European cuckoo bees of the tribe Dioxyini (Hymenoptera, Megachilidae): distribution, annotated checklist and identification key

Abstract

Altogether, ten species of cuckoo bees of the tribe Dioxyini have been recorded from Europe, with two species distributed widely in the continent while others are restricted in distribution to only one or several countries in southern Europe. These ten representatives are classified into five genera: Aglaoapis , Dioxys , Ensliniana , Metadioxys and Paradioxys . Dioxys atlanticus is reclassified from a subspecies to a valid species, and new occurrence records of this species are reported. New synonymy is established for Dioxys cinctus = D. montana syn. nov. The distribution, morphology, ecology and hosts of all species were reviewed from both published and unpublished sources. New red-list categories for each species were created according to the new records of occurrence. An identification key including all ten species and photographs of their whole bodies and main identification characteristics was prepared, and distribution maps for all species were created.

Full text

European cuckoo bees of the tribe Dioxyini
(Hymenoptera, Megachilidae): distribution,
annotated checklist and identification key
Petr Bogusch1
1Department of Biology, Faculty of Science, University of Hradec Králové, Rokitanského 62, CZ-500 03
Hradec Králové, Czech Republic
Corresponding author: Petr Bogusch (bogusch.petr@gmail.com)
Academic editor: Jack Ne  |  Received 14 April 2023  |  Accepted 27 June 2023  |  Published 25 July 2023
https://zoobank.org/16A4A165-5185-4C89-960D-614A74E6D394
Citation: Bogusch P (2023) European cuckoo bees of the tribe Dioxyini (Hymenoptera, Megachilidae): distribution,
annotated checklist and identication key. Journal of Hymenoptera Research 96: 599–628. https://doi.org/10.3897/
jhr.96.104957
Abstract
Altogether, ten species of cuckoo bees of the tribe Dioxyini have been recorded from Europe, with two spe-
cies distributed widely in the continent while others are restricted in distribution to only one or several coun-
tries in southern Europe. ese ten representatives are classied into ve genera: Aglaoapis, Dioxys, Enslini-
ana, Metadioxys and Paradioxys. Dioxys atlanticus is reclassied from a subspecies to a valid species, and new
occurrence records of this species are reported. New synonymy is established for Dioxys cinctus = D. montana
syn. nov. e distribution, morphology, ecology and hosts of all species were reviewed from both published
and unpublished sources. New red-list categories for each species were created according to the new records
of occurrence. An identication key including all ten species and photographs of their whole bodies and
main identication characteristics was prepared, and distribution maps for all species were created.
Keywords
Europe, Aglaoapis, Dioxys, Ensliniana, Metadioxys, Paradioxys, maps, ecology, hosts, conservation
Introduction
Bees (Anthophila) form a group of seven families within the monophylum Aculeata
inside the highly diversied order Hymenoptera. is group coevolved with owering
plants, whose pollen, nectar and oils serve as the main food sources for both adults
JHR 96: 599–628 (2023)
doi: 10.3897/jhr.96.104957
https://jhr.pensoft.net
Copyright Petr Bogusch. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0),
which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
RESEARCH ARTICLE

Petr Bogusch / Journal of Hymenoptera Research 96: 599–628 (2023)
600
and their brood (Grimaldi and Engel 2005; Michener 2007). ere are many nest-
ing strategies for bees that vary according to the placement of the nest, the care of the
brood, the level of social behaviour, and the specialisation of the pollen (Klein et al.
2007; Michener 2007; Bogusch et al. 2020a). While the majority of bees create their
own nests, there is a group of interspecic nest kleptoparasites known as cuckoo bees,
which is distributed across all regions of the world and comprises approximately 15%
of all bee species (Batra 1984; Michener 2007; Sless et al. 2022). Cuckoo bees are not
a taxonomical group, but rather are ecological groups of phylogenetically unrelated
genera across multiple families, whose females lay their eggs into the nests of other bee
species. ey utilize a variety of strategies for invading host nests and destroying host
broods (Michener 2007; Habermannová et al. 2013; Westrich 2018).
Interspecic cuckoo behaviour is known in four families: Colletidae, Halictidae, Meg-
achilidae and Apidae, including the species-rich subfamily Nomadinae of the family Api-
dae which is comprised of many genera and species (Sless et al. 2022). Within the Megach-
ilidae, several genera with kleptoparasitic behaviour are distributed across more groups of
this family, while the tribe Dioxyini represents the least species-rich group of cuckoo bees
within Megachilidae (Gonzalez et al. 2012; Westrich 2018). is group of cuckoo bees
comprises eight genera of usually rare cuckoo bees, which are distributed in the Old World
and in North America (Popov 1936; Michener 2007). Currently, 36 species are known
worldwide (Michener 2007). Of this number, ten species in ve genera have been recorded
from Europe (Warncke 1977; Tkalcu 2001), most of which are very rare and restricted in
their distribution to a small area in the south of the continent. Many of these species have
their main distribution area in North Africa and/or the Middle East, and their occurrence
in Europe represents only the very small, northernmost part of their distribution (Warncke
1977; Baldock et al. 2018; Lhomme et al. 2020; Varnava et al. 2020; Ghisbain et al. 2023).
Warncke (1977) summarised the taxonomy of all species of the western part of the
Palaearctic region. However, this study is somewhat outdated and does not contain all
known species (i.e., Dioxys lanzarotensis Tkalcu, 2001, described after the publication
of Warncke’s (1977) study). e identication key is useful, especially because of the
quite simple identication of this group based on very specic dierences among the
species, but is very dierent from modern keys that include photographs. Additionally,
current surveys in several countries in southern Europe have improved the knowledge
of the distribution and ecology of this group.
e goal of this study is to review the taxonomy, distribution, ecology and conser-
vation of all species of tribe Dioxyini recorded from Europe. An identication key for
all species is included.
Materials and methods
e specimens for the study were collected by the author from the eld or obtained
from other collections. A large part of the material studied included pinned specimens
from both private and museum collections. Additionally, material from the following

Dioxyini of Europe 601
museums and institutions was studied: Národní muzeum, Praha (Czech Republic, cu-
rator Jan Macek), Moravské zemské muzeum, Brno (Czech Republic, curator Igor
Malenovský), Natural History Museum, London (United Kingdom, curator Joseph
Monks), Naturhistorisches Museum, Wien (Austria, curator Manuela Vizek), Biolo-
giezentrum Linz (Austria, curator Esther Öckermüller), Naturhistorisches Museum
Berlin (Germany, curator Stefanie Krause), and Naturkundemuseum Bayern Munich
(Germany, curator Stefan Schmidt). Several records were obtained from internet sourc-
es, especially from photos on Flickr (https://www.ickr.com/) and iNaturalist (https://
www.inaturalist.com/). In this case, only records from specialists or photos enabling
identication to the species level were included. However, the goal of this study was
not to create detailed maps of distribution, because this work will be done in several
European bee projects in the near future and occurrence records from local authorities,
taxonomic specialists and museums are still not completely collected and validated.
e author examined only a part of the type material of both valid species and syno-
nyms and all the original descriptions of all species and their synonyms. e type mate-
rial was not studied in most cases because the description of the species is comprehensive
and clear, and/or there was rich material available in private or museum collections for
study, so the examination of the original types was not necessary. All of the type material
studied by the author is indicated in the taxonomic treatment sections of species.
Morphology was studied using a digital Keyence VHX-700 photographing mi-
croscope with measuring tools. Only specimens clearly exhibiting diagnostic features
were imaged. e identication keys for females and males are dichotomous, using the
following standard abbreviations before the corresponding number: S – metasomal
sternum, T – metasomal tergum, and F – agellomere. Figure abbreviations in brackets
(Fig. 1A, Figs 2–4) are used in the text. Head measurements were performed from the
labrum base to the occipital ridge at the end of the head. If the same character applied
to both females and males, it was photographed only once (in the female). If the spe-
cies has a restricted range, it is also mentioned in the key in parentheses.
Species distributions were determined using studied material and literary sources
on Dioxyini of Europe. Distribution maps were created in QGIS 3.6. e red-list cat-
egories from Nieto et al. (2014) are indicated, and updates to conservation status are
proposed for several species.
Results
Identification key for the females of European Dioxyini
1 Scutellum without lateral projections (Fig. 15C); S6 and T5 elongated, more
than three times longer than wide (Fig. 15D) ...............................................
............................................... Ensliniana bidentata (Spain and Portugal)
– Scutellum with lateral toothlike projections (Fig. 1C), often with medial
tooth; S6 usually shorter (Fig. 1D) .............................................................. 2

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602
2 Head and thorax with dense reddish-brown hair; metasoma without apical
bands of whitish hair; metasoma and legs completely reddish (Fig. 3A) ........
........................................................... Dioxys ardens (Spain and Portugal)
– Head and thorax hairy or only sparsely haired, colour of hairs whitish or
brownish; metasoma and legs usually completely or partly black; metasomal
terga usually with narrow but well-developed apical bands of whitish hair
(Fig. 7A) .....................................................................................................3
3 Forecoxa with a toothlike carina anteriorly (Fig. 1E) ................................... 4
– Forecoxa rounded anteriorly (Fig. 7C) ........................................................5
4 Postscutellum with a medial toothlike projection, which is narrow and elon-
gated (Fig. 1C); T6 rounded and attened medially; S6 slightly emarginate
(Fig. 1D); body completely black with whitish hair (Fig. 1A) .......................
................................................................................... Aglaoapis tridentata
– Postscutellum with a medial toothlike process, which is not elongated (Fig.
17C); T6 not attened medially; S6 without an emargination (Fig. 17D);
legs, metasoma and antennae usually partly reddish (Fig. 17A) ....................
........................................................................ Metadioxys graeca (Greece)
5 S6 and T6 sharp and elongated (Fig. 19C); metasomal bands often not well de-
ned; metasoma and legs partly reddish (Fig. 19A) ....... Paradioxys pannonicus
– S6 and T6 not elongated (Fig. 7D); metasomal bands well or ill-visible; body
often completely black ................................................................................ 6
6 Apex of T6 truncate; S6 very slightly emarginate apically (Fig. 7D); black
with whitish hairs and often basal terga of metasoma reddish (Fig. 7A) ........
............................................................................................. Dioxys cinctus
– Apex of T6 convex (Fig. 11C); S6 with or without emargination; body com-
pletely black or partly reddish; legs can be also partly or completely reddish ..... 7
7 Metasoma completely black (Fig. 5A); small species (5–7 mm) .....................
......................................................................................... Dioxys atlanticus
– Metasoma partly reddish (Fig. 13A); legs partly reddish or dark; small to
larger species ............................................................................................... 8
8 Legs at least partly reddish (Fig. 13A, D); T6 coarsely, contiguously punc-
tate (Fig. 13C); S2 with medial part slightly protruded (Fig. 13D); head and
mesosoma in part with appressed, squamose pubescence ......Dioxys pumilus
– Legs black (Fig. 11A); T6 nely, densely, but not contiguously, punctate (Fig.
11C); S2 without protruded medial part (Fig. 11E); head and mesosoma
with pubescence semi-erect, not squamose ........................... Dioxys moestus
Identification key for the males of European Dioxyini
1 Scutellum without lateral projections; postscutellum without apical tooth-
like process in the middle (Fig. 15C); S7 and T7 with tooth-like processes
laterally (Fig. 15E) .................. Ensliniana bidentata (Spain and Portugal)
– Scutellum with lateral toothlike projections, often with medial tooth-like
process (Fig. 1D); S7 and T7 usually without tooth-like processes .............. 2

Dioxyini of Europe 603
2 Head and thorax with dense reddish-brown hair; metasoma without apical
bands of whitish hair, metasoma and legs completely reddish (Fig. 3B) .........
........................................................... Dioxys ardens (Spain and Portugal)
– Head and thorax hirsute or only sparsely haired, colour of hairs whitish or
brownish; metasoma and legs usually completely or partly black; metasomal
terga usually with narrow but well-developed apical bands of whitish hair
(Fig. 7B) .....................................................................................................3
3 Forecoxa with a toothlike carina anteriorly (Fig. 1E) ................................... 4
– Forecoxa rounded anteriorly (Fig. 7C) ........................................................5
4 Postscutellum with a medial toothlike projection, which is narrow and elon-
gate (Fig. 1C); T7 emarginated (Fig. 1F); body completely black with whitish
hair (Fig. 1B) ...............................................................Aglaoapis tridentata
– Postscutellum with a medial toothlike process, which is not elongated
(Fig. 17C); T7 not attened medially; last tergum not emarginated (Fig. 17E);
legs, metasoma and antennae usually partly reddish (Fig. 17B) .....................
.........................................................................Metadioxys graeca (Greece)
5 Metasoma completely reddish with semi-transparent apical parts of terga,
without well-visible apical bands (Fig. 19B); apex of S4 with two toothlike
processes medially (Fig. 19D, E); T7 with straight apex (Fig. 19F); S7 with
lateral teeth (Fig. 19E) .............................................Paradioxys pannonicus
– Metasoma black or partly reddish (Fig. 7B); rarely completely reddish but
then with well-visible apical bands of whitish or yellowish hair, apex of S4
straight or emarginated but without so prominent toothlike processes, S7
without lateral teeth .................................................................................... 6
6 Apex of S4 medially swollen, bidentate (Fig. 11F); matt and roughly punctate;
T7 shiny with coarse but sparse punctures (Fig. 11G) ........... Dioxys moestus
– Apex of S4 straight (Fig. 13E), emarginated or with small toothlike processes
(not as prominent as those of P. pannonicus); T7 more densely punctate .....7
7 Legs and metasoma partly or completely reddish (Fig. 13B, D); T7 triangular
with acute apex (Fig. 13F); apex of S4 waved (Fig. 13E) ...... Dioxys pumilus
– Legs dark, metosoma dark or partly reddish; T7 broad (Fig. 7F); with straight
or slightly curved apex; apex of S4 dierent ................................................ 8
8 Larger species (7–10 mm); at least part of T1 reddish (Fig. 7B); apex of S4
slightly emarginated (Fig. 7E); T7 coarsely and densely punctate (Fig. 7F) ...
..............................................................................................Dioxys cinctus
– Smaller species (5–7 mm); completely black (Fig. 5B); apex of S4 with two
toothlike processes (Fig. 5E); T7 with shiny interspaces .............................. 9
9 Mesonotum, scutellum and T1–T3 densely and deeply punctate, punctures
larger; mesosternum medially nely rugose (Fig. 5B, C); emargination between
toothlike processes on apex of S4 narrow (Fig. 5E) .............. Dioxys atlanticus
– Mesonotum, scutellum and T1–T3 more sparsely and nely punctate, punc-
tures smaller and shallow; mesosternum shiny (Fig. 9B, C); emargination
between toothlike processes on apex of S4 wider (Fig. 9D) ...........................
..................................... Dioxys lanzarotensis (Canary Islands: Lanzarote)

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Extralimital species
Altogether, 10 species of ve genera of tribe Dioxyini are known to occur in Europe.
ese are representatives of the genera Aglaoapis Cameron, Ensliniana Alfken,
Metadioxys Popov and Paradioxys Mocsáry (one species of each genus) and six species
of the genus Dioxys Lepeletier & Serville.
In the neighbouring regions, representatives of the genera Allodioxys Popov, Eudi-
oxys Mavromoustakis and Prodioxys Friese are recorded, but no species of these genera
has ever been recorded from Europe. Representatives of these genera dier from Euro-
pean species by the following characteristics (adopted from Warncke (1977)).
Both sexes of Allodioxys have the mesonotum with elongated projections on both
sides and postscutellum with a spine-like process medially. Four species occur in the
Middle East and/or North Africa (Libya, Algeria, Israel, Syria): Allodioxys ammobius
(Mavromoustakis), A. limbifera Pérez, A. moricei (Friese) and A. schulthessi Popov
(Warncke 1977; Lhomme et al. 2020).
Two species of Eudioxys occur in neighbouring regions: Eudioxys quadridentata
(Friese) in North Africa (Egypt) and E. schwarzi Mavromoustakis in the Middle East
(Iran) (Warncke 1977; Lhomme et al. 2020). Both sexes of this genus have axillae with
two spines laterally and the scutellum with one spine on both sides.
ree species of Prodioxys occur in North Africa: Prodioxys carneus Gribodo, P. lon-
giventris Pérez and P. ruventris Lepeletier (Warncke 1977; Lhomme et al. 2020). All
species are generally similar to Dioxys ardens with a completely reddish metasoma and
legs and brownish hirsute head and mesosoma, females dier by the very long and nar-
row last tergum and sternum, males have last sternum of dierent shape than D. ardens.
Accounts of European species
Genus Aglaoapis Cameron
Aglaoapis Cameron, 1901: 262. Type species: Aglaoapis brevipennis Cameron, 1901,
monobasic.
Dioxoides Popov, 1947: 89. Type species: Coelioxys tridentata Nylander, 1848, by origi-
nal designation.
Notes. e genus Aglaoapis is distributed in Europe, the Middle East, India and South
Africa. ree species are known worldwide (Michener 2007), and only one species oc-
curs in Europe (Ghisbain et al. 2023).
Aglaoapis tridentata (Nylander)
Coelioxys tridentata Nylander, 1848 (nec Apis tridentata Fabricius, 1775): 254.
Dioxys fasciata Schenck, 1861: 383.

Dioxyini of Europe 605
Dioxys kuntzei Noskiewicz, 1940: 99.
Dioxoides tridentata ssp. limassolica Mavromoustakis, 1949: 587.
Diagnosis. Larger species (9–12 mm), both sexes are black with well-developed white
bands of short hair on metasomal terga (Fig. 1A, B). Both sexes have the fore coxa with
a carina anteriorly (Fig. 1E) and a short projection and the scutellum with a medial
toothlike projection (Fig. 1C). Females have a longer last tergum than females of the
genus Dioxys and an emarginated last sternum (Fig. 1D). e last tergum is emargin-
ated for males (Fig. 1F).
Distribution. Aglaoapis tridentata is a Palaearctic species that occurs in Europe,
from Spain in the west to Russia in the east (Fig. 2), and in Asia from the Caucasus,
Kyrgyzstan and China, Kazakhstan, Siberia and Russian Far East. is species reaches
the farthest north of any species of the tribe, with records from Finland and Sweden
(Ornosa et al. 2008; Madsen and Calabuig 2010; Ascher and Pickering 2023).
Biology and hosts: Species recorded especially in steppic formations, sunny slopes, for-
est steppes and other open or semiopen habitats. Occurs also in abandoned sandpits, spoil
Figure 1. Aglaoapis tridentata A female, dorsal view B male, dorsal view C female, mesosoma dorsal view
D female, metasoma, dorsal view E female, fore coxa F male, T7, dorsal view. Red scale bars represent
the length of 1 mm.

Petr Bogusch / Journal of Hymenoptera Research 96: 599–628 (2023)
606
Figure 2. Aglaoapis tridentata, distribution in Europe.
heaps and other habitats of anthropogenic origin. is species attacks nests of bees of the
family Megachilidae, especially those nesting underground or making their own nests near
the ground surface. Hoplitis anthocopoides (Schenck), Hoplitis ravouxi (Pérez), probably also
Hoplitis adunca (Panzer) and Megachile pilidens Alfken, in southern parts of Europe, and
also Chalicodoma parietina (Georoy) were conrmed as hosts of this species (Westrich
2018). Scheuchl and Willner (2016) also listed Megachile leachella Curtis as a likely host.
Conservation status. Nieto et al. (2014) classied this species as LC – data de-
cient. is species is relatively rare throughout its range. Its distribution in Europe
is the largest of any species, and it is still being recorded in most countries – it was
reported to be regionally extinct only in Belgium and Finland (Ghisbain et al. 2023).
It should therefore be classied as LC.
Dioxys Lepeletier & Serville
Dioxys Lepeletier & Serville, 1825: 109, type species: Trachusa cincta Jurine, 1807,
monobasic.
Hoplopasites Ashmead, 1898: 284, type species: Phileremus productus Cresson, 1879,
by original designation.
Chrysopheon Titus, 1901: 256, type species: Chrysopheon aurifuscus Titus, 1901,
monobasic.

Dioxyini of Europe 607
Notes. Dioxys is a Holarctic genus distributed in most of Europe and North Africa to
central Asia in the east and the southwestern USA and adjacent Mexico in the west-
ern hemisphere (Hurd 1958; Michener 2007). Five species occur in North America,
and approximately 13 occur in the Palaearctic region (Warncke 1977; Ghisbain et al.
2023). Six species occur in Europe.
Dioxys ardens Gerstaecker
Dioxys ardens Gerstaecker, 1869: 161.
Dioxys ruspina Pérez, 1895:26.
Diagnosis. Larger species, body length 8–10 mm. In both sexes, typical in its colouration,
the metasoma is completely reddish without terminal or basal bands, and the legs and a-
gellum are reddish (Fig. 3A, B). T6 of females is elongated and narrowed posteriorly, with a
rounded apex. e mesosoma is ferruginously hirsute. Axillae with teeth and postscutellum
with short but sharp tooth in the middle. e same characteristics are also typical for males.
Distribution. Spain, Portugal (Fig. 4) and North Africa (Morocco to Libya).
Biology and hosts. Species recorded from semideserts and other arid open habi-
tats. Hosts unknown.
Conservation status. is species is known only from several records from south-
ern parts of Spain and one record from Portugal. Nieto et al. (2014) listed this species
as DD – data decient. According to the distribution records, it can be VU-vulnerable
because of its long-lasting rarity. However, we know only a little about the populations
and occurrence of this species in recent years.
Note. In general, a similar species, Dioxys chalicoda Lucas, was recorded from
North Africa (Algeria and Libya). One very old record is from Gibraltar, but this speci-
men was erroneously identied and belongs to D. ardens (coll. Biologiezentrum Linz,
Austria). is species diers in the colouration of the metasoma, which has black col-
ouration of at least the last three segments. Females have mandibles with lateral tuber-
cles, and males have ends of S4 with an emargination and S5 and S6 convex.
Dioxys atlanticus Saunders, 1904, stat. nov.
Dioxys atlanticus Saunders, 1904: 232.
Diagnosis. is species is small (5–7 mm in total length) and completely black with
well-developed apical bands on metasomal terga (Fig. 5A, B). It is generally similar to
Dioxys lanzarotensis, from which it can be identied by denser punctation on metaso-
mal terga (Fig. 5C) and clypeus. It is morphologically similar to D. moestus, which is
probably its near relative, rather than to D. cinctus, into which it was previously clas-
sied as a subspecies (Warncke 1977). e dark colouration of tibial spurs reported

Petr Bogusch / Journal of Hymenoptera Research 96: 599–628 (2023)
608
Figure 3. Dioxys ardens A female, dorsal view B male, dorsal view.
Figure 4. Dioxys ardens, distribution in Europe.
by Warncke (1977) was not observed in any specimen I have studied (Fig. 5D). Based
on the morphology and distribution, it is clearly a separate species. Syntypes (a male
and a female from Santa Cruz, Tenerife) from the Natural History Museum London
were studied.

Dioxyini of Europe 609
Distribution. is species was described from a male and a female from the Ca-
nary Islands (Tenerife), where it was also recorded on two other islands – Lanzarote
and Gran Canaria (Hohmann et al. 1993) (Fig. 6). It was also recorded in Egypt
(Warncke 1977) and currently in Sardinia (Orroli, 02.vi.2011, 6 ♀♀, G. Pagliano lgt.,
P. Bogusch det., coll. Biologiezentrum Linz, Austria). Based on the records, the spe-
cies occurs in the Canary Islands and several parts of North Africa, South Europe and
perhaps the Middle East, but it is very rare and hard to nd. Specimens from Sardinia
correspond in size, morphology and colouration with those of the Canary Islands and
with both syntypes.
Biology and hosts. e species occurs in open habitats – steppes, semideserts, in
rocky areas with shrubby vegetation. Little is known about its biology. Hosts unknown.
Conservation status: Nieto et al. (2014) listed this species as DD – data decient. ere
are quite recent records from the Canary Islands and new records from Sardinia (Italy). In
my opinion, it should be VU – vulnerable because of its restricted distribution area.
Figure 5. Dioxys atlanticus A female, dorsal view B male, dorsal view C female, mesosoma dorsal view
D female, tibial spur on fore leg E male, last metasomal segments, ventral view. Red scale bars represent
the length of 1 mm, blue scale bars 100 µm.

Petr Bogusch / Journal of Hymenoptera Research 96: 599–628 (2023)
610
Dioxys cinctus (Jurine)
Trachusa cincta Jurine, 1807: 253.
Dioxys pyrenaica Lepeletier, 1841: 515.
Dioxys maura Lepeletier, 1841: 516.
Dioxys cruenta Gerstaecker, 1869: 166.
Dioxys spinigera Pérez, 1884: 299.
Dioxys cincta var. jucunda Mocsáry, 1894: 36.
Dioxys cincta ab. friederikae Mader, 1933: 125.
Dioxys montana Heinrich 1977: 11–12, syn. nov.
Notes. Type specimens of this species and the description were studied in Biologiezen-
trum Linz, Austria. Both type specimens (a male and a female from the Sertavul Pass
in Turkey) do not morphologically dier from typical specimens of D. cinctus.
Diagnosis. Larger species, body length variable between 5–12 mm, probably de-
pending on the host. In both sexes, the body is black with the rst two metasomal terga
entirely or partly reddish and narrow apical bands of whitish short appressed hair (Fig.
7A, B). In several cases, red colouration is present on T3–T4. Mesosoma bears long
whitish hair, apex of T6 of females straight and only shortly elongated, shorter than
in similar species (Fig. 7D). e legs and antennae are black. is species has a very
large distribution area and is connected with many host species. It causes variability in
size (5–12 mm), while populations from southern Europe and North Africa are often
smaller. e colouration is very variable, too – normally both sexes have rst metaso-
mal terga entirely reddish but usually populations from the north of the distribution
area are darker and sometimes are entirely black with no reddish pattern.
Figure 6. Dioxys atlanticus, distribution in Canary Islands.

Dioxyini of Europe 611
Distribution. A species with a western Palaearctic distribution known from cen-
tral and southern Europe from Portugal to Greece and Romania (Fig. 8). Outside of
Europe, it is found in North Africa, Israel and as far east as the Caucasus (Dusmet
1921; Popov 1936; Warncke 1977; Standfuss et al. 2003; Ornosa et al. 2008; Ascher
and Pickering 2023).
Biology and hosts. Species occurring in a variety of open and semi-open habitats:
steppe, semideserts, forest steppes and many others. It was also recorded in sites of
anthropogenic origin – former sandpits, quarries, spoil heaps and military exercising
areas. is species has more host species in its large distribution area: Chalicodoma
parietina, Chalicodoma pyrenaica Lepeletier, Hoplitis adunca and Hoplitis anthocopoides
were conrmed (Scheuchl and Willner 2016). Its hosts nest underground, create nests
of mud or resin, or nest in various types of cavities above the ground. Parasitising
females of this species were often recorded around bee hotels associated with nests of
H.adunca in the Czech Republic (P. Bogusch, unpublished records). e actual num-
ber of hosts is certainly higher, and the preferred host species dier among the localities
within the large distribution area.
Figure 7. Dioxys cinctus A female, dorsal view B male, dorsal view C female, fore coxa D female, meso-
soma dorsal view E male, last metasomal segments, ventral view F male, T4–T6, dorsal view. Red scale
bars represent the length of 1 mm.

Petr Bogusch / Journal of Hymenoptera Research 96: 599–628 (2023)
612
Conservation status. Nieto et al. (2014) classied this species as LC – least con-
cern. It is distributed in most of southern and central Europe, with the northern
distribution border in France, Germany, the Czech Republic and Ukraine. In many
countries, it is not rare, and the numbers of recorded individuals are even higher than
those of A. tridentata. In the Czech Republic, the species has spread in the last 20 years
(Straka and Bogusch 2017). us, it should stay in the category LC.
Dioxys lanzarotensis Tkalcu
Dioxys lanzarotensis Tkalcu, 2001: 49–50.
Diagnosis. Small species very similar in morphology to D. atlanticus (Fig. 9A) but diers
by sparser punctures on metasomal terga (Fig. 9B) and on clypeus. Only a single male
(holotype) was recorded from the island Lanzarote of the Canary Islands (Spain) (Tkalcu
2001). e holotype should be deposited in Übersee-Museum Bremen, Germany, but
was discovered in the collection of Francisco La Roche in San Cristóbal de La Laguna,
Tenerife, Spain. According to the studies of the holotype, this species is very similar to
males of D. atlanticus but diers by the above reported characteristics, as well as several
others (length and position of toothlike processes on S4 (Fig. 9D), shape of head). It is
certainly a separate species; however, its distribution area is restricted to one island.
Distribution. Lanzarote, Canary Islands, Spain (Fig. 10) (Tkalcu 2001).
Figure 8. Dioxys cinctus, distribution in Europe.

Dioxyini of Europe 613
Figure 9. Dioxys lanzarotensis A male, dorsal view B male, metasoma, dorsal view C male, mesosoma
dorsal view D male, last metasomal segments, ventral view. Red scale bars represent the length of 1 mm.
Figure 10. Dioxys lanzarotensis, distribution in Canary Islands.
Biology and hosts. Unknown.
Conservation status. Nieto et al. (2014) classied this species as DD – data de-
cient. It is the only category in which this species can be classied because we know
only one specimen, the type.

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614
Dioxys moestus Costa
Dioxys moesta Costa, 1883: 96.
Dioxys rotundata Pérez, 1883: 300.
Diagnosis. Middle-sized species, body length 5–8 mm. Species with typical general
appearance for this genus, black with rst 2–3 metasomal terga entirely or partly
reddish, with narrow apical bands of whitish short appressed hair (Fig. 11A, B).
Mesosoma with long whitish hair, apex of metasomal T6 rounded (Fig. 11C). e
legs and antennae are black. e last metasomal terga in males was not as narrow as
that in D. cinctus. In general, similar to D. cinctus but diers by several character-
istics: it is usually smaller and the reddish colouration is more distributed (usually
on T1–3, in D. cinctus on T1–T2). Females of D. cinctus have a straight apex of T6,
while females of D. moestus have a round apex of T6. Males of D. moestus have two
Figure 11. Dioxys moestus A female, dorsal view B male, dorsal view C female, mesosoma dorsal view
D female, metasoma, dorsal view E female, metasoma lateral F male, last metasomal segments, ventral
view G male, T4–T6, dorsal view. Red scale bars represent the length of 1 mm, blue scale bars 100 µm.

Dioxyini of Europe 615
tooth-like processes on S4 medio-posteriorly (Fig. 11F), and males of D. cinctus
lack this characteristic. Punctation of T2–T3 of D. moestus is ner and sparser than
in D. cinctus.
Distribution. Mediterranean species described from Sardinia. Recorded from
Portugal to Greece (Fig. 12). Outside of Europe, it is recorded in North Africa, from
Morocco to Tunisia, and Israel (Warncke 1977; Ornosa et al. 2008).
Biology and hosts. is species occurs in open habitats. It was collected in
open habitats with shrubby vegetation, steppic formations or rocky landscapes
with almond orchards. Hosts are Hoplitis benoisti (Alfken), Hoplitis fertoni (Pérez)
and Hoplitis zandeni (Teunissen & van Achterberg) (Bogusch et al. 2020b), prob-
ably also Hoplitis ochraceicornis (Ferton) (I. Cross, unpublished record). Several
specimens were reared from nests of H. fertoni placed inside snail shells (Bogusch
et al. 2020b).
Conservation status. Nieto et al. (2014) classied this species as DD – data de-
cient. is species occurs in most of southern Europe, where it is rare but still fre-
quently recorded. It can be classied as LC – least concern.
Note. Dioxys heinrichi Warncke occurs in North Africa (Morocco and Algeria) and
is similar to D. moestus. Female D. heinrichi have a longer F2 and more convex clypeus,
and males do not have a swollen end of S4 and lack the two small teeth. e end of S5
is slightly emarginated.
Figure 12. Dioxys moestus, distribution in Europe.

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616
Dioxys pumilus Gerstaecker
Dioxys pumilus Gerstaecker, 1869: 167.
Dioxys varipes De Stefani, 1887: 113.
Dioxys maroccana Popov, 1936: 16.
Dioxys cypriaca Popov, 1944: 121.
Diagnosis. Smaller species, total body length 4–6 mm. Species with typical general ap-
pearance for this genus, black with rst 2–4 metasomal terga entirely or partly reddish,
with narrow apical bands of whitish short appressed hair (Fig. 13A, B). Mesosoma with
long whitish hair, apex of metasomal T6 rounded (Fig. 13C). e legs and antennae are
at least partly reddish. Last metasomal terga in males not as narrowned as in D. cinctus,
last tergum more curved than in D. moestus and D. cinctus. Species in general similar
to smaller individuals of D. moestus and D. cinctus dier by reddish legs and agellum.
Females have a T6 that is longer than it is wide (distinctly longer than the T6 of both
Figure 13. Dioxys pumilus A female, dorsal view B male, dorsal view C female, metasoma, dorsal view
Dfemale, metasoma lateral E male, last metasomal segments, ventral view F male, last metasomal seg-
ments, dorsal view. Red scale bars represent the length of 1 mm.

Dioxyini of Europe 617
similar species), males do not have a sharp medio-posterior projection on S4 (present
in D. moestus) but the apex of S4 is waved, not straight as in D. cinctus (Fig. 13E).
Punctation of T2–T3 is coarser and denser than in D. moestus and D. cinctus. Speci-
mens from Cyprus (described by Popov, 1944 as a separate species) look Smore colour-
ful at rst view but do not dier in their morphology, and the diagnostic characteristics
of D. cypriaca are variable and form a continuous line to D. pumilus. us, D. cypriaca
is currently supposed to be a synonym of D. pumilus.
Distribution. is is a western Palaearctic species. e nominate subspecies oc-
curs in the eastern Mediterranean basin (Greece, Cyprus, Turkey) (Fig. 14) and spreads
towards Asia Minor (Israel, Syria) and Iran. e subspecies D. p. varipes occurs in the
western Mediterranean basin (Sicily, Spain, Morocco, Algeria, Tunisia). e taxonomic
statuses of these subspecies are unclear, but they do not dier in morphology, other
than the specimens from western parts of the distribution area often being darker than
those from the east.
Biology and hosts. is species was recorded in a variety of open and semi-open
habitats – steppes, forest steppes, semideserts, open landscapes with shrubby vegeta-
tion and many others. Heriades crenulatus Nylander was reported as a likely host of this
species in Cyprus (Mavromoustakis 1959). In Portugal, it was recorded in association
with Hoplitis annulata Latreille (Baldock et al. 2018). Small species of Osmiini are also
supposed to be host species of D. pumilus.
Figure 14. Dioxys pumilus, distribution in Europe.

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618
Conservation status. Nieto et al. (2014) classied this species as DD – data de-
cient. is species occurs in many countries in southern Europe, while in several locali-
ties, it has been recorded in large series. It is more local than D. moestus but probably
more numerous at the localities. It can be classied as LC – least concern.
Ensliniana Alfken
Ensliniana Alfken, 1938: 431, type species: Ensliniana cuspidata Alfken, 1938 = Stelis
bidentata Friese, 1899, by original designation.
Dioxoides Popov, 1947: 89. Type species: Coelioxys tridentata Nylander, 1848, by origi-
nal designation.
Notes. is genus is distributed in North Africa and the Middle East, from Morocco
in the west to Turkmenistan in the east. ree species were described, one of which
was reported from Europe – Portugal and Spain (Popov 1936, 1953; Michener 2007).
Ensliniana bidentata (Friese)
Stelis bidentata Friese, 1899: 285.
Paradioxys pannonica var. rupes Friese, 1899: 285.
Dioxys richaensis Friese, 1911: 139.
Dioxys bidentata Friese in Schulthess, 1924: 319.
Ensliniana cuspidata Alfken, 1938: 431.
Diagnosis. Larger species, body length 7–10 mm. e only species of the genus re-
corded from Europe. It is typical by the characteristics of the genus; both sexes are
generally similar to Dioxys species (Fig. 15A, B) but lack axillar teeth (Fig. 15C). T5 of
females is shiny, and T6 and S6 are elongated with two lateral teeth, similar to Paradi-
oxys pannonica (Fig. 15D). P. pannonica diers in the black or dark brown colour of the
entire body, with a reddish pattern only on the rst three metasomal terga and distinct
apical bands of whitish short appressed hair on the metasomal terga.
Distribution. In Europe, only several specimens are known from Spain and Por-
tugal (Ornosa and Ortiz-Sánchéz 2014; Torres 2020; Ascher and Pickering 2023) (Fig.
16). It was described from Israel (Jericho and Oran). Outside of Europe, it is known
from Morocco, Algeria, Tunisia, Turkey, Syria, Israel and Jordan (Grace 2010).
Biology and hosts. is species probably occurs in open habitats – steppic grass-
lands, rocky slopes, semideserts and other habitats. Baldock et al. (2018) listed Hoplitis
zaianorum (Benoist) as a likely host of this species.
Conservation status. Nieto et al. (2014) classied this species as DD – data de-
cient. is species is known in Europe only from Spain and Portugal, where it was
recorded both in the past and in recent years. It can be classied as VU – vulnerable
because of its restricted distribution area in Europe.

Dioxyini of Europe 619
Figure 15. Ensliniana bidentata A female, dorsal view B male, dorsal view C female, last metasomal seg-
ments, dorsal view D female, mesosoma, dorsal view E male, last metasomal segments, dorsal view. Red
scale bars represent the length of 1 mm.
Figure 16. Ensliniana bidentata, distribution in Europe.

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620
Metadioxys Popov
Metadioxys Popov, 1947: 88, type species: Dioxys formosa Morawitz, 1875, by original
designation.
Notes. is genus has a similar distribution to the previous genera in North Africa and
the Middle East, from Morocco in the west to Uzbekistan in the east. ree species
were described, one of which was reported from Europe – Greece (Popov 1936, 1953;
Michener 2007).
Metadioxys graeca (Mavromoustakis)
Dioxys formosa graeca Mavromoustakis, 1963: 696.
Figure 17. Metadioxys graeca A female, dorsal view B male, dorsal view C female, mesosoma dorsal view
D female, metasoma, dorsal view E male, last metasomal segments, dorsal view. Red scale bars represent
the length of 1 mm.

Dioxyini of Europe 621
Figure 18. Metadioxys graeca, distribution in Europe. Doubtful record from Italy in orange.
Diagnosis. Larger species, body length 8–10 mm. Species generally very similar to
those of the genus Dioxys, black with rst 2–4 metasomal terga entirely or partly red-
dish (rarely whole metasoma reddish and other body parts reddish), with narrow api-
cal bands of whitish short appressed hair (Fig. 17A, B). Whole body with scale-like
whitish short appressed hair, similar to those of Epeolus species. Apex of metasomal T6
more elongated than in Dioxys and Aglaoapis, rounded (Fig. 17D). e legs and anten-
nae are at least partly reddish. Typical with sharp carina on coxa of front leg laterally,
postscutellum without teeth medioapically, only with an ill-visible tubercle (Fig. 17C).
In both sexes, T6 was without emargination (Fig. 17D, E).
Distribution. In the European region, this species is known from Greece (essaly
and Crete) (Fig. 18). Its range extends out of the European region to Asiatic Turkey,
Morocco and Israel (Warncke 1977; Grace 2010; Ascher and Pickering 2023; Kuhl-
mann et al. 2023).
Biology and hosts. is species probably occurs in open habitats – steppic grass-
lands, rocky slopes, semideserts and other habitats. Hosts unknown.
Conservation status. Nieto et al. (2014) classied this species as DD – data de-
cient. ere are only old records from Greece and one doubtful record from Italy
(certainly this species but probably wrongly labelled), and no recent nds are known.
Because of its unknown population trend, we must leave this species in category DD
– data decient.
Note. Metadioxys formosa Morawitz occurs in North Africa (Morocco) and the
Middle East (Israel and Turkmenistan). It is smaller than M. graeca (6–8 mm), and

Petr Bogusch / Journal of Hymenoptera Research 96: 599–628 (2023)
622
both sexes have scale-like whitish short appressed hair distributed on more parts of
the body than M. graeca. Female has last tergum broadened laterally; male has deeply
emarginated last sternite.
Paradioxys Mocsáry
Paradioxys Mocsáry, 1894: 35, type species: Dioxys pannonica Mocsáry, 1877,
monobasic.
Notes. is genus is reported from southeastern Europe and the Middle East. Its oc-
currence ranges from Austria in the west to Iran in the east. Two species are known, one
of which occurs in Europe (Popov 1936; Michener 2007).
Figure 19. Paradioxys pannonicus A female, dorsal view B male, dorsal view C female, last metasomal seg-
ments, dorsal view D male, metasoma, ventral view E male, last metasomal segments, ventral view F male,
last metasomal segments, dorsal view. Red scale bars represent the length of 1 mm, blue scale bars 100 µm.

Dioxyini of Europe 623
Paradioxys pannonicus (Mocsáry)
Paradioxys pannonica Mocsáry, 1894: 35.
Diagnosis. Middle-sized species, body length 7–10 mm. e only species of the genus
within Europe. Typical by the characteristics of the genus, similar to species of the ge-
nus Dioxys. It is uniform in appearance, females black with rst four metasomal terga
entirely reddish amd males’ whole metasoma reddish. Legs are also reddish (Fig. 19A,
B). Metasoma of females have narrow apical bands of whitish short appressed hair;
similar hairs are also distributed on the rst metasomal terga laterally. e metasoma
of males have bands of whitish short appressed hair that are sparse and barely visible.
e male body has a tomentum-like light brown or yellowish hair. Females have a
narrowed T6 and a very long and sharp S6, projecting behind the T6 (Fig. 19C). In
males, the end of the metasoma is straight, not curved (Fig. 19F), and the apex of S4
has two prominent toothlike projections, which are larger than in males of Dioxys
(Fig. 19D, E).
Distribution. is species was described from Hungary and is a Euro-Asiatic spe-
cies that occurs from central Europe to the north-eastern Mediterranean (Fig. 20) and
outside of Europe towards the Middle East and Iran (Popov 1936; Warncke 1977;
Bogusch et al. 2007; Gusenleitner et al. 2012; Ascher and Pickering 2023).
Figure 20. Paradioxys pannonicus, distribution in Europe.

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624
Biology and hosts. is species attacks nests of species of the family Megachilidae.
Scheuchl and Willner (2016) reported Chalicodoma hungarica (Mocsáry) as its main
host, and Gusenleitner et al. (2012) included C. parietina as its host.
Conservation status. Nieto et al. (2014) classied this species as DD – data de-
cient. is is a typical Pannonian species occurring in the Pannonian basin. However,
there is a lack of records from recent years, and the most recent nds are from the
1970s and 1980s. We suppose this species is CR – critically endangered in Europe.
Discussion
Cuckoo bees of the tribe Dioxyini are represented by ten species in Europe. ese
cuckoo bees are usually rare, and sightings are not common. Only two species occur in
a large part of the continent. Aglaoapis tridentata is a relatively rare species of steppic
habitats whose distribution covers large parts of Europe, including Scandinavia (Swe-
den and Finland) and Russia. Dioxys cinctus occurs in most of southern and central
Europe, where it spreads towards the north – the rst record from the northwestern
part of the Czech Republic (Bohemia) comes from 2012 (Straka et al. 2015) and
from Germany from 2019 (Saure and Petrischak 2020). Other species occur only in
southern Europe, while two of them were recorded from a larger part of this region.
Dioxys moestus has a larger distribution but is usually recorded in small numbers of
individuals, in contrast to D. pumilus, which has a smaller distribution but numerous
series of individuals from multiple localities. Paradioxys pannonicus is known from
central and southeastern Europe historically but there are no recent records on the
occurrence of this species in Europe (the latest record I have revised is from Bulgaria
from 1983). Two species occur in North Africa and reach their distribution area to
Spain and Portugal. Ensliniana bidentata was recorded and observed several times af-
ter 2000 (Baldock et al. 2018), but for Dioxys ardens the latest record from Spain dates
back to 1976. A similar case is seen in Metadioxys graeca, which occurs in the Middle
East and in Greece in Europe, where it was recently recorded only in Crete. Dioxys
atlanticus and D. lanzarotensis were thought to be endemic to the Canary Islands.
Dioxys lanzarotensis is known only from a single male from Lanzarote, in contrast,
D. atlanticus was collected on Lanzarote, Gran Canaria and Tenerife, while several
specimens were collected on Gran Canaria around the year 2020 (P. Bogusch, own
observations). But D. atlanticus was also recorded from Egypt (Warncke 1977) and
recently from Sardinia.
Several other species were recorded in North Africa or the Middle East. All these
species are rare, and their occurrence in Europe is unlikely. e record of Dioxys chali-
coda from Gibraltar belongs to D. ardens (P. Bogusch revised). us, in future, studies
on the distribution of D. atlanticus in southern Europe (if the species will be discovered
elsewhere than in Sardinia and Egypt) and attempts to nd specimens or populations
of P. pannonica would be interesting. Although this species has not recorded for a long
time, it is likely to occur in Slovakia, Hungary, Bulgaria, Greece or Romania.

Dioxyini of Europe 625
Dioxyini are cuckoo bees. is means that discovering these bees is more compli-
cated than discovering nesting bee species. eir hosts are bees of the family Megachi-
lidae, which usually do not nest in aggregations, and thus their nests are harder to nd.
However, several species have quite visible nests made of mud (genus Chalicodoma) or
can be recorded in bee hotels (hosts of D. cinctus as well as this cuckoo bee species).
Furthermore, nests of Hoplitis fertoni parasitised by D. moestus were reported from
empty snail shells in Spain (Bogusch et al. 2020b).
To preserve the fauna of this tribe of cuckoo bees, it is necessary to take care of
their habitats. Most species were recorded mainly in open habitats of steppic or semi-
desert characteristic. Conservation of these habitats across Europe can go hand in hand
with the conservation of the hosts and be helpful for these rare, beautiful and interest-
ing cuckoo bees.
Acknowledgements
I would like to thank to all curators and collection owners for helping with the ma-
terial. e study was supported by the Excellence Research Project of University of
Hradec Kralove Nr. 2202/2023.
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