, head, frontal
view.
BOGUSCH & STRAKA
28 · Zootaxa 3311 © 2012
Magnolia Press
FIGURES 48–54. Females. 48—Sphecodes ferruginatus, pronotum, oblique view; 49—S. geoffrellus, pronotum, oblique
view; 50—S. hyalinatus, propodeum, dorsal view; 51—S. ferruginatus, thorax, ventral view; 52—S. hyalinatus, thorax, ventral
view; 53—S. ferruginatus, head, frontal view; 54—S. hyalinatus, head, frontal view.
Zootaxa 3311 © 2012 Magnolia Press · 29
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
FIGURES 55–61. Females. 55—Sphecodes pseudofasciatus, head, frontal view; 56—S. pseudofasciatus, head, dorsal view;
57—S. zangherii, head, frontal view; 58—S. zangherii, head, dorsal view; 59—S. croaticus, head, frontal view; 60—S. croati-
cus, head, dorsal view; 61—S. croaticus, sculpture of tergite 2.
BOGUSCH & STRAKA
30 · Zootaxa 3311 © 2012
Magnolia Press
FIGURES 62–73. Females. 62—Sphecodes dusmeti, head, frontal view; 63—S. crassus, head, frontal view; 64—S. geoffrel-
lus, head, frontal view; 65—S. crassus, dorsal part of mesopleura; 66—S. geoffrellus, dorsal part of mesopleura; 67—S. cras-
sus, sculpture of tergum 2 and 3; 68—S. geoffrellus, sculpture of tergum 2 and 3; 69—S. crassus, antenna; 70—S. geoffrellus,
antenna; 71—S. miniatus, antenna; 72—S. miniatus, head, lateral view; 73—S. nomioidis, head, lateral view.
Zootaxa 3311 © 2012 Magnolia Press · 31
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
FIGURES 74–86. Females. 74—Sphecodes miniatus, head, frontal view; 75—S. marginatus, head, frontal view; 76—S. nomi-
oidis, head, frontal view; 77—S. miniatus, scutum; 78—S. marginatus, scutum; 79—S. nomioidis, scutum; 80—S. miniatus,
sculpture of tergum 2; 81—S. marginatus, sculpture of tergum 2; 82—S. nomioidis, sculpture of tergum 2; 83, 84—S. miniatus,
pygidium, oblique and dorsal view; 85—S. marginatus, pygidium, dorsal view; 86—S. nomioidis, pygidium, dorsal view.
BOGUSCH & STRAKA
32 · Zootaxa 3311 © 2012
Magnolia Press
FIGURES 87–96. Males. 87—Sphecodes spinulosus, hind tibia, lateral view; 88—S. monilicornis, hind tibia, lateral view;
89—S. rufiventris, mesopleura; 90—S. rufiventris, antenna; 91—S. cristatus, scutum; 92—S. olivieri, scutum; 93—S. scabri-
collis, head and scutum, dorsal view; 94—S. cristatus, antenna; 95—S. olivieri, antenna; 96—S. albilabris, scutum.
Zootaxa 3311 © 2012 Magnolia Press · 33
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
FIGURES 97–108. Males. 97—Sphecodes monilicornis, head, frontal view; 98—S. reticulatus, head, frontal view; 99—S.
monilicornis, sculpture of tergites 1–3; 100—S. gibbus, head, sculpture of vertex; 101—S. reticulatus, head, sculpture of ver-
tex; 102—S. schenckii, head, sculpture of vertex and antenna; 103—S. gibbus, antenna; 104—S. reticulatus, sculpture of tergite
4; 105—S. alternatus, sculpture of tergite 4; 106—S. crassanus, sculpture of tergite 4; 107—S. alternatus, terminal flagellom-
eres; 108—S. crassanus, terminal flagellomeres.
BOGUSCH & STRAKA
34 · Zootaxa 3311 © 2012
Magnolia Press
FIGURES 109–118. Males. 109—Sphecodes ruficrus, sculpture of tergites 1–2; 110—S. ephippius, sculpture of tergites 1–2;
111—S. ruficrus, all body, dorsal view; 112—S. puncticeps, sculpture of tergites 1–2; 113—S. pellucidus, sculpture of tergites
1–2; 114—S. pellucidus, antenna; 115—S. puncticeps, antenna; 116—S. dusmeti, antenna; 117—S. ephippius, sculpture of
scutellum; 118—S. longulus, sculpture of scutellum.
Zootaxa 3311 © 2012 Magnolia Press · 35
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
FIGURES 119–132. Males. 119—Sphecodes hyalinatus, antenna; 120—S. ferruginatus, antenna; 121—S. pseudofasciatus,
antenna; 122—S. marginatus, antenna; 123—S. nomioidis, antenna; 124—S. pseudofasciatus, head, frontal view; 125—S. cras-
sus, head, frontal view; 126—S. miniatus, head, frontal view; 127—S. crassus, sculpture of tergum 1; 128—S. croaticus, sculp-
ture of tergum 1; 129—S. pseudofasciatus, scutum; 130—S. pinguiculus, scutum; 131—S. intermedius, scutum; 132—S.
pinguiculus, head, frontal view.
BOGUSCH & STRAKA
36 · Zootaxa 3311 © 2012
Magnolia Press
FIGURES 133–138. Males. 133—Sphecodes geoffrellus, sculpture of tergites 1–3; 134—S. miniatus, sculpture of tergites 1–3;
135—S. zangherii, head, dorsal oblique view; 136—S. zangherii, sculpture of tergum 1; 137—S. miniatus, head, dorsal oblique
view; 138—S. zangherii, scutum.
Zootaxa 3311 © 2012 Magnolia Press · 37
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
FIGURES 139–174. Male genitalia, dorsal and lateral views. 139, 140—Sphecodes albilabris; 141, 142—S. alternatus;
143, 144—S. crassanus; 145, 146—S. crassus; 147, 148—S. croaticus; 149, 150—S. cristatus; 151, 152—S. ephippius; 153,
154—S. ferruginatus; 155, 156—S. geoffrellus; 157, 158—S. gibbus; 159, 160—S. hyalinatus; 161, 162—S. intermedius; 163,
164—S. majalis; 165, 166—S. marginatus; 167, 168—S. miniatus; 169, 170—S. monilicornis; 171, 172—S. longulus; 173,
174—S. nomioidis.
BOGUSCH & STRAKA
38 · Zootaxa 3311 © 2012
Magnolia Press
FIGURES 175–204. Male genitalia, dorsal and lateral views. 175, 176—Sphecodes olivieri; 177, 178—S. pellucidus; 179,
180—S. pinguiculus; 181, 182—S. pseudofasciatus; 183, 184—S. puncticeps; 185, 186—S. reticulatus; 187, 188—S. rubicun-
dus; 189, 190—S. ruficrus; 191, 192—S. rufiventris; 193, 194—S. scabricollis; 195, 196—S. schenckii; 197, 198—S. spinulo-
sus; 199, 200—S. zangherii; 201, 202—S. dusmeti; 203, 204—S. niger.
Acknowledgement
We would like to thank colleagues and collectors who helped us with the material. Special thanks belong to Maxi-
milian Schwarz (Ansfelden, Austria) and Christian Schmid-Egger (Berlin, Germany), who helped us with material
for this study and acccess to their collections. Additional thanks to Mike Herrmann (Konstanz, Germany) and one
Zootaxa 3311 © 2012 Magnolia Press · 39
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
anonymous referee for valuable comments to the article. Institutional support to JS was given by Ministry of Edu-
cation of the Czech Republic, project no. MSM0021620828; grant support to PB and JS was given by Czech Sci-
ence Foundation, project no. 206/09/0522.
References
Alfken, J.D. (1912) Die Bienenfauna von Ostpreussen. Schrift des physisch-ökonomischen Vereins in Königsberg, 53, 114–182.
Alfken, J.D. (1913) Die Bienenfauna von Bremen. Abhandlungen des naturwissenschaflichen Vereins Bremen, 22, 1–
220.Alfken, J. D. (1914) Beitrag zur Kenntnis der Bienenfauna von Algerien. Mémoires de la Société Entomologique Bel-
gique, 22, 185–237.
Amiet, F., Müller, A. & Neumeyer, R. (1999) Hymenoptera Apidae. 2. Teil: Gattungen Colletes, Dufourea, Hylaeus, Nomia,
Nomioides, Rhophitoides, Rophites, Sphecodes, Systropha. Fauna Helvetica 4. Centre Suisse de Cartographie de la Faune
& Schweitzerische Entomologische Gesellschaft, Luzern, 219 pp.
Bischoff, H. (1927) Biologie der Hymenopteren. J. Springer, Berlin, 598 pp.
Blüthgen, P. (1923a) Beiträge zur Systematik der Bienengattung Sphecodes Latr. Deutsche Entomologische Zeitschrift, 1923,
441–514.
Blüthgen, P. (1923b) Zur Biologie der Bienengattung Sphecodes Latr. (Hym.). Zeitschrift für Wissenschaftlicher Insekten-Bio-
logie, 18, 19–23.Blüthgen, P. (1923c) Sphecodes zablockii nov. spec. Weibchen und Sph. croaticus Meyer Weibchen
(Hym. Apidae). Polskie Pismo Entomologiczne, 2, 188–190.
Blüthgen, P. (1924) Beiträge zur Systematik der Bienengattung Sphecodes Latr. II. Deutsche entomologische Zeitschrift, 1924,
457–516.
Blüthgen, P. (1927) Beiträge zur Systematik der Bienengattung Sphecodes Latr. III. Zoologische Jahrbuch, Abteilung für Sys-
tematik, 53, 23–112.
Blüthgen, P. (1934) Die Wirte der Paläarktischen Sphecodes-Arten. Zeitschrift Wissenschaftlicher Insekten-Biologie, 27, 33–42,
61–66.Blüthgen, P. (1935) Halictus, Nomioides und Sphecodes, pp. 360–367. In: Popov, V.V. (ed.) Beiträge zur Bienen-
fauna von Tadzhikistan. Trudy Tajikskoy Bazy Akademie Nauk SSSR, 5, 1–408.
Blüthgen, P. (1938) Neue Halictidi aus Zypern. Konowia, 16, 41–54.
Blüthgen, P. (1949) Neues oder Wissenswertes über mitteleuropäische Aculeaten und Goldwespen. Beiträge zur taxonomis-
chen Zoologie, 1, 77–100.
Bogusch, P. (2003) Biologie vybraných druh
ů
kleptoparazitických v
č
el (Hymenoptera: Apocrita, Apoidea). [Biology of selected
cuckoo bee species (Hymenoptera: Apocrita, Apoidea)]. Charles University in Prague, Faculty of Sciences, MSc. thesis,
unpublished, 111 pp. (in Czech).
Bogusch, P., Kratochvíl, L. & Straka, J. (2006) Generalist cuckoo bees are species specialist in an individual level
(Hymenoptera: Apoidea, Sphecodes). Behavioral Ecology and Sociobiology, 60, 422–429.
Bogusch, P., Straka, J. & Kment, P. (2007) Annotated checklist of the Aculeata (Hymenoptera) of the Czech Republic and Slo-
vakia. Komentovaný seznam žahadlových blanok
ř
ídlých (Hymenoptera: Aculeata)
Č
eské republiky a Slovenska. Acta
Entomologica Musei Nationalis Pragae, Supplementum 11, 1–300.
Burger, F., Meitzel, T. & Ruhnke, H. (2006) Aktuelles zur Bienenfauna (Hymenoptera: Apidae) Sachsen-Anhalts und
Deutschlands. Entomologische Nachrichten und Berichte, 50, 129–133.
Burger, F. & Reum, D. (2004) Dritter Nachtrag zur Bienenfauna Thüringens (Hymenoptera: Apidae). Check-Listen Thüringer
Insekten- und Spinnentiere, 12, 33–39.
Celary, W. (1991) Review of the cuckoo bees of the genus Sphecodes Latr. (Hymenoptera, Apoidea, Halictidae) in Poland. Pol-
ske pismo entomologiczne, 61, 1–20.
Christ, J.L. (1791) Naturgeschichte, Klassifikation und Nomenklatur der Insekten vom Bienen, Wespen und Ameisengeschlecht.
Herrmann, Frankfurt am Main, 535 pp.
Cockerell, T.D.A. (1911) Bees in the collection of the United States National Museum. 2. Proceedings of the United States
National Museum, 40, 241–264.
Cockerell, T.D.A. (1931) Descriptions and Records of Bees. 127. Annals and Magazine of Natural History, 7, 344–351.
Dalla Torre, K.W. (1877) Die Apiden Tirols. Zeitschrift des Ferdinandeums, 21, 159–196.
Dalla Torre, K.W. (1896) Catalogus Hymenopterorum hucusque descriptorum systematicus et synonymicus. Vol. X. Apidae
(Anthophila). Sumptibus Guilelmi Engelmann, Lipsiae, VIII + 643 pp.
Erichson, W.F. (1835) Beschreibung von 19 neuen Hymenopteren aus Andalusien, pp. 101–109. In: Waltl, J. (ed.) Reise durch
Tyrol, Oberitalien und Piemont nach dem südlichen Spanien, nebst einem Anhange zoologischen Inhalts. 2. Auflage. Ver-
lag der Pustetschen Buchhandlung (J. F. Winkler), Passau, 381 pp.
Erlandsson, S. (1979): Hymenoptera aculeata from the european part of the mediterranean countries. II. Acta Entomologica
Jugoslavica, 15, 111–130.
Fabricius, J.C. (1793) Entomologia systematica emendara et aucta Secundum classes, ordines, gen., spec., adjectis syn-
onymidis, locis, observationibus, descriptionibus. Vol. 2. Christian Gottlieb Proft, Hafniae, 8 + 518 pp.
Fabricius, J.C. (1804) Systema Piezatorum, secundum ordines, gen., spec., adjectis synonymidis, locis, observationibus,
descriptionibus. Carolum Reichard, Brunsvigae. xiv + 439 pp.
BOGUSCH & STRAKA
40 · Zootaxa 3311 © 2012
Magnolia Press
Field, J. (1996) Patterns of provisioning and iteroparity in a solitary halictinae bee, Lasioglossum (Evylaeus) fratellum (Perez),
with notes on L. (E.) calceatum (Scop.) and L. (E.) villosulum) (K.). Insectes Sociaux, 43, 167–182.Frey-Gessner, E.
(1903) Hyménoptères du Valais. Famille Apidae. Bulletin de la Murithienne, 30, 78–154.
Friese, H. (1888) Die Schmarotzerbienen und ihre Wirte. Zoologischer Jahrbuch der Systematik, 5, 751–860.Fuessly, J. C.
(1775) Verzeichnis der ihm bekannten Schweitzerischen Inseckten. J. C. Fuessly, Zürich, xii + 62 pp.
Geoffroy, E.L. (1785), pp. 233–544. In: Fourcroy, A. F. (ed.) Entomologia parisiensis. Sive Catalogus Insectorum quae in Agro
Parisiensis reperiuntur. Bd. 2. Parisiis, Aedibus Serpentineis, Pars prima, Paris, viii + 231 pp.
Germar, E.F. (1819) Fauna Insectorum Europae. Vol. 5. Kümmel, Halae, 25 Tables.
Gistel, J. (1857) Achthundert und zwanzig neue oder unbeschriebene wirbellose Thiere. Vacun a, 2, 513–606.
Gmelin, J.F. (1790) Caroli a Linné Systema naturae, sive regna tria naturae systematice proposita per classes, ordines, genera et
species. Tom. I. Pars IV, Beer, Lipsiae, pp 1517–2224.
Gribodo, G. (1894) Note Imenotterologiche. Nota 2. Nuove generi e n. specie di Imenotteri antofili ed osservazioni sopra
alcune species gia conosciute. Bollettino della Società Entomologica Italiana, 26, 262–314.
Grozdani
ć
, S. (1971) Biologische Untersuchungen an den Bienen. Deutsche Entomologische Zeitschrift, 18, 217–226.
Hagens, J. (1874) Ueber die Genitalien der männlichen Bienen, besonders der Gattung Sphecodes. Berliner entomologische
Zeitschrift, 1874, 25–43.
Hagens, J. (1875) Weitere Beiträge zur Kenntniss der deutschen Sphecodes-Arten. Deutsche Entomologische Zeitschrift, 19(2),
315–319.
Hagens, J. (1882) Ueber die männlichen Genitalien der Bienengattung Sphecodes. Deutsche Entomologische Zeitschrift, 26,
209–229.Harris, M. (1776) An exposition of English Insects, including the several classes of Neuroptera, Hymenoptera, &
Diptera, or bees, flies, & libellulae. M. Harris, London, pp. i–viii + 9–72.
Heide, A. (1992) Zur Bionomie von Lasioglossum (Evylaeus) fratellum (Pérez), einer Furchenbiene mit ungewöhnlich lan-
glebigen Weibchen (Hymenoptera, Halictinae). Drosera, 1992(2), 171–188.
Herrmann, M. (2006) Wirtsbindung und Habitate der Blutbiene Sphecodes ruficrus (Erichson 1835). Mitteilungen der entomol-
ogische Vereins in Stuttgart, 41, 55–60.
Herrmann, M., Burger, F., Müller, A. & Tischendorf, S. (2003) Verbreitung, Lebensraum und Biologie der Furchenbiene Lasio-
glossum pallens (Brullé 1832) und ihrer Kuckucksbiene Sphecodes majalis Pérez 1903 in Deutschland (Hymenoptera,
Apidae, Halictinae). Carolinea, 61, 133–144.
Illiger, K. (1806) William Kirbys Familien der bienenartigen Insekten mit Zusätzen, Nachweisungen und Bemerkungen. Maga-
zin für Insektenkunde, 5, 28–175.
Kirby, W. (1802) Monographia Apum Angliae. II. Ipswich, London, 388 pp.
Lepeletier, A. (1841) Histoire Naturelle des Insectes, Hyménoptères. Vol. 2. Roret, Paris, 680 pp.
Linnaeus, C. (1758) Systema Naturae. Editio 1. Laur. Salvii, Holmiae, 826 pp.
Linné, C. (1767) Systema Naturae. Editio 12. Laur. Salvii, Holmiae, 1327 pp.
Macek, J., Straka, J., Bogusch, P., Dvo
ř
ák, L., Bezd
ěč
ka, P. & Tyrner, P. (2010) Blanok
ř
ídlí
Č
eské Republiky. I. Žahadloví.
[Hymenoptera: Aculeata of the Czech Republic]. Academia, Praha, 524 pp. (in Czech).
Meyer, R. (1920) Apidae-Sphecodinae. Archiv für Naturgeschichte, 85A(1), 79–160.
Meyer, R. (1922) Nachtrag I zur Bienengattung Sphecodes Latr. Archiv für Naturgeschichte, 88A 8, 165–174.
Meyer, R. (1925) Zur Bienengattung Sphecodes. Archiv für Naturgeschichte, 90A(12), 1–12.
Morawitz, F. (1876) [Fauna of Bees of the Caucasus]. Trudy Russkago éntomologicheskago obshchestva, 12, 3–69 (in Russian).
Morawitz, F. (1894) [Fauna of Bees of the Turkmenistan]. Trudy Russkago éntomologicheskago obshchestva, 28, 1–76 (in Rus-
sian).
Möschler, A. (1938) Ein Beitrag zur Bienenfauna in Ostpreussen insbesondere der Kurischen Nehrung. Schrift der physischen
und ökonomischen Gesselschaft in Königsberg, 70, 243–288.
Neumeyer, R. & Obrist, M.K. (2000) Lasioglossum rufitarse (Zetterstedt, 1838) as a new host of Sphecodes geoffrellus (Kirby,
1802)? Bembix, 13, 9–10.
Nobile, V. & Campadelli, G. (1998) Il genere Sphecodes Latreille 1804 in Italia (Hymenoptera, Apoidea, Halictidae). Bolletino
dell'Istituto di Entomologia della Università di Bologna, 52, 85–103.
Nobile, V. & Turrisi, G.F. (2004) Contribution to the knowledge of Italian cleptoparasitic Bees. X. The genus Sphecodes
Latreille, 'pinguiculus Pérez' group, with description of new species (Hymenoptera, Apoidea, Halictidae). Entomofauna,
25(8), 117–132.
Noskiewicz, J. (1931) Sphecodes zangherii n. sp. (Hymenoptera, Apidae). Annales Musei Zoologici Polonici, 9, 139–145.
Nurse, C.G. (1903) New species of Indian Aculeate Hymenoptera. Annual Magazine of natural History, 11(7), 393–403, 511–
526, 528–549.
Olivier, A.G. (1789) Encyclopedie méthodique, dictionnaire des insectes. Bd. 4. Panckoucke, Paris, 331 pp.
Panzer, G.W.F. (1798) Fauna insectorum germanicae initia, oder Deutschlands Insecten. Heft 54. Felssecker, Nürnberg, 24 pp
+ 24 plates.
Pauly, A., Pesenko, Y.A. & La Roche, F. (2002) The Halictidae of the Cape Verde Islands (Hymenoptera Apoidea). Bulletin de
l’Institut Royal des sciences Naturelles de Belgique, 72, 201–2011.
Pérez, J. (1903) Espèces nouvelles de Mellifères (paléarctiques). Procès-verbaux des séances de la Société Linnéenne de Bor-
deaux, 58, LXXVIII–XCIII, CCVIII–CCXXXVI.
Pesenko, Y.A. (1979) A new species of cleptoparasitic bees of the genus Sphecodes Latr. from a nest of Nomioides minutissi-
Zootaxa 3311 © 2012 Magnolia Press · 41
REVIEW OF THE CUCKOO BEES OF CENTRAL EUROPE
mus (Rossi) (Hymenoptera, Halictidae). Entomological Reviews (Washingto n), 58 (4), 136.
Pittioni, B. (1950) Hymenoptera aculeata I. On the insectfauna of Cyprus. Results of the Expedition of 1939 by Harald, Hakan
and P. H. Lindberg. Commentationes Biologicae, Societas Scientiarum Fennica, 10, 1–94.
Riemann, H. (1987) Beitrag zur Stechimmenfauna niedersächsischer Sandgruben (Hymenoptera: Aculeata). Braunschweiger
Naturkundliche Schriften, 3, 213–242.
Rozen, J.G.Jr. (1965) The Biology and Immature Stages of Melitturga clavicornis (Latreille) and of Sphecodes albilabris
(Kirby) and the Recognition of the Oxaeidae at the Family Level (Hymenoptera, Apoidea). American Museum Novitates,
2224, 1–18.
Schenck, A. (1853) Nachtrag zu der Beschreibung nassauischer Bienenarten. Jahrbücher des Vereins für Naturkunde im Her-
zogthum Nassau, 9, 88–306.
Schönitzer, K. & Klinksik, C. (1990) The Ethology of the Solitary Bee Andrena nycthemera Imhoff, 1866 (Hymenoptera:
Apoidea). Entomofauna, 11(23/1), 377–427.
Schulz, W.A. (1906) Spolia Hymenopterologica. Druck und Verlag der Junfermannschen Buchhandlung, Paderborn, 355 pp.
Schwarz, M. & Gusenleitner, F. (2012) Zur Kenntnis der von Nobile V. & G.F. Turrisi (2004) aus Italien beschriebenen Sphe-
codes-Arten (Hymenoptera, Apidae). Entomofauna, 33(8), 73–80.
Schwarz, M., Gusenleitner, F., Westrich, P. & Dathe, H.H. (1996) Katalog der Bienen Österreichs, Deutschlands und der
Schweiz (Hymenoptera, Apidae). Entomofauna, 8, 1–398.
Sichel, J. (1865) Études hyménoptèrologiques. Annales de la Société Entomologique de France, 5(4), 331–492.
Sick, M., Ayasse, M., Tengö, J., Engels, W., Lübke, G. & Francke, W. (1994) Host-parasite relationship in six species of Sphe-
codes bees and their Halictid hosts: Nest intrusion, intranidal behavior, and Dufour’s gland volantiles (Hymenoptera: Hal-
ictidae). Journal of Insect Behavior, 7, 101–117.
Smith, F. (1845) Descriptions of the British species of Bees belonging to the genus Sphecodes of Latreille. Zoologist, 3, 1011–
1015.
Smith, F. (1848) Descriptions of the British species of Bees belonging to the genus Halictus of Latreille. Zoologist, 6, 2037–
2044, 2100–2108, 2167–2175.
Smith, F. (1853) Catalogue of Hymenopterous Insects in the Collection of the British Museum. Andrenidae and Apidae. Part 1.
Trustees, London, 197 pp.
Smith, F. (1879) Descriptions of new species of Hymenoptera in the collection of the British Museum. Trustees, London, 21 +
240 pp.
Sowa, S. & Mostowska, I. (1978) Przyczynek do biologii Andrena labialis Kb. (Hym., Apoidea) w województwie olsztynskim.
Polske pismo entomologiczne, 48, 439–444.
Spinola, M. (1838) Des Hyménoptères recueillis par M. Fischer pendant son voyade en Egypte, et communiqués par. M. le
Docteur Waltl. Annales de la Société Entomologique de France, 7, 512.
Spinola, M. (1843) Sur quelques Hyménoptères peu connus, recueillis en Espagne, pendant l’année 1842, par M. Victor Ghil-
iani, voyageur-naturaliste. Annales de la Société Entomologique de France, (2)1, 111–144.
Stöckhert, F.K. (1933) Die Bienen Frankens (Hym. Apid.). Deutsche entomologische Zeitschrift, Beiheft, 1932, 1–294.
Strand, E. (1921) Apidologisches, insbesondere über paläarktische Halictus-Arten, auf Grund von Material des Deutschen
Entomologischen Museums. Archiv für Naturgeschichte, 87A(3), 305–322.
Svensson, B.G. (1982) Blodbiet Sphecodes cristatus ny för Nordvästeuropa. [Cuckoo bee Sphecodes cristatus new for north-
west Europe]. Entomologisk Tidskrift, 103, 23–24. (in Swedish).
Šustera, O. (1959) Bestimmungstabelle der Tschechoslowakischen Arten der Bienengattung Sphecodes Latr. Acta Societatis
Entomologiceae
Č
echosloveniae, 56(2), 169–180.
Thomson, C.G. (1870) Opuscula entomologica. Bd. 2. Håkan Ohlson, Lund, 452 pp.
Torka, V. (1925) Seltene Bienen Oberschlesiens. International entomologischer Zeitschrift, 18, 255–257.Torka, V. (1926) Die
Bienenfauna Oberschlesiens. Internationale entomologische Zeitschrift, 20, 125–130.
Tournier, H. (1901) Descriptions de quelques Hyménoptères d’Europe et confins. Boletín de la Sociedad espanola Historia nat-
uralis, 1, 258.
Vegter, K. (1985) De tweede generatie van Andrena barbilabris in Drenthe. [Second generation of Andrena barbilabris in
Drenthe]. Entomologische Berichten (Amsterdam), 45, 3–5. (in Dutch).
Vegter, K. (1993) Gastheren van enige soorten Sphecodes in Drenthe (Hymenoptera: Apidae). [Hosts of some Sphecodes in
Drenthe]. Entomologische Berichten (Amsterdam), 53, 67–70. (in Dutch).
Verhoeff, C. (1890) Ein Beitrag zur deutschen Hymenopteren-Fauna. Entomologische Nachrichten Berlin, 16, 329.
Warncke, K. (1992) Die Westpaläarktischen Arten der Bienengattung Sphecodes Latr. (Hymenoptera, Apidae, Halictinae). Ber-
icht der Naturforschenden Gesellschaft Augsburg, 52, 9–64.
Wesmael, C. (1835): Observations sur les espèces du genre Sphécode. Bulletin de l’Académie des sciences, des lettres et des
beaux-arts de Belgique, 2, 279–287.
Westrich, P. (1989) Die Wildbienen Baden-Württembergs. Band. 1 und 2. Eugen Ulmer Verlag, Stuttgart, 972 pp.
Westrich, P. (2006) Beobachtungen an einem Nistplatz von Lasioglossum marginellum (Schenck, 1853) (Hym., Apidae). Ento-
mologische Nachrichten und Berichte, 50, 55–61.
Witt, R. (1992) Zur Bionomie der Sandbiene Andrena barbilabris (Kirby 1802) und ihrer Kuckucksbienen Nomada alboguttata
Herrich-Schäffer 1839 und Sphecodes pellucidus Smith 1845. Drosera, 1992(1), 47–81.
