A revision of the Andrena (Hymenoptera: Andrenidae) of Lebanon with the description of six new species

Abstract

Lebanon has a rich but chronically understudied bee fauna. As part of ongoing efforts to better understand bee diversity in Lebanon we present a critical revision of the speciose genus Andrena based primarily on the examination of specimens collected from contemporary surveys, as well as previously undetermined material from museum collections. Andrena (Aciandrena) abruptifovea n. sp. from Lebanon, A. (incertae sedis) cedricola n. sp. from Lebanon and Syria, A. (Euandrena) scrophulariae n. sp. from Lebanon, and A. (incertae sedis) prodigiosa n. sp., from Lebanon, are described. A. (Chlorandrena) edentula n. sp. and A. (Rufandrena) parvispinae n. sp., both from northwestern Syria, are described due to their proximity to northern Lebanon. The previously unknown male of A. (Pallandrena) christineae Dubitzky, 2006 is also described. A. (Poliandrena) unifasciata Friese, 1899 stat. rev. which was described from Lebanon is removed from synonymy with A. (Poliandrena) caspica Morawitz, 1886, and the relationship between these two taxa and A. (Poliandrena) unicincta Friese, 1899 is clarified. Altogether, these changes increase the number of Andrena species known from Lebanon to 86, including species known previously only from Turkey or Israel. In total, 25 and 23 species are also reported as newly recorded for Jordan and Syria, respectively. Based on the diversity of Andrena in neighbouring countries in the Levant, it is probable that the true number of Andrena species in Lebanon is well over 100.

Full text

Full Terms & Conditions of access and use can be found at
https://www.tandfonline.com/action/journalInformation?journalCode=tase20
Annales de la Société entomologique de France (N.S.)
International Journal of Entomology
ISSN: (Print) (Online) Journal homepage: https://www.tandfonline.com/loi/tase20
A revision of the Andrena (Hymenoptera:
Andrenidae) of Lebanon with the description of six
new species
Thomas J. Wood , Mira Boustani & Pierre Rasmont
To cite this article: Thomas J. Wood , Mira Boustani & Pierre Rasmont (2020): A revision of the
Andrena (Hymenoptera: Andrenidae) of Lebanon with the description of six new species, Annales
de la Société entomologique de France (N.S.)
To link to this article: https://doi.org/10.1080/00379271.2020.1794960
View supplementary material
Published online: 23 Sep 2020.
Submit your article to this journal
View related articles
View Crossmark data

A revision of the Andrena (Hymenoptera: Andrenidae) of Lebanon with the description of six
new species
Thomas J. Wood*, Mira Boustani & Pierre Rasmont
Laboratoire de Zoologie, Université de Mons, 7000, Mons, Belgium
(Accepté le 7 juillet 2020; publié en ligne le 23 septembre 2020)
Summary. Lebanon has a rich but chronically understudied bee fauna. As part of ongoing efforts to better understand bee
diversity in Lebanon we present a critical revision of the speciose genus Andrena based primarily on the examination of
specimens collected from contemporary surveys, as well as previously undetermined material from museum collections.
Andrena (Aciandrena) abruptifovea n. sp. from Lebanon, A. (incertae sedis) cedricola n. sp. from Lebanon and Syria, A.
(Euandrena) scrophulariae n. sp. from Lebanon, and A. (incertae sedis) prodigiosa n. sp., from Lebanon, are described.
A. (Chlorandrena) edentula n. sp. and A. (Rufandrena) parvispinae n. sp., both from north-western Syria, are described
due to their proximity to northern Lebanon. The previously unknown male of A. (Pallandrena) christineae Dubitzky, 2006
is also described. A. (Poliandrena) unifasciata Friese, 1899 stat. rev. which was described from Lebanon is removed from
synonymy with A. (Poliandrena) caspica Morawitz, 1886, and the relationship between these two taxa and A.
(Poliandrena) unicincta Friese, 1899 is claried. Altogether, these changes increase the number of Andrena species
known from Lebanon to 86, including species known previously only from Turkey or Israel. In total, 25 and 23 species are
also reported as newly recorded for Jordan and Syria, respectively. Based on the diversity of Andrena in neighbouring
countries in the Levant, it is probable that the true number of Andrena species in Lebanon is well over 100.
Résumé. Révision des Andrena (Hymenoptera : Andrenidae) du Liban, avec la description de six espèces nouvelles.
Le Liban a une riche faune d’abeilles trop peu étudiée de manière chronique. Comme contribution à l’effort actuel pour
mieux comprendre la diversité des abeilles du Liban, nous présentons ici une révision critique du genre Andrena, connu
pour être riche en espèce. Cette révision est basée sur l’examen de spécimens collectés durant des recherches récentes,
ainsi que de matériel indéterminé de collections de musées. Les espèces nouvelles suivantes sont décrites du Liban :
Andrena (Aciandrena) abruptifovea n. sp. (Liban), A. (incertae sedis) cedricola n. sp. (Liban et Syrie), A. (Euandrena)
scrophulariae n. sp. (Liban), and A. (incertae sedis) prodigiosa n. sp. (Liban). Les espèces suivantes sont décrites du
nord-ouest de la Syrie, à proximité du Liban : A. (Chlorandrena) edentula n. sp. et A. (Rufandrena) parvispinae n. sp. Le
mâle encore inconnu d’A. (Pallandrena) christineae Dubitzky, 2006, est aussi décrit. A. (Poliandrena) unifasciata Friese,
1899 stat. rev., décrit du Liban, est retiré de la synonymie d’A. (Poliandrena) caspica Morawitz, 1886, tandis que les
relations entre ces deux derniers taxons et A. (Poliandrena) unicincta Friese, 1899, sont clariées. Tout compté, ces
changements augmentent le nombre d’espèces d’Andrena connues du Liban à 86, y compris des espèces qui auparavant
n’étaient connues que de Turquie et Israël. Au total, 25 et 23 espèces sont aussi comptées pour la première fois de Jordanie
et de Syrie, respectivement. En se basant sur la diversité des Andrena dans les autres pays du Levant, il est probable que le
nombre total d’espèces du Liban excède 100.
http://www.zoobank.org/urn:lsid:zoobank.org:pub:9FC27F37-A201-4C17-BE46-1C2AAA337CCB
Keywords: Levant; endemic species; alpine; solitary bees; taxonomy
The genus Andrena Fabricius, 1775 is the second most
species-rich genus of bees worldwide after
Lasioglossum Curis, 1833 (Ascher & Pickering 2020).
The genus is richest in Mediterranean and xeric areas,
so unsurprisingly the Eastern Mediterranean including
the Levant is a hotspot of Andrena diversity. The
Andrena fauna of the Eastern Mediterranean was exten-
sively revised by Klaus Warncke in the 1960s and
1970s, with important papers focusing on Greece
(Warncke 1965a), Israel (Warncke 1969a), and Turkey
(Warncke 1965b, 1969b, 1975a, 1975b). Despite these
extensive efforts, new Andrena species continue to be
described from this region at regular intervals (e.g.
Gusenleitner 1998; Scheuchl et al. 2004; Grünwaldt
et al. 2005; Scheuchl & Hazir 2012; Schwenninger
2015; Kratochwil 2015; Pisanty et al. 2016, 2018),
and it is still too early to put a number on the true
species richness of Eastern Mediterranean Andrena.
*Corresponding author. Email: thomasjames.wood@umons.ac.be
Annales de la Société entomologique de France (N.S.), 2020
https://doi.org/10.1080/00379271.2020.1794960
© 2020 Société entomologique de France
Published online 23 Sep 2020

Lebanon is a small country in the Levant, but it
has a wide variety of ecological regions and biotypes
(Abi-Saleh & Sa 1988), and a rich but chronically
understudied bee fauna. For example, there are only
four published papers that report detailed Andrena
records from Lebanon (Friese 1899; Mavromoustakis
1962; Warncke 1969a; Schuberth et al. 2001). This
situation is not helped by historical changes in the
geographic scope of Lebanon. In the early part of
the 20th century Lebanon was part of Ottoman Syria
until the end of the First World War, where what is
today modern Lebanon, Syria, and Hatay province in
Turkey was ceded to France as part of the Mandate for
Syria and the Lebanon following the Sykes–Picot
agreement between Britain and France (Traboulsi
2012). Therefore, literature prior to 1918 would refer
simply to ‘Syria’ (e.g. Friese 1899) without allowing
contemporary authors to identify a locality or distin-
guish between modern day Lebanon or Syria. Even
later works would simply give very little distributional
information, for example Friese (1922) who reported
Andrena cubiceps Friese, 1914 var. fulvopilosa from
“Syrien, Adana [Turkey]”, making it impossible to
know if this taxon is present in Lebanon or not.
However, though there is a lack of clarity over the
exact country referred to in what historic literature is
available, this does not greatly change the overall
picture of a lack of literature itself and historical
study. Only three species of Andrena have been
described with a locus typicus in what is now modern
day Syria (one currently considered valid) and two
from modern day Lebanon (two considered valid, see
entry for Andrena unifasciata Friese, 1899 below)
compared to the nearly 200 species described from
Israel and Turkey (e.g. Warncke 1969b; Warncke
1975b; Gusenleitner 1998; Scheuchl & Hazir 2012;
Pisanty et al. 2016, 2018). Against this background,
the contemporary study is not particularly limited by
the confusion over historical geographic terminology.
As part of efforts to improve our understanding of
Lebanese bees, the country has been surveyed for
multiple years and museum material located in order
to critically revise the national fauna. Collection
efforts focused on important plant areas such as
Tannourine Cedars, Bcharre Cedars and Horch Ehden
in northern Lebanon which harbour a high proportion
of endemic plants (Bou Dagher-Kharrat et al. 2018),
and which may therefore also contain range-restricted
and previously undescribed bee species, as is the case
for other invertebrate groups (Larsen 1974; Németh
2019). As part of this effort, here we revise the
Andrena fauna of Lebanon, describe six new species
from Lebanon and Syria, and present additional data
on some species newly recorded in the nearby coun-
tries of Jordan and Syria. Collecting sites for material
examined by this project are shown in Figure 1, with
full geographic coordinate details in online supple-
mentary material. This paper is not intended to be a
revision of the Syrian or Jordanian Andrena faunas,
but as part of an extensive revision of undetermined
museum material from the Levant by the lead author a
number of new country records were detected.
Therefore, where species listed here as present in the
Lebanese fauna were also newly detected in these two
other countries, this information is given. The
Jordanian and Syrian Andrena faunas will be revised
in later publications.
Methodology
Revisions to the Andrena subgeneric system of classication
are ongoing (Pisanty et al. 2020), and classications are
likely to change substantially in the future. However, this
will require extensive revisionary work over many years,
and so for now we broadly follow the subgeneric system of
Warncke (1968) with modications for when changes to this
system are simple, clear, and well-supported (Gusenleitner &
Schwarz 2002; Pisanty et al. 2020).
Specimens collected during this study were collected by
hand net and killed with ethyl acetate or by freezing. At
some collection localities, blue, yellow, and white pan traps
were used. These were lled to the ¾ mark with water plus
colourless and odourless soap and left for 48 hours before
collection. Each collection station used three sets of triplets
(one of each colour, separated by 1 m) which were them-
selves separated by 10 m.
All data were digitised using Data Fauna Flora 5.1.2
(Barbier et al. 2000) and mapped with Quantum GIS
2.18.27. All geographic co-ordinates are given using the
datum World Geodetic System of 1984 (WGS 1984). Where
detailed records are available from the literature, these are
included (Friese 1899; Mavromoustakis 1962; Schuberth
et al. 2001). An additional publication (Grace 2010) presented
an overview of the wild bee species of the Eastern
Mediterranean, reporting 55 species of Andrena from
Lebanon. However, the underlying data supporting these list-
ings is unclear and so here we take a conservative position
and only list this publication where other lines of evidence
exist, never in isolation. Twenty-three unconrmed species
from Grace (2010) are listed in online supplementary material
with notes. Species marked with an asterisk * in the main
manuscript have not previously been reported from Lebanon
in any form.
Morphological terminology follows Michener (2007).
Specimens were measured from the vertical plane of the front
of the head to the tip of the metasoma to the nearest 0.5 mm.
Photographs were taken using an Olympus E-M1 Mark II with a
60 mm macro lens and were stacked using Zerene Stacker 1.04
(Zerene Systems, USA) and plates were prepared in GNU
Image Manipulation Program (GIMP) 2.10.
2T. J. Wood et al.

Abbreviations
DACN = Department of Agriculture collection, Nicosia, Cyprus
MSC = Maximillian Schwarz personal collection, Ansfelden,
Austria
OÖLM = Oberösterreich Landesmusum, Linz, Austria
RBIN = Royal Belgian Institute of Natural Sciences, Brussels,
Belgium
SOILS = Soils Permaculture Association, Saidoun, Lebanon
TCFNR = Tannourine Cedar Forest Nature Reserve,
Tannourine, Lebanon
TJW = Personal collection of Thomas Wood, Mons,
Belgium
USEK = Holy Spirit University of Kaslik, Jounieh, Lebanon
UMONS = University of Mons, Mons, Belgium
XVA = Xavier van Achter personal collection, Mons,
Belgium
Figure 1. Map of Lebanon with the collecting locations of material examined during this project (black dots). Additionally, specic
collection locations for species newly described here from Lebanon are indicated (red dots), including one additional location from Syria.
Full sampling details can be found in online supplementary material. Geographic relief is indicated, with altitude measured in metres above
sea level.
Annales de la Société entomologique de France (N.S.) 3

Results
Descriptions of new species
Andrena (Aciandrena) abruptifovea Wood, n. sp.
http://www.zoobank.org/urn:lsid:zoobank.org:act:97069DFB-
7648-4BA8-A0BD-B37FF27F55EB
Type material. Holotype: ♂, LEBANON, Mount
Lebanon, Chouf Biosphere Reserve, Barouk trails, 1772 m
[33°41'10''N 35°41'56''E, Figure 1, location 5], 16.V.2019,
leg. Boustani, Thlaspi spp. Deposited at the RBIN.
Paratypes: LEBANON: Mount Lebanon, Chouf
Biosphere Reserve, Barouk trails, 1772 m, 16.V.2019, 5♂
[#1–5], 1♀ [#6], leg. Boustani, Thlaspi spp.; N Lebanon,
Harrisa, Al Jawar, 1758 m, 11.V.2019, 1♂ [#7], 1♀ [#8], leg.
Boustani, Brassica spp. (Figure 1, location 2); N Lebanon,
Tannourine Reserve, 1781 m, 13.V.2019, 1♀ [#9], leg.
Boustani, Thlaspi spp. (Figure 1, location 2); N Lebanon,
Arz Tannourine Gate area, 1754 m, 2.IV.2018, 1♀ [#10],
leg. Boustani (Figure 1, location 2); N Lebanon, Hadath El
Jebbe, border of the Cedar forest, 1618 m, 5.V.2017, 1♀
[#11], leg. Boustani (Figure 1, location 2); N Lebanon, Arz
Bcharre, Forest of the Cedars of God, 1897 m, 20.V.2019,
1♂ [#12], leg. Boustani, Alyssum spp. (Figure 1, location 1).
Paratypes are deposited at the RBIN [#1–5; #9; #11], the
DACN [#7–8], and in the personal collection of TJW [#6;
#10; #12].
Description of female. Body length 6 mm (Figure 2).
Head. Slightly longer than broad, frons, paraocular areas,
gena, and vertex dull metallic green, clypeus black
(Figure 3). Clypeus clearly domed and weakly punctured,
punctures separated by 2–3 puncture diameters.
Underlying surface shagreened and weakly shining,
becoming shinier at the apical margin. Process of labrum
narrow, triangular, pointed, clearly longer than wide
(Figure 4). Face, gena, vertex, and scape with short
white hairs, these not exceeding the length of the scape.
Antennal segments 1–5 black, segments 6–12 predomi-
nantly orange below and above, segment 3 slightly
exceeding 4 + 5 in length. Fovea narrow, equalling the
width of an antenna above where they occupy half the
distance between a lateral ocellus and the top of the
compound eye, fovea abruptly narrowed to approximately
30% of their maximum width about 2/3rd of the way
from the top of the compound eye to the level of the
antennal insertions. Ocelloccipital distance extremely
short, almost non-existent, posterior ocellus almost touch-
ing vertex.
Mesosoma. Scutum and scutellum densely shagreened,
weakly shining, with a subtle green-purple metallic
sheen (Figure 4), weakly and shallowly punctured, punc-
tures separated by 2–3 puncture diameters. Pronotum
non-carinate. Episternum and propodeum nely
shagreened, weakly shining, propodeal triangle well dif-
ferentiated, nely reticulate, weakly shining. Episternum
and propodeum laterally with long white hairs, these not
exceeding the length of the scape, scutum and scutellum
with shorter white hairs. Legs dark, apical tarsal segments
lightened orange, pubescence white. Floccus, femoral,
and tibial scopa white. Wings hyaline, venation amber,
nervulus antefurcal.
Metasoma. Terga dark with hints of a bronze metallic
sheen, evenly shagreened and weakly shining, apical
margins of T2–4 slightly depressed and lightened pale
yellow (Figure 5). T2–3 with weak apical lateral hair
fringes, these widely interrupted. T5 and hairs anking
pygidial plate orange-golden. Pygidial plate triangular,
pointed, attened and without a raised central area.
Description of male. Body length 5–5.5 mm (Figure 6).
Head. Black, as wide as long. Clypeus domed, evenly
shagreened, weakly shining, clearly punctured, punctures
separated by 1–2 puncture diameters laterally, becoming
sparser centrally to form a broad almost impunctate line.
Clypeus most commonly predominantly yellow marked
(Figure 8) but this can vary from 80–90% coverage to
50% coverage to completely black (Figure 9). Pubescence
as in the female. Antennal segments 1–6 dark, segments
7–13 slightly lightened orange below, segment 3 slightly
shorter than 4 + 5 in length. Ocelloccipital distance short,
wider than female, at most ¼ width of posterior ocellus.
Mesosoma. As in the female.
Metasoma. Tergites as in the female, weakly shining with
uniform even shagreenation (Figure 7). T2–4 with weak
apical lateral hair fringes, also interrupted widely.
Genitalia simple (Figure 10), gonocoxites with weak
rounded apical teeth, penis valve slightly widened cen-
trally, gonostyli with parallel sides, forming a slight api-
cal point, becoming slightly translucent at their apexes.
Diagnosis. Andrena abruptifovea can be placed into the
subgenus Aciandrena in the female sex by the
combination of small size, black integument, narrow
facial fovea, and a shagreened propodeal triangle, and in
the male sex by the additional character of a yellow
marked clypeus.
Is most similar to Andrena (Aciandrena) pulicaria
Warncke, 1975 from Greece and Turkey, most notably
in the male sex where the clypeus is domed and partly
yellow marked (Figure 8, though the coloration can be
variable, see Figure 9), in contrast to other Aciandrena
species where the male clypeus is completely at or
attened and completely or almost completely yellow
marked (e.g. A. pratincola Warncke, 1974 and A. lamiana
Warncke, 1965, respectively). It can easily be separated
by a comparison of the genitalia as A. pulicaria has a
4T. J. Wood et al.

clear, rounded protrusion on the inner margin of the
gonostyli (Figure 11; also illustrated in Warncke 1975b).
The genital capsule is similar to the recently described A.
israelica Scheuchl & Pisanty, 2016 with moderately pro-
nounced gonocoxal teeth, but the gonostyli of A.
abruptifovea are longer and come to a clearer point
(Figure 10, but not as clear as in species like A. aciculata
Morawitz, 1886, see illustrations in Warncke 1972). The
two species can also be separated because the clypeus of
A. israelica is attened and black. In a series of six males
Figures 2–9. Andrena (Aciandrena) abruptifovea n. sp. 2, Female prole, 3, female face, 4, female labrum, 5, female tergites, 6, male
prole, 7, male tergites, 8, male face, yellow marked form, 9, male face, black form.
Annales de la Société entomologique de France (N.S.) 5

of A. abruptifovea taken from the type locality, two had a
predominantly yellow clypeus, three had a partially yel-
low clypeus, and one was completely black. However,
they were all domed, and the two species can be separated
on this basis rather than simply by colour.
In the female sex, A. abruptifovea can also be placed
close to A. pulicaria because of the domed clypeus that is
basally and centrally shagreened and weakly shiny but
becoming clearly shiny at the apical margin. However,
the two can be clearly separated by the labral process
which like an equilateral triangle in A. pulicaria (roughly
as long as wide) but like an isosceles triangle in A.
abruptiofovea (clearly longer than wide, Figure 4). The
foveae are also more sharply and abruptly narrowed
below (Figure 4, also clearly distinguishing this taxon
from A. israelica and A. judaea Scheuchl & Pisanty,
2016) and the tergal integument is much less strongly
sculptured, shagreened and therefore weakly shining
(Figure 5), hammer shagreened and dull in A. pulicaria.
Additional Aciandrena material is available from lower
elevations from Israel and Syria that is very close morpholo-
gically to A. abruptifovea. However, it differs in the male sex
because the clypeus is atter, and the genital capsule is shorter,
with comparatively shorter and less pointed gonostyli, and the
penis valve is thinner and more parallel sided without the
slightly bulging sides medially that can be seen in A. abrupti-
fovea (Figure 10). In the female sex, the clypeus is slightly less
domed and the fovea, whilst constricted below like A. abrup-
tifovea, are narrower overall (occupying slightly less than half
of the space between a lateral ocellus and the compound eye at
their maximal width, for A. abruptifovea occupying half this
space). These differences are subtle, particularly in the female
sex, but given the difference in male genitalia morphology we
take the position that these are not the same taxon as A.
abruptifovea, and their status needs to be established, prefer-
ably with molecular techniques.
Other material examined (Andrena (Aciandrena)
species close to A. abruptifovea). ISRAEL: Jerusalem
env., 20.III.1993, 3♀, leg. D. Abel; SYRIA: An Nasrah
env. [Al-Nasrah], 8–13.IV.2005, 2♀, leg. S. Jald; Tartus,
250 m, St. Georg-Klost., 3.IV.1988, 1♀, leg. L. Blank;
Tartus, Sata, 10 km N, 300 m, 3.IV.1988, 2♂, 1♀, leg. L.
Blank; Aleppo, 500 m, Simeons-Kloster, 19.IV.1988, 1♀,
leg. L. Blank; Latakia, 750 m, Qaranjah, 3.IV.1988, 1♀,
leg. L. Blank, currently all TJW.
Distribution. Known from high altitude sites in central
and northern Lebanon.
Floral preferences. All oral records were made from
Brassicaceae. Other Aciandrena species for which data
are available show exclusive use of Brassicaceae (TJW
unpublished data), and this may be the case for A.
abruptifovea but more study is required.
Etymology. The name abrupti (abrupt, sudden) + fovea
(the facial fovea that help to characterise Andrena in the
female sex) was chosen to describe the abruptly
constricted fovea that run along the inner margins of the
compound eyes, helping distinguishing this species from
other Levantine members of the subgenus.
Andrena (Chlorandrena) edentula Wood, n. sp.
http://www.zoobank.org/urn:lsid:zoobank.org:pub:25E04964-
3516-4666-9871-82DD924CC373
Type material. Holotype: ♂, SYRIA occ., An Nasrah
env [Al-Nasrah, 34°47'37''N 36°17'23''E], 8–13.IV.2005,
leg. J. Saki. Deposited at the OÖLM.
Female. Unknown.
Description of male. Body length 11 mm (Figure 12).
Head. As long as wide, dark but with a very subtle slight
metallic sheen, most obvious in the centre of the clypeus
(Figure 13). Clypeus black, with shallow raised reticula-
tion forming weak lateral carinae, underlying surface
microreticulate, weakly shining. Process of labrum
broadly trapezoidal, twice as wide as long, front margin
widely and shallowly emarginate. Face, gena, vertex, and
scape with light brown-yellow hairs, the longest of these
equalling the scape in length. Antennae dark, antennal
Figures 10–11. Andrena (Aciandrena), male genitalia. 10, A. (A.) abruptifovea n. sp.; 11, A. (A.) pulicaria Warncke, 1975.
6T. J. Wood et al.

segments 3–13 below with extremely short and dense
hairs giving a lightened silvery impression, segment 3
shorter than 4 + 5. Ocelloccipital distance slightly exceed-
ing diameter of posterior ocellus.
Mesosoma. Scutum and scutellum strongly shagreened,
densely punctate, punctures separated by 0.5–1 puncture
diameter, underlying surface weakly shining. Pronotum
non-carinate. Scutellum centrally with a very small area
free of punctures, underlying surface smooth and shining.
Episternum and propodeum reticulate, underlying surface
weakly shining, propodeal triangle well marked with a
weak carina and by a change in sculpture, basally with
short longitudinal carinae, dull. Episternum, propodeum,
scutum, and scutellum with long light brown-yellow
hairs, the longest of these on the episternum clearly
exceeding the length of a scape. Legs dark, nal tarsal
segments slightly lightened red-brown, pubescence
yellow. Wings hyaline, venation and stigma amber, ner-
vulus interstitial.
Metasoma. Tergites dark, apical margins of T2–5 slightly
lightened red-brown (Figure 14). Tergites clearly and
densely punctured, punctures of typical Chlorandrena
crater form with raised rim, this most apparent on T1,
punctures on following tergites progressively with less
clearly marked rims, punctures separated by 1 puncture
diameter. Underlying surface on tergal discs weakly
shagreened, shining, tergal margins with clear lateral
microreticulation, dull. Pubescence weak, without clear
pattern, T1–5 generally with sparse yellow to dark brown
hairs, T6 with apical margin covered by a loose fringe of
yellow hairs, these not obscuring the underlying surface.
S8 in dorsal view greatly widened, with short laterally
projecting teeth basally, apically with a short lamellate
projection, weakly pointed (Figure 15). S8 in ventral view
Figures 12–17. Andrena (Chlorandrena) edentula n. sp. 12, Male prole; 13, male face; 14, male tergites; 15, male sternite 8; dorsal
view; 16, male sternite 8; ventral view; 17, male genitalia.
Annales de la Société entomologique de France (N.S.) 7

with thick laterally projecting hair tufts that clearly
obscure the underlying surface (Figure 16). Centrally
and ventrally at the level of the laterally teeth with a
thin and inconspicuous ventrally projecting hair tuft, per-
pendicular to the sternal plate itself. Genitalia with
weakly pronounced gonocoxite teeth, these almost absent
(Figure 17). Penis valve very narrow, almost the same
width throughout its length. Gonostyli broad, only
slightly narrowing apically where they form a attened
and lamellate apex with a small projecting point, internal
margin with a raised carina.
Diagnosis. Andrena edentula can be quickly placed into the
Chlorandrena because of the deep and distinct ‘crater
punctures’ on the terga that are impressed with a slightly
raised rim. It can be further placed into the rhenana-group
because sternite 8 is broad with lateral tooth-like projections
(Figure 15; Schwenninger 2015). The rhenana-group was
recently revised in the West Palearctic (Schwenninger
2015), but A. edentula is unlike any previously described
member, as sternite 8 is extremely short and has thick
lateral hair tufts (Figure 16), centrally and ventrally at the
level of the lateral teeth with a tuft of long hairs that projects
perpendicularly and ventrally away from the sternal plate, and
the genitalia are unique with the normally clear gonocoxal
teeth greatly reduced and an extremely narrow penis valve
(Figure 17). The gonostyli show an afnity to other members
of the rhenana-group as their apexes are attened and
lamellate, forming only a weak point, with a raised carina
along their inner margin, like A. orientana Warncke, 1965
and A. taraxaci Giraud, 1861, but the combination of weak
gonocoxal teeth and the narrow penis valve are unique. Using
the key of Schwenninger (2015), A. edentula keys to couplet
10 at which point it matches neither species as the genitalia
are markedly divergent.
Distribution. Known only from the locus typicus in
north-western Syria. This locality is just 20 km north of
the Lebanese border, and so the species may be present in
northern Lebanon.
Floral preferences. None recorded but members of the
Chlorandrena are very strongly tied to Asteraceae
(Cichorieae or Anthemideae, Schmid-Egger & Scheuchl
1997; Amiet et al. 2010; TJW unpublished data).
Etymology. The name edentula (toothless) was chosen
because the male genitalia have greatly reduced
gonocoxal teeth which is unusual for members of the
rhenana-group (Schwenninger 2015).
Andrena (Euandrena) scrophulariae Wood, n. sp.
http://www.zoobank.org/urn:lsid:zoobank.org:pub:
A8E26F5F-D7DB-4AE8-A82F-FA3C0C098122
Type material. Holotype: ♀, LEBANON: N Lebanon,
Arz Tannourine, Gate Area, 1754 m [34°12'25''N 35°
55'55''E, Figure 1, location 2], 20.V.2018, leg. Boustani,
Scrophularia spp. Deposited at the RBIN. BOLD
sequence entry number: HYMAA061-20.
Paratypes: LEBANON: N Lebanon, Arz Tannourine,
Gate Area, 1754 m, 20.V.2018, 3♀ [#1-3], leg. Boustani,
Scrophularia spp. Paratypes are deposited at the RBIN [#1],
the DACN [#2], and in the personal collection of TJW [#3].
Description of female. Body length 9–9.5 mm
(Figure 18).
Head. Dark, but with a clear greenish metallic sheen, this
particularly pronounced on the basal part of the clypeus as the
colour transitions from weak metallic green (clypeus mar-
gins) to black (centre of the clypeus, Figure 19). Clypeus
domed and centrally attened, clearly and strongly punctured,
punctures separated by 1 (laterally) to 2 (centrally) puncture
diameters. Clypeus with a faint longitudinal impunctate line
centrally. Clypeus shagreened laterally, transitioning to
smooth and shining centrally. Process of labrum twice as
long as broad, clearly emarginate apically (Figure 20). Face
centrally with white hairs, laterally with black hairs. Gena and
scape with white hairs, vertex with a mixture of black and
white hairs, the longest of these hairs equalling the length of
the scape. Antennae dark, segments 5–12 slightly lightened
grey below, segment 3 exceeding 4 + 5 in length, slightly
shorter than 3 + 4 + 6. Foveae comma shaped, equalling the
width of an antenna above where they occupy half of the
space between a lateral ocellus and the top of the compound
eye, narrowing below to 60% of their maximum width, lled
with a light brown to dark brown pilosity depending on the
angle of observation. Ocelloccipital distance slightly shorter
than width of posterior ocellus.
Mesosoma. Scutum densely shagreened, weakly shining,
clearly punctured, punctures separated by 1–2 puncture
diameters, underlying integument with faint metallic
sheen but not as pronounced as on the face. Pronotum
non-carinate. Scutellum laterally the same as the scutum,
but centrally smooth and shining, without shagreenation.
Episternum and propodeum dull, weakly reticulate, pro-
podeal triangle visible by a change in sculpturing, almost
without rugosity except basally. Episternum, propodeum,
scutum, and scutellum with long whitish branched hairs,
these equalling the length of the scape. Legs dark, tarsi
not noticeably lightened, pubescence light brown. Floccus
and femoral scopa white, tibial scopa light orange, com-
posed of simple hairs. Wings slightly yellowed, venation
amber, nervulus antefurcal.
Metasoma. Tergites dark, narrow apical part of tergal
margins lightened translucent yellow (Figure 21). T1
without shagreenation, smooth and shiny, following ter-
gites with subtle shagreenation, predominantly shiny. T1
8T. J. Wood et al.

with sparse but clear and small punctures, punctures
separated by 3 puncture diameters, following tergites
more densely and weakly punctured, punctures separated
by 2 puncture diameters. T1 with a fringe of long hairs on
its apical margin, these hairs clearly exceeding the length
of the margin itself. T2–4 with fringes of shorter and
denser hairs, these only slightly exceeding the length of
the margins themselves and obscuring the underlying sur-
face. T5 and the hairs anking the pygidial plate dark
brown. Pygidial plate at, without central raised portion,
densely punctured, with impunctate and slightly raised
margin.
Male. Unknown.
Diagnosis. Andrena scrophulariae clearly has comma-
shaped fovea that are relatively broad above and
narrowed below, and would initially seem to be a good
t for the Euandrena. However, it has an elongate face
(Figure 19) and the process of the labrum is much more
elongate than is typical for Euandrena (Figure 20),
suggesting possibly afnity with Didonia (Warncke
1968). Most specically, A. scrophulariae resembles A.
(Didonia) solenopalpa from France and Spain that has a
similarly elongate face. However, COI barcode data
unequivocally places A. scrophulariae within the
Euandrena (C. Praz and TJW unpublished data).
Additionally, provisional molecular Andrena phylogenies
(S. Bossert in litt.) suggest that A. solenopalpa may be
nested within the Euandrena and the Ptilandrena (Pisanty
et al. 2020; see also Praz et al. 2019 for discussion over
the paraphyly of these two subgenera). Indeed, previous
authors have found problems with the concept of Didonia
and have split off several previous Didonia species into a
new subgenus Hamandrena based on the presence of
strong hooked bristles on the galea (not including A.
solenopalpa that lacks these bristles, Dubitzky et al.
2010). Given these outstanding problems surrounding
Andrena classication, we take the approach of placing
this taxon within the large Euandrena group, and await
future subgeneric reclassication efforts with interest.
Andrena scrophulariae can easily be separated from
A. solenopalpa by the shape of the labral process, which
is widely triangular in A. solenopalpa but elongate trape-
zoidal in A. scrophulariae with a clearly emarginate front
margin. It has a longer face than any currently described
species of Eastern Mediterranean Euandrena.
Distribution. Known only from Arz Tannourine
(Harissa) in northern Lebanon, but given what is known
about other high altitude Cedar forests and their
associated Andrena communities this species is likely to
be more widespread across this region.
Floral preferences. All specimens were collected from
unidentied Scrophularia (Scrophulariaceae) species. The
one available pollen load contained pure Scrophularia
pollen. The elongated face of the female suggests a
strong association with owers with deep corollas, but
Figures 18–21. Andrena (Euandrena) scrophulariae n. sp. 18, Female prole; 19, female face; 20, female labrum; 21, female tergites.
Annales de la Société entomologique de France (N.S.) 9

more study is required before rm conclusions can be
drawn.
Etymology. The name scrophulariae (of Scrophularia) was
chosen because of the use of this genus as a pollen source.
Andrena (incertae sedis) cedricola Wood, n. sp.
http://www.zoobank.org/urn:lsid:zoobank.org:
pub:53413ABA-919A-4D87-BC94-0545BAF4191A
Type material. Holotype: ♀, LEBANON, N Lebanon,
Hadath El Jebbe, border of the Cedar forest, 1646 m [34°
12'58''N 35°56'13''E, Figure 1, location 2], 13.V.2017, leg.
Boustani, Brassicaceae. Deposited at the RBIN.
Paratypes: LEBANON: N Lebanon, Hadath El Jebbe,
border of the Cedar forest, 1618 m, 5.V.2017, 1♂ [#1], 1♀
[#2], leg. Boustani; Mount Lebanon, Falougha, Cedar
woods, 1480 m, 27.IV.2019, 1♀ [#3], leg. Boustani
(Figure 1, location 4); Mount Lebanon, Laqlouq, Matoube,
1657 m, 30.IV.2017, 1♀ [#4], leg. Boustani (Figure 1, loca-
tion 3); N Lebanon, Harrisa, Al Jawar, 1758 m, 11.V.2019,
2♂ [#5-6], leg. Boustani, white Brassica (Figure 1, location
2); SYRIA: Faouar [probably Camp Faouar UNDOF, Golan
Heights, c. 1000 m – it is not clear exactly where this
sampling point is, and the author may have been at a higher
altitude towards Mount Hermon], 31.III.2001, 1♂ [#7], leg.
J. Plass (Figure 1, location 7). Paratypes are deposited at the
RBIN [#2; #6], the DACN [#3; #5], the OÖLM [#7], and the
personal collection of TJW [#1; #4].
Description of female. Body length 7.5–8 mm
(Figure 22).
Head. Black, as long as wide (Figure 23). Clypeus
slightly domed, shagreened, weakly shining, with scat-
tered but clear punctures, punctures separated by 1–3
puncture diameters. Process of labrum triangular, slightly
longer than wide (Figure 24). Face, gena, and scape with
short white hairs, these becoming longer and more yel-
lowish on the vertex, the length of these hairs approach-
ing but not exceeding the length of the scape. Antennae
predominantly dark, segments 9–12 becoming very
slightly lightened orange below, segment 3 slightly longer
than 4 + 5, shorter than 4 + 5 + 6. Fovea relatively broad
above, exceeding the width of an antenna, but only occu-
pying 1/3rd of the distance between a lateral ocellus and
the top of the compound eye, narrowing below to 50% of
their maximum width. Ocelloccipital distance narrow, at
most 2/3 width of posterior ocellus.
Mesosoma. Scutum and scutellum shagreened, weakly to
strongly shining centrally, surface clearly and evenly
punctured, punctures separated by 1–2 puncture dia-
meters. Pronotum non-carinate. Episternum and propo-
deum shagreened and reticulated, dull, propodeal
triangle clearly delineated with a shallow marginal carina,
internal surface with clear raised reticulation. Episternum
and propodeum laterally with white hairs, scutum and
scutellum dorsally with shorter light brown hairs. Legs
uniformly dark, pubescence whitish. Floccus, femoral and
tibia scopa white. Wings hyaline, venation brown, nervu-
lus antefurcal.
Metasoma. Tergites dark, T1 entirely and T2–4 with
apical areas without shagreenation, smooth and shiny
(Figure 25). T2–4 basally with shagreenation and extre-
mely scattered and sparse punctures. T1 sparsely but
nely punctured, punctures separated by 2–3 puncture
diameters. T2–4 both basally and apically with loose
lateral hair patches and hair fringes respectively
(Figure 25). T5 and hairs anking pygidial plate light
brown, laterally with scattered white hairs. Pygidial
plate at, without raised central portion, densely punctate
with a thin impunctate margin.
Description of male. Body length 6.5–7 mm (Figure 26).
Head. Black, slightly wider than long (Figure 27).
Clypeus black, sculpturing as in the female. Process of
labrum roughly square, slightly wider than long. Pilosity
and ocelloccipital distance as in the female, antennae
uniformly dark, segment 3 shorter than 4 + 5.
Mesosoma. Scutum and scutellum strongly shagreened,
weakly shining, shallowly and unevenly punctured, punc-
tures with slightly raised rims, separated by 1–2 puncture
diameters. Episternum and propodeal sculpturing, meso-
somal pilosity, and pronotal carination as in the female.
Legs dark, apical tarsal segments slightly lightened.
Wings hyaline, venation dark brown, nervulus slightly
antefurcal.
Metasoma. As in the female, T1 and T2–5 apically
smooth and shiny, without shagreenation, contrasting
with the basally shagreened and dull parts (Figure 28).
Pubescence as in the female. Genitalia long and slim,
gonocoxites with weak rounded teeth, penis valve narrow,
and gonostyli parallel sided, slightly wider at their apex
than in their middle (Figure 29).
Diagnosis. Andrena cedricola morphologically resembles
members of the subgenus Poecilandrena based on the
criteria outlined by Pisanty et al. (2018), specically A.
stenofovea Scheuchl & Pisanty, 2018. However, this
subgenus has been treated as a ‘wastebasket’ for species
lacking apomorphies, and is strongly paraphyletic
(Pisanty et al. 2020). As such, we do not formally place
A. cedricola into a subgenus at this time, but it can be
grouped with the Poecilandrena more generally for
species recognition and identication purposes.
In the female sex it is most similar to A. stenofovea
because of the aberrant small and triangular labral process
(Figure 24). However, the tergites are completely
10 T. J. Wood et al.

different, with wide and shiny marginal areas (Figure 25),
whereas in A. stenofovea the tergites are uniformly
shagreened, with the margins barely differentiated and
difcult to see. Though they do narrow below, the facial
foveae are also wider than the width of an antenna above,
whereas they are narrower than the width of an antenna
throughout their entire length in A. stenofovea.
The male genitalia are simple (Figure 29) and closest
to species like A. sedumella Scheuchl & Pisanty, 2018
and A. limassolica Mavromoustais, 1948, but are unique
amongst Levantine species by the combination of gono-
coxal teeth and uncurved gonostyli that broaden slightly
at their apex, as well as by the unique sculpturing of the
tergal margins as in the female.
Figures 22–29. Andrena (incertae sedis) cedricola n. sp. 22, Female prole; 23, female face; 24, female labrum; 25, female tergites;
26, male prole; 27, male face; 28, male tergites; 29, male genitalia.
Annales de la Société entomologique de France (N.S.) 11

Modifying the key of Pisanty et al. (2018), the female
of A. cedricola keys to couplet 3 at which point it meets
some of the characters of A. stenofovea (triangular lab-
rum), but not others (facial fovea extremely narrow and
elongate) but also does not agree with the alternative
(facial fovea uniformly broad, labral process trapezoidal
to rectangular). The male of A. cedricola keys to couplet
5 at which point it does not agree with either pathway
(gonocoxites have a dorsal lobe, but the clypeus is not
yellow marked). Both sexes can therefore be easily sepa-
rated from other similar Levantine Andrena.
Distribution. Known from high altitude sites in central
and northern Lebanon and the Golan Heights, Syria.
Floral preferences. One female and two males were
collected from Brassicaceae. A different female
specimen (N Lebanon, Hadath El Jebbe, border of the
Cedar forest, 1618 m, 5.V.2017) with no oral visitation
data had a scopa full of pollen. Pollen analysis of this
load showed that it contained pure Brassicaceae pollen
(18 μm in diameter, Thlaspi-type). More study is required
before any rm conclusions can be drawn for this species.
Etymology. The name cedri (Cedar, Cedrus, genus of
Old World trees) + cola (inhabiting, living in a place)
was chosen because Lebanese specimens were collected
from high altitude sites characterised by stands of Cedrus
libani (Pinaceae), the national emblem of Lebanon.
Andrena (Rufandrena) parvispinae Wood, n. sp.
http://www.zoobank.org/urn:lsid:zoobank.org:pub:
DACFB7F0-5A14-4E86-A8C9-25CB79428CB8
Type material. Holotype: ♂, SYRIA, Latakia, Qaranjah,
750 m [probably 35.783°N 35.900°E], 3.IV.1988, leg. L.
Blank. Deposited at the OÖLM.
Paratypes: SYRIA: Latakia, Qaranjah, 750 m, 3.
IV.1988, 4♂, leg. L. Blank; Tartus, Sata, 10 km E, 300 m
[probably Alhulu west forests], 1♂, leg. L. Blank. Paratypes
are deposited at the OÖLM with one retained in the personal
collection of TJW.
Female. Unknown.
Description of male. Body length 5 mm (Figure 30).
Head. Black, clearly wider than long (Figure 33). Clypeus
arched, weakly shagreened basally and laterally, centre
weakly shiny, entire surface yellow with coloration
extending onto the lower paraocular areas. Clypeus sur-
face weakly and shallowly punctured, centrally with
punctures separated by 3–4 puncture diameters, laterally
by 2–3 puncture diameters. Process of labrum weakly
trapezoidal, twice as broad as long. Face, gena, vertex,
and scape with white hairs, these not exceeding the length
of the scape. Antennae black, antennal segments slightly
lightened orange-brown below from segment 5 onwards,
segment 3 slightly exceeding 4 + 5 in length, shorter than
4 + 5 + 6. Ocelloccipital distance short, at most ½ width
of posterior ocellus.
Mesosoma. Scutum and scutellum strongly shagreened
but weakly shining, sparsely, shallowly, and subtly punc-
tured, punctures separated by 2–3 puncture diameters.
Pronotum non-carinate. Episternum shagreened, weakly
shining. Propodeal triangle slightly marked by a weakly
raised external carina, centrally with weak rugosity.
Mesosoma with white hairs, these densest and longest
on the episternum, the longest hairs equalling the length
of the scape. Legs dark, lightened to dark red-orange at
the apex of the tarsi, pubescence white. Wings hyaline,
venation amber, nervulus clearly postfurcal.
Metasoma. Tergites dark, apical margins lightened yellow
to amber, hyaline (Figure 32). Tergal integument
shagreened, the strength of the shagreenation decreasing
from the T1 to T5, therefore becoming shinier. T1 almost
impunctate, with the density of punctures increasing
across tergites, T5 therefore moderately but very shal-
lowly punctate, punctures separated by 2 puncture dia-
meters. Tergites laterally with small patches of white
hairs, very widely interrupted, these most pronounced
on T2–4. S7 widely emarginate, S8 widened and at-
tened, mallet shaped, laterally with two short but clearly
projecting spines (Figure 31). Genitalia long, gonocoxa
with moderately pronounced and rounded teeth, gonostyli
curved with a thin, hyaline outer margin (Figure 34).
Penis valve centrally with two lateral translucent semi-
circular projections.
Diagnosis. Andrena parvispinae can be instantly
recognised as a member of the subgenus Rufandrena
because of the characteristic structure of sternite 8 in the
male sex which is strongly widened, and which possesses a
clear spine on each side that clearly projects through
surrounding hairs (Figures 30, 31). This character is so
clear and distinctive that it is actually the rst couplet in
Warncke’s (1968) key to West Palearctic Andrena
subgenera. Only two species have been placed into the
Rufandrena, A. ruventris Lepeletier, 1841 (Morocco to
Libya) and A. orbitalis Morawitz, 1871 (north-western
Africa, Iberia, France, and Italy, Gusenleitner & Schwarz
2002). As well as being separable simply by their much
larger size (10–11 mm in the male sex against 5 mm in A.
parvispinae), both A. ruventris and A. orbitalis have quite
different genitalia with a marked emargination in the apical
part of the gonostyli (Figures 34, 37, 40). The three species
share a similar broader than long head shape, and similar
12 T. J. Wood et al.

white facial markings, though those in A. parvispinae are
more extensive (Figures 33, 36, 39).
Distribution. Known only from Syria from Tartus to
Latakia. The locality labels suggest inland locations in
wooded districts. The species may be in northern
Lebanon and south-eastern Turkey as both localities are
within 20 km of the respective borders, but these areas
must be searched.
Floral preferences. None recorded. Its relative Andrena
orbitalis may be a specialist of Plantago (Plantaginaceae,
TJW unpublished data), and so Plantago species could be
searched in the spring. Nothing is known of the oral
preferences of A. ruventris.
Etymology. The name parvi (small) + spinae (spines) was
chosen to describe the two tiny spines in the sides of
sternite eight that place this bee in the subgenus
Rufandrena. The name can also have a double meaning,
referring to the small body size of the species itself relative
to the other two members of this subgenus (i.e. the small
bee in the group of Andrena with spines on sternite eight).
Figures 30–40. Andrena (Rufandrena). 30–34, A. (R.) parvispinae n. sp.: 30, male prole; 31, male sternite eight; 32, male tergites;
33, male face; 34, male genitalia; 35–37, A. (R.) orbitalis Morawitz, 1871: 35, male tergites; 36, male face; 37, male genitalia; 38–40,
A. (R.) ruventris Lepeletier, 1841: 38, male tergites; 39, male face; 40, male genitalia.
Annales de la Société entomologique de France (N.S.) 13

Andrena (incertae sedis) prodigiosa Wood, n. sp.
http://www.zoobank.org/urn:lsid:zoobank.org:pub:
F7A264EC-C11D-4009-B6E7-1C1683AA9C9F
Type material. Holotype: ♂, LEBANON, Beka’a,
Qaraoun Lake [33°33'57''N 35°42'28''E, Figure 1, location
6], 24.III.2013, leg. M. Kasparek. Deposited at the OÖLM.
Female. Unknown.
Description of male. Body length 8 mm (Figure 41).
Head. Black, clearly broader than long (Figure 42).
Clypeus arched, apically truncate. Underlying surface
very weakly shagreened in the apical half and shiny,
strongly shagreened basally and laterally, dull. Shiny
areas of clypeus sparsely punctured, punctures separated
by 2–3 puncture diameters, dull areas of clypeus densely
punctate, punctures separated by 0.5–1 puncture
diameters. Process of labrum rectangular, three times
wider than long, weakly shining. Face, gena, vertex, and
scape with whitish to yellowish hairs, some of these
exceeding the length of the scape. Antennal segments
1–3 entirely dark, segments 4–13 orange below.
Segment 3 equals segments 4 + 5. Ocelloccipital distance
equalling width of posterior ocellus.
Mesosoma. Scutum and scutellum densely shagreened,
dull, with irregular shallow punctures with clearly raised
rims, punctures separated by 0.5–2 puncture diameters.
Pronotum non-carinate. Episternum strongly shagreened,
dull, with dense shallow punctures without rim, punctures
separated by 0.5–1 puncture diameters. Propodeum
shagreened and reticulate, dull, propodeal triangle
smooth, therefore indicated by this lack of reticulation.
Episternum, propodeum, scutum, and scutellum with long
whitish to yellowish hairs, these exceeding the length of
the scape. Legs dark, tarsal segments 3–5 lightened
Figures 41–46. Andrena (incertae sedis) prodigiosa n. sp. 41, Male prole; 42, male face; 43, male tergites; 44, male genitalia; 45,
male genitalia in prole; 46, male genitalia in reverse view, view of bifurcate penis valve.
14 T. J. Wood et al.

orange-red, pubescence whitish to yellowish. Wings hya-
line, venation amber, nervulus interstitial.
Metasoma. Tergites dark, shagreened, weakly shining,
apical margins lightened yellow-brown (Figure 43).
Tergal margins laterally with loose fringes of whitish
yellow hairs. Sternite 8 arched (Figure 45), covered in
golden hairs (Figure 46). Genitalia abnormal (Figure 44),
gonocoxites with strong apical teeth, ground colour
brown transitioning into translucent yellow as the gono-
coxites meet the gonostyli which are attened and shovel
Figures 47–54. Andrena (Pallandrena) christineae Dubitzky, 2006. 47, Female prole; 48, female tergites; 49, female labrum; 50,
female scopa; 51, male prole; 52, male tergites; 53, male genitalia in prole; 54, male genitalia.
Annales de la Société entomologique de France (N.S.) 15

shaped, coming to an apical point that is covered in
yellow hairs (Figure 44). Penis valve apically strongly
bifurcate apically (Figure 46).
Diagnosis. The genitalia of this specimen are unlike any
West Palearctic Andrena species, and even placing it into
a subgenus is difcult, as its combination of characters
does not t any of the current subgeneric concepts.
Structurally, the specimen is unremarkable except for
the extraordinary genitalia (Figures 44–46) that has
hugely enlarged and attened gonostyli with apical hair
tufts, and a penis valve that is apically bifurcate
(Figure 37). Sternite 8 is centrally arched and covered
in projecting golden hairs (Figures 45, 46). The genitalia
are reminiscent of Parandrenella with strong gonocoxites
leading to partially attened gonostyli with hair tufts on
their internal margin in combination with a penis valve
that forms a triangle basally. The subgenus also often has
sternite 8 ornamented with elaborate hair patterns (e.g.
Andrena nisoria Warncke, 1969, see Scheuchl et al. 2011
for good illustrations), but the gonostyli are never this
attened, the penis valve is never bifurcate, and the male
clypeus is always yellow in all species described to date.
We would hesitantly associate this species with
Parandrenella on the basis of the overall shape of the
genital construction, but we feel that placement in this
subgenus before a female can be located would be
presumptive.
Distribution. Known only from the locus typicus in
eastern Lebanon.
Etymology. The name prodigiosa (bizarre, prodigious,
amazing) was chosen because of the remarkable male
genitalia.
Andrena species of Lebanon
Andrena (Aenandrena) bisulcata Morawitz, 1877
Distribution. Central Europe eastwards to the Caucasus,
Turkey, and the Near East (Gusenleitner & Schwarz
2002).
Literature. Grace (2010): Lebanon.
Material examined. JORDAN: Pella env. [Tabaqat Fahl],
−80 m, 29.IV.2006, 1♀, leg. K. Deneš, OÖLM; North Shuna
environs, 29.IV.1996, 1♀, leg. Mi. Halada, OÖLM; South of
Irbid, 13.IV.2009, 1♂, leg. M. Snižek, OÖLM; 10 km N, NE
of Jarash [Jerash], 20.IV.2002, 1♀, leg. M. Snižek, OÖLM;
LEBANON: Mount Lebanon, Fidar, 25.IV.2017, 1♀, leg. Z.
Mahfouz, USEK; Jubayl [Byblos], 5.IV.2017, 1♂, leg. T.
Iskandar, USEK; SYRIA: Tartus, St. Georg-Kloster, 250 m,
3.IV.1988, 1♂, leg. L. Blank, MSC.
Notes. This species is reported as new for Jordan and
Syria.
Andrena (Brachyandrena) colletiformis Morawitz, 1874
Distribution. Southern Europe and North Africa to the
Near East and Central Asia (Gusenleitner & Schwarz
2002).
Literature. Grace (2010): Lebanon.
Material examined. JORDAN: Pella env. [Tabaqat Fahl],
−80 m, 29.IV.2006, 1♀, leg. K. Deneš, OÖLM; 30 km
WWN of Aljun, 30.IV.2006, 1♀, leg. F. Kantner, OÖLM;
LEBANON: Mount Lebanon, Chamis, Wadi Cheber,
372 m, 26.IV.2019, 1♀, leg. P. Rasmont, UMONS,
Chaetosciadium trichospermum; N Lebanon, Kfar Hay,
Monastere St. Youhanna Maroun, 327 m, 3.V.2017, 1♀,
leg. Boustani, TCFNR.
Notes. This species is reported as new for Jordan.
Andrena (Chlorandrena) exquisita Warncke, 1975 *
Distribution. Bulgaria, Turkey and Israel (Gusenleitner
& Schwarz 2002; Pisanty et al. 2018).
Material examined. JORDAN: Aljoun, 28.IV.2012, 1♀,
leg. M. Kafka, OÖLM; Aljun environs, 1.V.2006, 1♀, leg.
K. Deneš, OÖLM; 15 km W Jerash, Dibbin, 2.V.2006, 4♀,
leg. K. Deneš, OÖLM; LEBANON: Bekaa, Kefraya, Horch
El Kaser, 992 m, 9.IV.2019, 1♂, 3♀, leg. Boustani, TCFNR,
Crepis sancta; Bekaa, West Bekaa, Ammiq, 871 m, 27.
IV.2019, 1♀, leg. Boustani, TCFNR, Sonchus oleraceus;
N Lebanon, Tannourine El Tahta, Wadi Ain Al Raha,
900 m, 27.III.2017, 2♂, 2♀; 11.IV.2017, 1♂; 18.IV.2017,
3♀; 3.V.2017, 2♀, all leg. Boustani, TCFNR; N Lebanon,
Tannourine El Tahta, Al Mahbase, 893 m, 5.V.2019, 1♂,
leg. Boustani, TCFNR; N Lebanon, Fehta, El Biara, 1664 m,
11.V.2019, 1♀, leg. Boustani, TCFNR, Crepis sancta; N
Lebanon, Arz Tannourine, Gate, 1796 m, 11.V.2019, 1♂,
leg. Boustani, TCFNR; N Lebanon, Ehden, Horch, Trail 1,
1534 m, 2.V.2017, 2♀, leg. Boustani, TCFNR; SYRIA:
Latakia, Qaranjah, 750 m, 3.IV.1988, 11♂, leg. L. Blank,
MSC; Tartus, St. Georg-Kloster, 250 m, 3.IV.1988, 1♀, leg.
L. Blank, TJW; Aleppo, 500 m, Simeons-Kloster, 19.
IV.1988, 1♀, leg. L. Blank, MSC.
Notes. This species is reported as new for Jordan and
Syria. It was only recently reported from Israel for the
rst time (Pisanty et al. 2018), so it is clearly quite
widespread but previously unnoticed in the Levant.
Andrena (Chlorandrena) humabilis Warncke, 1965 *
Distribution. Balkans, Turkey and Israel (Gusenleitner &
Schwarz 2002)
16 T. J. Wood et al.

Material examined. JORDAN: 15 km W Jerash, Dibbin,
2.V.2006, 1♀, leg. K. Deneš, OÖLM; LEBANON: S
Lebanon, Saidoun, El Mrouj, 26.III.2017, 1♀, leg. A.
Baghdadi, SOILS; Saidoun, 10-11.II.2018, 1♂, 2♀, leg
A. Baghdadi, SOILS/TJW; SYRIA: Tartus, St. Georg-
Kloster, 250 m, 3.IV.1988, 1♀, leg. L. Blank, MSC.
Notes. This species is reported as new for Jordan and
Syria.
Andrena (Chlorandrena) humilis Imhoff, 1832 *
Distribution. Europe, north-western Africa, Turkey, and
eastwards into Russia and the Caucasus (Gusenleitner &
Schwarz 2002; Hazir et al. 2014).
Material examined. LEBANON: N Lebanon, Fehta, El
Biara, 1664 m, 24.IV.2018, 1♂, leg. Boustani, TJW,
Crepis spp.; 11.V.2019, 2♂, leg. Boustani, TCFNR,
Crepis sancta; N Lebanon, Arz Tannourine, Tannourine
Forest Reserve Outskirts, 1794 m, 6.V.2017, 5♂, leg.
Boustani, TCFNR, Geranium spp.; N Lebanon, Ehden,
Jord, 1983 m, 7.V.2019, 1♂, leg. Boustani, TCFNR,
Medicago spp.; N Lebanon, Arz Bcharre, Forest of the
Cedars of God, Forest Limit, 1873 m, 6.V.2019, 2♀, leg.
Boustani, TCFRN/TJW, Crepis spp.
Notes. The taxonomic situation surrounding A. humilis is
complicated. It is the most widespread of the West
Palearctic members of the Chlorandrena and indeed is the
only species of this subgenus found in northern Europe, where
it is easily identied due to a unique combination of
characters. However, in southern Europe the situation is
complicated by the presence of many similar species, and
also by A. humilis itself which is variable. The usually
consistently yellow male clypeus can show a reduced
yellow marking, or be entirely black (e.g. Switzerland,
Amiet et al. 2010). Because of this variation, Warncke
(1975a, 1975b) described several subspecies including
Andrena humilis indigena Warncke, 1975 and Andrena
humilis prunella Warncke, 1975 from central and south-
eastern Turkey respectively. Gusenleitner (1998) re-
examined the later form and elevated it to species rank as
Andrena tadauchii Gusenleitner, 1998 based on the clearly
divergent male genitalia.
Examination of material collected from high elevation
sites in Lebanon is difcult to immediately assign to a
species as whilst the male genitalia conform, they also
have entirely black clypei whereas the yellow marking is
reduced but present in subspecies such as A. h. indigena.
Given the variation in clypeal coloration across the range of
A. humilis, Lebanese specimens were assigned to this taxon
because of the strong similarity in genital construction. True
A. humilis is widespread in Turkey (Hazir et al. 2014), and
the seeming restriction of Lebanese specimens to high ele-
vation sites would be consistent with the idea that the
species is on the very edge of its southern range. Andrena
tadauchii is known from lower elevation sites Turkey, Syria,
and Israel (Gusenleitner & Schwarz 2002) and may well be
present in Lebanon, but this must be conrmed.
Andrena (Chlorandrena) orientana Warncke, 1965 *
Distribution. South-eastern Europe and Ukraine to the
Near East (Schwenninger et al. 2015).
Material examined. LEBANON: Bekaa, Kefraya,
1009 m, 27.IV.2019, 2♀, leg. Boustani, TCFNR/TJW,
Crepis spp.
Andrena (Chlorandrena) panurgimorpha
Mavromoustakis, 1957
Distribution. Greece, Turkey and the Caucasus, Cyprus,
Ukraine and Israel (Gusenleitner & Schwarz 2002).
Literature. Grace (2010): Lebanon.
Material examined. JORDAN: Pella env. [Tabaqat Fahl],
−80 m, 4.V.1995, 1♀; 29.IV.2006, 1♀, both leg. K.
Deneš, OÖLM; North Shuna environs, 29–30.IV.1996,
1♂, leg. Mi. Halada, OÖLM; 20 km N of Karak,
1000 m, 27.IV.2006, 1♂, leg. K. Deneš, OÖLM; NW of
Ajlun, 850 m, 20.V.2007, 4♀, leg. Z. Kevjal, OÖLM;
Ajlun, 30 km W Jarash, 2.VI.2006, 2♀, leg. Z. Kevjal,
OÖLM; Aljun environs, 1.V.2006, 2♂, 2♀, leg. F.
Kantner, OÖLM; Aljun environs, 5.V.1995, 1♀, leg. K.
Deneš, OÖLM; Ajlun env [Aljoun], 840 m, 1.V.2006, 5♂,
3♀, leg. K. Deneš, OÖLM; Irbid, Saham village, 19–25.
IV.2003, 1♂, 2♀, leg. I. Pljushtch, OÖLM; 10 km W
Jarash, 1.V.1996, 1♀, leg. Ma. Halada, OÖLM; 15 km
W Jerash, Dibbin, 2.V.2006, 2♀, leg. K. Deneš, OÖLM;
10 km N, NE of Jarash [Jerash], 20.IV.2002, 1♀, leg. M.
Snižek, OÖLM; 20 km NW of Amman, 420 m, 5.V.2006,
1♀, leg. K. Deneš, OÖLM; LEBANON: N Lebanon, Arz
Tannourine, Tannourine Forest Reserve Outskirts,
1739 m, 6.V.2017, 1♂, leg. Boustani, TJW; N Lebanon,
Jairoun, 29.V.2012, 2♀, leg. M. Kasparek, TCFNR;
SYRIA: Aleppo, 500 m, Simeons-Kloster, 19.IV.1988,
1♂, leg. L. Blank, TJW; 60 km S Damascus, Khahab,
14.V.1996, 1♀, leg. Ma. Halada, OÖLM.
Notes. This species is reported as new for Jordan and
Syria.
Andrena (Chlorandrena) pinkeunia Warncke, 1969 *
Distribution. Israel (Gusenleitner & Schwarz 2002).
Material examined. JORDAN: Pella env. [Tabaqat Fahl],
−80 m, 29.IV.2006, 1♀, leg. K. Deneš, TJW; Aljoun, 28.
IV.2012, 1♀, leg. M. Kafka, OÖLM; LEBANON: S
Lebanon, Saidoun, 8.V.2018, 1♀, leg. A. Baghdadi,
SOILS.
Annales de la Société entomologique de France (N.S.) 17

Notes. This species is reported as new for Jordan.
Andrena (Chrysandrena) hesperia Smith, 1853
Distribution. Circum-Mediterranean to Central Asia
(Gusenleitner & Schwarz 2002).
Literature. Grace (2010): Lebanon.
Material examined. JORDAN: Pella env. [Tabaqat Fahl],
−80 m, 29.IV.2006, 1♀, leg. K. Deneš, OÖLM; Aljoun,
28.IV.2012, 1♀, leg. M. Kafka, OÖLM; Aljun environs,
1.V.2006, 2♀, leg. K. Deneš, OÖLM; Irbid, Saham
village, 19–25.IV.2003, 4♂, 2♀, leg. I. Pljushtch,
OÖLM; 15 km W Jerash, Dibbin, 2.V.2006, 8♀, leg. K.
Deneš, OÖLM; LEBANON: Mount Lebanon, Ksaibe,
518 m, 18.IV.2019, 1♂, leg. Boustani, TCFNR, Sonchus
spp.; Mount Lebaon, Bentael, 369 m, 23.IV.2019, 1♀, leg.
Bous, Rasm, Neme, TCFNR, Crepis spp.; N Lebanon,
Tannourine El Tahta, Wadi Ain aA Raha, 900 m, 3.
V.2017, 1♀, leg. Boustani, TCFNR; N Lebanon, Fehta,
El Biara, 1664 m, 11.V.2019, 1♀, leg. Boustani, TCFNR,
Trifolium resupinatum; N Lebanon, Harissa: Chir El
Ribez, 1730 m, 31.V.2017, 2♀, leg. Boustani, TCFNR,
Crepis reuteriana; N Lebanon, Horch Ehden, Nabeh
Jouit, 1410 m, 20.V.2019, 1♂, 1♀, leg. Boustani,
TCFNR, Crepis spp.; N Lebanon, Bcharre, Dahr el
Adib, 2585 m, 27.V.2017, 2♂, leg. P. Rasmont &
Boustani, TCFNR; SYRIA: Latakia, Qaranjah, 750 m,
3.IV.1988, 1♂, leg. L. Blank, MSC.
Notes. This species is reported as new for Jordan and
Syria.
Andrena (Chrysandrena) merula Warncke, 1969 *
Distribution. Greece, Turkey and Israel (Gusenleitner &
Schwarz 2002).
Material examined. JORDAN: NW of Ajlun, 850 m, 20.
V.2007, 1♀, leg. Z. Kevjal, OÖLM; Aljoun, 27.IV.2012,
1♀, leg. M. Kafka, OÖLM; Irbid, Saham village, 19–25.
IV.2003, 1♂, 2♀, leg. I. Pljushtch, OÖLM; 15 km W
Jerash, Dibbin, 2.V.2006, 2♀, leg. K. Deneš, OÖLM;
10 km N, NE of Jarash [Jerash], 20.IV.2002, 2♀, leg.
M. Snižek, OÖLM; LEBANON: Mount Lebanon, Barja,
Marj Barja Daher, 350 m, 7.III.2017, 1♂, leg. Boustani,
TCFNR; N Lebanon, Tannourine el Tahta, Al Mahbase,
893 m, 25.IV.2019, 1♂, leg. Boustani, TCFNR; SYRIA:
40 km SW Hama, Masyat [Masyaf], 1.V.2000, 1♂, leg. F.
Kantner, OÖLM.
Notes. Very similar to A. hesperia but without orange
coloured legs and with slightly ner tergal sculpturing.
This species is reported as new for Jordan and Syria.
Andrena (Cordandrena) torda Warncke, 1965
Distribution. Greece, Turkey and Israel (Gusenleitner &
Schwarz 2002).
Literature. Grace (2010): Lebanon.
Material examined. JORDAN: North Shuna environs,
29.IV.1996, 1♀, leg. Mi. Halada, OÖLM; S of At Tala,
27–30.III.2013, 1♀, leg. M. Snižek, OÖLM; 20 km N of
Karak, 1000 m, 27.IV.2006, 1♀, leg. K. Deneš, OÖLM;
LEBANON: Bekaa, Quaraoun, 24.III.2013, 1♀, leg. M.
Kasparek, TJW; N Lebanon, South of Jairoun, 23.V.2012,
1♀, leg. M. Kasparek, TCFNR; SYRIA: Aleppo, SW,
700 m, 7.IV.1988, 1♀, leg. L. Blank, TJW.
Notes. This species is reported as new for Jordan and Syria.
Andrena (Cryptandrena) aruana Warncke, 1967 *
Distribution. Israel and probably Syria (Warncke 1967;
Gusenleitner & Schwarz 2002).
Material examined. LEBANON: Bekaa, Quaraoun, 24.
III.2013, 1♂, leg. M. Kasparek, TJW.
Notes. Extremely similar to A. monacha (below) in the
female sex, this species is best separated using the clearly
different male genitalia.
Andrena (Cryptandrena) brumanensis Friese, 1899
Distribution. Southern Europe to Turkey and the Near
East (Gusenleitner & Schwarz 2002).
Literature. Friese (1899): Mount Lebanon, Brumana
[Broummana, close to Beirut], 30.IV.1899, leg. F. Morice.
Friese listed the country as Syria, but the actual location is in
modern day Lebanon; Grace (2010): Lebanon.
Material examined. JORDAN: North Shuna environs,
29.IV.1996, 1♀, leg. Mi. Halada, OÖLM.
Notes. This species is reported as new for Jordan.
Andrena (Cryptandrena) monacha Warncke, 1965
Distribution. Greece, Turkey and Cyprus (Gusenleitner
& Schwarz 2002).
Literature. Grace (2010): Lebanon.
Material examined. LEBANON: Mount Lebanon,
Khaldah [Khalde], 18.IV.1973, 1♂, (no collector
information), TCFNR; SYRIA: Tartus, Sata, 10 km E,
300 m, 3.IV.1988, 1♂, leg. L. Blank, MSC.
Notes. This species has not yet been reported from Israel
or Jordan, and so it may be on the edge of its southern
range in Lebanon, and is reported as new for Syria.
18 T. J. Wood et al.

Andrena (Cryptandrena) ventricosa Dours, 1873
Literature. Grace (2010): Lebanon.
Material examined. LEBANON: N Lebanon, Jairoun,
29.V.2012, 1♀, leg. M. Kasparek, TCFNR.
Andrena (Euandrena) bicolor Fabricius, 1775 *
Distribution. Europe, north-western Africa, Turkey, the
Near East, and eastward to Central Asia (Gusenleitner &
Schwarz 2002)
Material examined. LEBANON: N Lebanon, Hadath El
Jebbe, Al Fouar, 1529 m, 12.IV.2019, 1♀, leg. Boustani,
TCFNR, Galium spp.
Notes. The Andrena bicolor-group, as well as the rest of
the subgenus Euandrena, are badly in need of an in-depth
revision across the Mediterranean region (Pisanty et al.
2018; Praz et al. 2019). There are several probably
undescribed Euandrena species present in Lebanon (see
Discussion), so here we only present one record of a
specimen that can be condently placed as A. bicolor.
The status of Euandrena in the Mediterranean basin is
likely to change dramatically in coming years.
Andrena (Euandrena) rutibialis limosa Warncke,
1969
Distribution. Jordan, Israel, Syria and Turkey
(Gusenleitner & Schwarz 2002).
Material examined. JORDAN: 15 km W Jerash, Dibbin,
750 m, 2.V.2006, 6♀, leg. K. Deneš, OÖLM/TJW; 20 km
N of Karak, 1000 m, 27.IV.2006, 1♂, leg. K. Deneš,
OÖLM; Ajlun S of Anjara, 27.IV.2002, 2♀, leg. M.
Snižek, OÖLM; Aljoun, 28.IV.2012, 1♂, 1♀, leg. M.
Kafka, TJW; Aljun environs, 5.V.1995, 1♀, leg. K.
Deneš, OÖLM; Aljun environs, 840 m, 1.V.2006, 2♂,
1♀, leg. K. Deneš & F. Kantner, OÖLM; Aljun Hills
env, 29.IV–1.V.2006, 1♀, leg. K. Deneš, OÖLM; Irbid,
Saham village, 25.IV.2003, 1♀, leg. I. Pljushtch, OÖLM/
TJW; LEBANON: Donnieh, 1600 m, 23.V.2012, 1♀, leg.
M. Kasparek, TJW; Donnieh, Wadi Cehennem oberth.
Quemmamine, 1393 m, 23.V.2012, 2♀, leg. M.
Kasparek, TJW; south of Jairoun, 1648 m, 23.V.2012,
1♀, leg. M. Kasparek, TJW.
Notes. A. rutibialis was originally described from
Jericho, in the modern-day West Bank, though the type
locality is listed as [Jordan] by Gusenleitner and Schwarz
(2002), so it is not actually clear if it has been previously
reported from Jordan or not. As its name suggests, one of
its dening characters are the red-coloured hind tibiae and
tarsi. Warncke later described the subspecies limosa with
type material from Turkey, Jordan and Israel (Warncke
1969a). This bee is darker, lacking the red-coloured legs
of the nominate type, but shares the shiny clypeus,
densely and clearly punctate tergites, and clear hair
fringes on the tergites. All examined material from
Jordan and Lebanon conformed to A. r. limosa, with no
specimens displaying red-coloured legs or tergites.
Andrena r. limosa is considered a valid taxon by some
authors (Rasmont et al. 2017), and further investigation
into A. rutibialis is needed to clarify the relationship
between these different colour forms.
Andrena (Euandrena) rufula Schmiedeknecht, 1883 *
Distribution. Southern Europe from France and Spain
eastwards to Ukraine and with a patchy distribution south
into Greece and Turkey (Gusenleitner & Schwarz 2002).
Material examined. LEBANON: N Lebanon, Horch
Ehden, Nabeh Jouit, 13.IV.2019, 1410 m, 1♀, leg.
Boustani, TCFNR, Crataegus cf monogyna; N Lebanon,
Horch Ehden, Nabeh Jouit, 24.IV.2019, 1337 m, 1♀, leg.
P. Rasmont, TCFNR, Salix libani; N Lebanon, Arz
Bcharre, Forest of the Cedars of God, 1913 m, 9.
V.2017, 2♀, leg. Boustani, TCFNR, Cotoneaster spp.
Notes. Andrena rufula was reported from the western
Taurus mountains of Turkey between Akseki and Beyşehir
at 1300 m (Warncke 1975a). Its presence in Lebanon whilst
unexpected is not unprecedented, and the high altitude
Lebanese sites are ecologically and climatically
comparable to the Taurus mountains. Andrena rufula is a
univoltine polylectic species that ies in the spring (Amiet
et al. 2010), and the ower records here would suggest that it
forages from owering broadleaved trees. Andrena rufula
can be an abundant visitor to owering apple trees in eastern
France, collecting pure loads of Rosaceae pollen (TJW,
unpublished data), so these observations are consistent
with this picture. This is the rst record of this species
from the Levant, but it may also be present at high altitude
sites in Syria and Israel.
Andrena (Fuscandrena) stenofovea Scheuchl &
Pisanty, 2018 *
Distribution. Israel (Pisanty et al. 2018).
Material examined. LEBANON: Mount Lebanon, Barja,
Marj Barja Daher, 350 m, 7.III.2017, 2♂, leg. Boustani,
TCFNR; Bekaa, Quaraoun, 24.III.2013, 1♀, leg. M.
Kasparek, TCFNR; N Lebanon, Tannourine El Tahta,
Wadi Ain El Raha, 900 m, 27.III.2017, 1♀, leg.
Boustani, TJW; SYRIA: Tartus, St. Georg-Kloster,
250 m, 3.IV.1988, 2♀, leg. L. Blank, MSC.
Annales de la Société entomologique de France (N.S.) 19

Notes. Originally placed in the Poecilandrena, this
species is better placed in the Fuscandrena (Pisanty
et al. 2020). This species is reported as new for Syria.
Andrena (Holandrena) forsterella Osytchnjuk, 1978
Distribution. The exact distribution of A. forsterella is
unclear because its putative rst generation was split off
as a distinct species A. wilhelmi Schuberth, 1995. The two
taxa have a similar distribution (Schuberth 1995) and A.
forsterella is probably distributed from Italy, through the
Balkans, to Turkey and further eastwards. True A.
forsterella has not been recorded from Israel, where
records conform to A. wilhelmi (Pisanty et al. 2018).
See Notes.
Literature. Grace (2010): Lebanon.
Material examined. LEBANON: Bekaa, Der el Ahmar,
989 m, 3.VII.2019, 1♀, leg. Boustani, TJW.
Notes. Andrena wilhelmi is a spring-ying species, on the
wing in March to June, whereas true A. forsterella ies
between June and August (Schuberth 1995). This record
represents the most southerly extent of its range, as re-
examination of specimens reported as A. forsterella by
Warncke (1969) collected in March and April conrm
their identity as A. wilhelmi (Pisanty et al. 2018). Both
taxa are likely to be present in Lebanon.
Andrena (Holandrena) labialis (Kirby, 1802) *
Distribution. Europe and north-western Africa eastwards
to Turkey and Central Asia, south to Israel (Gusenleitner
& Schwarz 2002; Pisanty et al. 2018).
Material examined. LEBANON: Mount Lebanon, Horch
el Barouk, Chouf Biosphere Reserve, 1678 m, 2.
VII.2019, 1♀, X. van Achter, VXA, Medicago spp.; N
Lebanon, Hadath El Jebbeh, Chemin Wadi Ain El Raha,
1519 m, 29.VI.2017, 1♂, 1♀, leg. Boustani, TCFNR,
Ononis natrix; N Lebanon, Harissa: Al Jawar, 1738 m,
28.VI.2017, 1♂, leg. Boustani, TCFNR; N Lebanon, Arz
Tannourine Gate area, 1754 m, 23.VI.2018, 1♂, 1♀, leg.
Boustani, TCFNR, Allium phanerantherum; N Lebanon,
Hadath El Jebbe, Border of the Cedar forest, 1632 m, 5.
VII.2019, 1♀, leg. Boustani, TCFNR, Stachys cretica; N
Lebanon, Tannourine Reserve Trail 4, 6.VII.2017,
1747 m, 1♂, leg. Boustani, TCFNR, Campanula stricta;
N Lebanon, Horch Ehden, Nabeh Jouit, 1410 m, 27.
VI.2019, 1♀, leg. Boustani, TCFNR, Stachys cretica;
Arz Bcharre, Forest of the Cedars of God, 22.VI.2017,
1815 m, 22.VI.2017, 1♀, leg. Boustani, TCFNR, Vicia
tenuifolia; Arz Bcharre, Bcharre Reforestation Area,
1993 m, 22.VI.2017, 1♂, 2♀, leg. Boustani, TCFNR,
Medicago spp.; 2.VII.2019, 1♂, leg. G. Ghisbain,
UMONS; N Lebanon, Bcharre, 2042 m, 12.VII.2019,
1♀, leg. X. van Achter, XVA, Vicia spp.; Bekaa, Ainata,
1556 m, 30.V.2017, 5♂, leg. P. Rasmont, TCFNR.
Notes. Andrena labialis was recently reported from Israel
for the rst time from Mount Hermon at comparable
altitudes (1600–1650 m, Pisanty et al. 2018) to the
Lebanese sites reported here.
Andrena (Holandrena) variabilis Smith, 1853
Distribution. LEBANON: Southern Europe and north-
western Africa eastwards to Turkey, the Near East, and
into Central Asia (Gusenleitner & Schwarz 2002).
Literature. Grace (2010): Lebanon.
Material examined. Mount Lebanon, Laqlouq, 1752 m,
30.VI.2019, 1♀, leg. G. Ghisbain, UMONS.
Andrena (Hoplandrena) trimmerana (Kirby, 1802) *
Distribution. The true distribution of A. trimmerana is
obscured by long-standing taxonomic confusion with
another member of the Hoplandrena that has been
referred to as A. carantonica Pérez, 1902, but it appears
to be distributed across Europe into north-western Africa,
Turkey and Israel (Gusenleitner & Schwarz 2002; Pisanty
et al. 2018).
Material examined. LEBANON: N Lebanon, Arz Bcharre,
Forest of the Cedars of God, 1913 m, 9.V.2017, 1♂, leg.
Boustani, TCFNR; Bekaa, Donnieh, Wadi Cehennem, (no
date), 1♀, leg. M. Kasparek, TCFNR; Bekaa, Ras Baalbeck,
5.V.2018, 1♂, leg. E. Harran, USEK.
Notes. True A. trimmerana is best and most condently
identied from spring males which have unidentate
mandibles (lacking an internal mandibular tooth) and
also possess a genal spine.
Andrena (Lepidandrena) statusa Gusenleitner, 1998 *
Distribution. Israel and Turkey (Gusenleitner & Schwarz
2002).
Material examined. LEBANON: N Lebanon,
Tannourine El Tahta, Wadi Ain El Raha, 936 m, 11.
IV.2017, 1♂, leg. Boustani, TJW.
Notes. Both A. statusa and A. elisaria Gusenleitner, 1998
(Turkey) were described in the same publication and from
the same type locality and date. They differ mostly by the
colour of the integument. Pisanty et al. (2018) argue that
they likely represent variation within a single species, as
both forms can be found together in Israel. For now, only
one specimen has been found in Lebanon so we can make
no comment as to the validity of this conclusion.
20 T. J. Wood et al.

Andrena (Margandrena) krausiella Gusenleitner, 1998
Distribution. Israel and Jordan (Gusenleitner & Schwarz
2002; Al-Ghzawi et al. 2006).
Literature. Grace (2010): Lebanon.
Material examined. LEBANON: S Lebanon, Saidoun,
20.I.2018, 2♀, leg. A. Baghdadi, SOILS/TJW.
Andrena (Melanapis) fuscosa Spinola, 1838 *
Distribution. Circum-Mediterranean to Central Asia
(Gusenleitner & Schwarz 2002).
Material examined. LEBANON: Bekaa, American
University of Beirut farm, 15.IV.1962, 1♂, leg. K.
Bedirian, AUB; 12.V.1962, 1♀, leg. S. Khan, AUB;
SYRIA: Aleppo, 500 m, Simeons-Kloster, 19.IV.1988,
3♂, leg. L. Blank, MSC.
Notes. Seemingly not previously reported from Syria.
Andrena (Melandrena) albopunctata (Rossi, 1792) *
Distribution. Circum-Mediterranean to Central Asia
(Gusenleitner & Schwarz 2002).
Material examined. LEBANON: S Lebanon, Sidon
[Saida], 25.V.1981, 1♀, (no collector information),
TCFNR; Mount Lebanon, Chhim, Khallat Chiim, 25.
IV.1981, 1♂, (no collector information), AUB; Bekaa,
American University of Beirut farm, 19.IV.1962, 1♀,
leg. N Samman; 27.IV.1962, 1♀, leg. K. Bedirian; 26.
V.1962, 1♀, leg. Z. Rai; 20.IV.1964, 1♀, leg. G.
Siddloul, all AUB; Bekaa, Baalbeck, Hoch Sneid, 21.
V.1964, 1♀, leg. B. Ayyash, AUB.
Andrena (Melandrena) elmaria Gusenleitner, 1998
Distribution. Cyprus, Israel, Syria, and Turkey
(Gusenleitner & Schwarz 2002).
Literature. Grace (2010): Lebanon.
Material examined. LEBANON: N Lebanon,
Tannourine El Tahta, Al Mahbase, 893 m, 12.IV.2019,
1♀, leg. Boustani, TJW; 5.V.2019, 1♀, leg. Boustani,
TCFNR; N Lebanon, Arz Tannourine Gate area,
1754 m, 2.IV.2018, 1♂, leg. Boustani, TCFNR; N
Lebanon, Ehden, Nabeh Jouit, 1336 m, 24.IV.2019, 1♀,
leg. P. Rasmont, UMONS, Salix cf libani.
Andrena (Melandrena) limata Smith, 1853 *
Distribution. Europe and north-western Africa to the
Near East, Turkey, and Central Asia (Gusenleitner &
Schwarz 2002).
Material examined. LEBANON: Mount Lebanon,
Falougha, 17.VII.1975, 1♂, (no collector information),
AUB; Bekaa, American University of Beirut farm, 2.
IV.1963, 1♀, leg. Babikir; 10.VI.1971, 1♂, leg. Anwar;
11.IV.1980, 1♀, leg. A. Fakher, all AUB.
Andrena (Melandrena) morio Brullé, 1832
Distribution. Europe, North Africa, the Middle East, and
into Central Asia (Gusenleitner & Schwarz 2002).
Literature. Mavromoustakis (1962): N Lebanon, Near
Becharré [Bcharre], 19.VI.1960, 1♀; Focke, 20.VI.1960,
2♀; N Lebanon, Kadisha river, 20.VI.1960, 1♀; Grace
(2010): Lebanon.
Andrena (Melandrena) nigroaenea (Kirby, 1802)
Distribution. Europe, Mediterranean, Turkey, the Near
East, and into Central Asia (Gusenleitner & Schwarz 2002).
Literature. Mavromoustakis (1962): N Lebanon, Near
Becharré [Bcharre], 27.VI.1960, 1♀; N Lebanon, Kadisha
river, 21-26.VI.1960, 6♀; Grace (2010): Lebanon.
Material examined. LEBANON: N Lebanon, Bcharre,
1900 m, 14.VII.2019, 1♀, leg. X. van Achter, XVA;
Mount Lebanon, Hboub, 29.III.2017, 1♀, leg. E. Harran,
USEK; N Lebanon, Tannourine El Tahta, Wadi Ain El Raha,
900 m, 1♂, leg. Boustani, TCFNR; N Lebanon, Hadath El
JEbbEh, Road to Wadi Al Fouar, 1553 m, 28.VI.2018, 1♀,
leg. Boustani, TCFNR; N Lebanon, Hadath el Jebbe, Al
Fouar, 1529 m, 12.VI.2019, 1♂, leg. Boustani, TCFNR; N
Lebanon, Tannourine Reserve, Trail 4, 1781 m, 3.VI.2019,
1♀, leg. Boustani, TCFNR; N Lebanon, Jairoun, 29.V.2012,
1♀, leg. M. Kasparek, TCFNR.
Andrena (Melandrena) pyropygia Kriechbaumer,
1873
Distribution. Eastern Mediterranean to Ukraine and into
Central Asia (Gusenleitner & Schwarz 2002).
Literature. Mavromoustakis (1962): Mount Lebanon,
Baabdate, 24–25.V.1953, 2♂, 1♀; S Lebanon, Djezzine
[Jezzine], 20.V.1953, 1♂; Grace (2010): Lebanon.
Andrena (Melandrena) thoracica (Fabricius, 1775)
Distribution. Europe, North Africa, the Near East, and
into Central Asia (Gusenleitner & Schwarz 2002).
Literature. Mavromoustakis (1962) (as A. t. kotschyi
Mavromoustakis, 1953): N Lebanon, Near Becharré
[Bcharre], 19–20.VI.1960, 3♀; N Lebanon, Kadisha
river, 25.VI.1960, 1♀; N Lebanon, Cedars [Arz
Bcharre], 4.VII.1960, 1♀; Grace (2010): Lebanon.
Material examined (A. t. kotschyi). LEBANON: Mount
Lebanon, Berbara, 10.IV.1961, 1♀, leg. H. Nasr, AUB;
Bekaa, American University of Beirut farm, 10.V.1961,
1♀, leg. R. Hajj, AUB; (A. thoracica sensu strictu)
Annales de la Société entomologique de France (N.S.) 21

Bekaa, American University of Beirut farm, 29.III.1964,
1♀, leg. Taylor, AUB.
Notes. There are outstanding taxonomic issues
surrounding A. thoracica and its described subspecies.
In typical nominate A. thoracica from northern Europe,
the hind tibial spurs are black, but in Mediterranean forms
such as A. t. kotschyi the hind tibial spurs are light red or
amber, similar to A. limata. Specimens from the same site
in Lebanon (Bekaa, American University of Beirut farm)
produced specimens with both colour forms. Andrena
thoracica could benet from molecular investigation
across its range.
Andrena (Melittoides) melittoides Friese, 1899 *
Distribution. Israel and Turkey (Gusenleitner & Schwarz
2002).
Literature. Friese (1899) described this species from
Jerusalem, but also listed below this “Syria”. It is not
clear exactly what he meant by this as he did not detail
specic specimens, and in the same paper he made
reference to “Jaffa (Syria)” (p. 343) which today is in
modern Israel. However, we can denitively report that
the species is present in Lebanon based on contemporary
material.
Material examined. LEBANON: Mount Lebanon, Wadi
Chahrour, 22.V.2009, 1♂, leg. N Nemer, USEK.
Andrena (Micrandrena) alfkenella Perkins, 1914 *
Distribution. Morocco and Europe to Turkey
(Gusenleitner & Schwarz 2002).
Material examined. LEBANON: Mount Lebanon, Chouf
Biosphere Reserve, Barouk Gate, 1428 m, 16.V.2019, 1♀,
leg. Boustani, TCFNR, Peltaria angustifolia; N Lebanon,
Tannourine Reserve, Trail 4, 1781 m, 30.VI.2019, 1♀,
leg. Boustani, TCFNR, Euphorbia spp.; N Lebanon, Arz
Tannourine, Main gate, 1796 m, 4.VII.2019, 14♀, leg.
Boustani, X. van Achter, & G. Ghisbain, TCFNR/TJW/
UMONS/XVA, Chaerophyllum aurantiacum; N Lebanon,
Arz Tannourine, reserve entrance, 1797 m, 12.VII.2019,
3♀, leg. X. van Achter, VXA, Chaerophyllum
aurantiacum; N Lebanon, Horch Ehden, Nabeh Jouit,
1336 m, 24.IV.2019, 2♀, leg. Boustani, TCFNR; N
Lebanon, Ehden, Horch Ehden, 1567 m, 1♀, leg.
Boustani & W. Yammine, TCFNR.
Notes. Andrena alfkenella is widespread throughout
Europe, extending to mountains in Turkey including the
Taurus Mountains. The species is bivoltine, though the
second generation is much more abundant and easier to
nd, showing a strong preference for Apiaceae pollen
(TJW, unpublished data). Its presence at altitude in
northern Lebanon is consistent with the trend shown in
other species more typically found in Europe.
Andrena (Micrandrena) alfkenelloides Warncke, 1965 *
Distribution. Balkans to Turkey and the Near East
(Gusenleitner & Schwarz 2002).
Material examined. LEBANON: N Lebanon, Tannourine
El Tahta, Wadi Ain El Raha, 900 m, 24–27.III.2017, 4♀, leg.
Boustani, TCFNR/TJW.
Andrena (Micrandrena) lindbergella Pittioni, 1950
Literature. Grace (2010): Lebanon.
Material examined. LEBANON: N Lebanon, Bcharre,
Dahr El Adib, 2585 m, 27.V.2017, 3♂, leg. P. Rasmont &
Boustani, TCFNR/TJW; N Lebanon, Bcharre, Dahr El
Adib, 2437 m, 8.VI.2017, 2♀, leg. Boustani, TCFNR/
TJW; N Lebanon, Arz Bcharre, Forest of the Cedars of
God, Forest Limit, 1897 m, 20.V.2019, 1♂, leg. Boustani,
TCFNR, Alyssum spp.; N Lebanon, Tannourine, Jabal Al
Mnaitra, 2469 m, 28.V.2017, 1♂, leg. P. Rasmont &
Boustani, TCFNR, Ranunculus demissus.
Notes. Recently recorded from Israel for the rst time
from Mount Hermon, also at altitude (1950–2200 m,
Pisanty et al. 2018). With a locus typicus in the Troodos
mountains in Cyprus, this is clearly an alpine species.
Andrena (Micrandrena) magunta Warncke, 1965 *
Distribution. Eastern Europe through the Balkans to
Turkey and Israel (Gusenleitner & Schwarz 2002;
Pisanty et al. 2018).
Material examined. LEBANON: Mount Lebanon,
Jamhour, 2.IV.1973, 1♀, (no collector information),
TCFNR; Bekaa, Quaraoun, 24.III.2013, 1♀, leg. M.
Kasparek, TCFNR; N Lebanon, Tannourine El Tahta,
Wadi Ain El Raha, 900 m, 5.V.2017, 1♀, leg. Boustani,
TJW; 11.V.2017, 2♂, leg. Boustani, TCFNR/TJW,
Carduus argentatus.
Andrena (Micrandrena) minutula (Kirby, 1802) *
Distribution. North-western Africa, through Europe to
Cyprus and Turkey (Gusenleitner & Schwarz 2002).
Material examined. LEBANON: N Lebanon, Hadath El
Jebbe, Al Fouar, 1529 m, 12.IV.2019, 1♀, leg. Boustani, TJW.
Notes. Another more typically ‘European’ species found
at altitude.
Andrena (Micrandrena) minutuloides Perkins, 1914 *
Distribution. Morocco and Europe to Turkey and the
Caucasus (Gusenleitner & Schwarz 2002).
22 T. J. Wood et al.

Material examined. LEBANON: N Lebanon, Ehden, Ain
El Naasa, 1598 m, 22.V.2019, 1♀, leg. Boustani, TJW.
Notes. Another more typically ‘European’ species found
at altitude.
Andrena (Micrandrena) oedicnema Warncke, 1975 *
Distribution. Greece and Turkey to Israel (Gusenleitner
& Schwarz 2002; Pisanty et al. 2018).
Material examined. JORDAN: Irbid, Saham villiage,
19–25.IV.2003, 3♀, leg. I. Pljushtch, OÖLM/TJW;
LEBANON: Mount Lebanon, Barja, Marj Barja Daher,
350 m, 7.III.2017, 1♂, leg. Boustani, TCFNR; Mount
Lebanon, Marej Barja, Kaleet El Besten, 358 m, 26.
IV.2019, 1♀, leg. Boustani, TCFNR; N Lebanon,
Tannourine El Tahta, Wadi Ain El Raha, 900 m, 24.
III.2017, 1♂, leg. Boustani, TCFNR; SYRIA: Tartus, St.
Georg-Kloster, 250 m, 3.IV.1988, 1♂, leg. L. Blank,
TJW; Latakia, Saladinburg [Citadel of Saladin], 900 m,
4.IV.1988, 1♂, leg. L. Blank, TJW.
Notes. Recently reported from Israel for the rst time
(Pisanty et al. 2018), this species is reported as new to
Jordan and Syria.
Andrena (Micrandrena) rugothorace Warncke, 1965 *
Distribution. Italy and the Balkans to Turkey and Israel
(Gusenleitner & Schwarz 2002; Pisanty et al. 2018).
Material examined. LEBANON: N Lebanon, Tannourine
El Tahta, Wadi Ain El Raha, 900 m, 11–18.IV.2017, 1♂, 1♀,
leg. Boustani, TCFNR; N Lebanon, Tannourine El Tahta, Al
Mahbase, 893 m, 25.IV–5.V.2019, 4♀, leg. Boustani,
TCFNR/TJW.
Notes. Predominantly found at altitude, as is the case in
Israel (Mount Hermon 1500 m, Pisanty et al. 2018).
Andrena (Micrandrena) simontornyella corpana
Warncke, 1965
Distribution. Southern Europe and north-western Africa
to Turkey and the Near East. The south-eastern form
found in Greece, Turkey, and the Levant is smaller and
was described as A. corpana (Warncke 1965).
Literature. Grace (2010): Lebanon.
Material examined. LEBANON: Bekaa, Qaraoun Lake,
24.III.2013, 1♀, leg. M. Kasparek, TJW.
Andrena (Micrandrena) spreta Pérez, 1895 aggregate
Distribution. The Andrena spreta complex is
taxonomically challenging and controversial. In a broad
sense, the species is found around the Mediterranean, and
north into central and northern Europe (Gusenleitner &
Schwarz 2002).
Literature. Grace (2010): Lebanon.
Material examined. LEBANON: N Lebanon,
Tannourine El Tahta, Wadi Ain El Raha, 934 m, 11.
V.2017, 1♂, leg. Boustani, TCFNR.
Notes. The taxonomic status of A. spreta and its
subspecies is unclear. It is currently in the process of
being revised and is likely to be broken up into multiple
species in the future. The form likely to be present in
Lebanon was described as A. s. scirpacea Warncke, 1975
from Turkey.
Andrena (Micrandrena) tiaretta Warncke, 1974
Distribution. North Africa to Israel and Syria
(Gusenleitner & Schwarz 2002).
Literature. Grace (2010): Lebanon.
Material examined. LEBANON: N Lebanon, Tannourine
El Tahta, Wadi Ain El Raha, 901 m, 11.IV.2017, 1♀, leg.
Boustani, TCFNR; N Lebanon, Tannourine El Tahta, Wadi
Ain El Raha, 878 m, 18.IV.2017, 1♀, leg. Boustani, TJW; N
Lebanon, Tannourine El Tahta, Al Mahbase, 893 m, 5.
V.2019, 1♀, leg. Boustani, TCFNR.
Notes. Andrena tiaretta can be differentiated from other
Near Eastern Micrandrena by the sculpturing of the
scutum, and by the male genitalia. Kratochwil (2015)
revised the A. tiaretta group, describing A. orientalis
Kratochwil, 2015 from Israel and Syria. We do not
follow this interpretation as the characters described for
separating the putative taxa are extremely slight, and we
take the position that these constitute acceptable variation
within a single species concept.
Andrena (Micrandrena) tringa Warncke, 1973 *
Distribution. Eastern Europe to Turkey and Israel
(Gusenleitner & Schwarz 2002; Pisanty et al. 2018).
Material examined. LEBANON: Mount Lebanon, Chouf
Biosphere Reserve, Maaser El Chouf Gate, 1726 m, 16.
V.2019, 1♀, leg. Boustani, TCFNR; Mount Lebanon,
Chouf Biosphere Reserve, Barouk Gate, 1428 m, 16.
V.2019, 1♀, leg. Boustani, TCFNR; Mount Lebanon,
Barouk, Chouf Biosphere Reserve, 1690 m, 4.VII.2019,
1♀, leg. X. van Achter, VXA; N Lebanon, Tannourine El
Tahta, Al Mahbase, 893 m, 12.IV.2019, 1♀, leg. Boustani,
TJW; N Lebanon, Harissa, Al Jawar, 1758 m, 11.V.2019,
1♂, 2♀, leg. Boustani, TCFNR; N Lebanon, Hadath El
Jebbeh, Road to Wadi Al Fouar, 1553 m, 6.VI.2018, 1♀,
leg. Boustani, TCFNR; N Lebanon, Tannourine Reserve,
Trail 4, 1781 m, 13.V.2019, 1♀, leg. Boustani, TCFNR; N
Annales de la Société entomologique de France (N.S.) 23

Lebanon, Arz Tannourine Gate area, 1754 m, 2.IV.2018,
2♀, leg. Boustani, TCFNR; N Lebanon Arz Bcharre,
Forest of the Cedars of God, Forest Limit, 1897 m, 20.
V.2019, 4♂, 5♀, leg. Boustani, TCFNR/TJW; N Lebanon,
Arz Bcharre, Forest of the Cedars of God Reforestation
Area, 1933 m, 18.IV.2018, 1♀, leg. Boustani, TCFNR.
Notes. Predominantly found at altitude, as is the case in
Israel (Mount Hermon 1300 m, Mount Meron 1000 m,
Pisanty et al. 2018).
Andrena (Nobandrena) anatolica Alfken, 1935
Literature. Grace (2010): Lebanon.
Distribution. Greece, Turkey, and the Near East to
Russia and the Caucasus (Gusenleitner & Schwarz
2002; Pisanty et al. 2018).
Material examined. LEBANON: N Lebanon, Arz
Bcharre, Forest of the Cedars of God, 1883 m, 27.V.2018,
3♀, leg. Boustani, TCFNR, Vicia tenuifolia; N Lebanon,
Arz Bcharre, Forest of the Cedars of God, Forest Limit,
1897 m, 2♂, 2♀, leg. Boustani, TCFNR, Erysimum spp.
Notes. This species was collected in association with
Erysimum (Brassicaceae), a likely pollen host as some
members of the Nobandrena are known to be specialised
on Brassicaceae (Amiet et al. 2010; TJW unpublished
data). This species was recently reported from Israel for
the rst time, also at a high altitude site (Mount Hermon
1500 m, Pisanty et al. 2018).
Andrena (Nobandrena) asiatica Friese, 1921
Distribution. Turkey and Lebanon (Schuberth et al.
2001); Grace (2010) Lebanon.
Literature. Schuberth et al. (2001): N Lebanon, Cedars
[Arz Bcharre], 10–23.VI.1962, 1♀, Sw, 1♂, Coll.
Schmiedeknecht, Zoological Museum Berlin.
Andrena (Notandrena) ungeri Mavromoustakis, 1952 *
Distribution. The map presented by Gusenleitner and
Schwarz (2002) presents incorrect data. Andrena ungeri
is found from south-eastern Europe through the Balkans
and Turkey to Israel and Syria (Warncke 1969a; Hazir
et al. 2014).
Material examined. JORDAN: Kerak, Mazra’a, −390 m,
19.III.1988, 18♀, MSC; LEBANON: Mount Lebanon,
Arz Al Barouk, 19.V.1972, 1♀, (no collector
information), AUB; Mount Lebanon, Dahr El Baïdar, 8.
III.1973, 1♀, leg. E. Baram, AUB; Bekaa, Bawarij, 10.
V.1975, 1♀, (no collector information), AUB; Bekaa,
Anjar, 27.IV.2016, 1♀, leg. Boustani, Y. Zgheib, TCFNR.
Notes. This species is reported as new for Jordan.
Andrena (Opandrena) schencki Morawitz, 1866
Distribution. Europe to Turkey, the Near East, and
Central Asia (Gusenleitner & Schwarz 2002).
Literature. Grace (2010): Lebanon.
Material examined. LEBANON: Beyrouth [Beirut], 1.
V.1979, 1♀, leg. Japrin, TCFNR.
Andrena (Orandrena) gallinula Warncke, 1975 *
Distribution. Turkey and Armenia (Gusenleitner &
Schwarz 2002).
Material examined. LEBANON: N Lebanon, Horch
Ehden, Nabeh Jouit, 1410 m, 20.V.2019, 1♀, leg.
Boustani, TJW, Diplotaxis spp.; N Lebanon, Horch
Ehden, Ain El Naasa, 1560 m, 22.V.2019, 2♀, leg.
Boustani, TCFNR, Diplotaxis spp.; SYRIA: 400 m NN
Homs E 20 km, 1.IV.1988, 1♀, leg. S.M. Blank, TJW.
Notes. Close to A. oralis Morawitz, 1876 but smaller.
Most similar to A. acrana Warncke, 1967 but can be
separated by the more densely shagreened scutum. This
species is reported as new for Syria.
Andrena (Orandrena) garrula lomvia Warncke, 1969 *
Distribution. Turkey and Israel (Gusenleitner & Schwarz
2002).
Material examined. JORDAN: Aljoun, 28.IV.2012, 1♀,
leg. M. Kafka, OÖLM; LEBANON: Mount Lebanon,
Barja, Marj Barja Daher, 350 m, 31.I.2016, 2♂, leg.
Boustani, Y. Zgheib, TCFNR; N Lebanon, Tannourine
El Tahta, Wadi Ain al Raha, 900 m, 24.III.2017, 4♀,
leg. Boustani, TCFNR/TJW; SYRIA: Tartus, St. Georg-
Kloster, 250 m, 3.IV.1988, 4♀, leg. L. Blank, MSC/TJW.
Notes. This taxon is reported as new for Jordan and Syria.
Andrena (Pallandrena) christineae Dubitzky, 2006 *
Distribution. Turkey (Ağrı; Hakkâri) and Iran
(Kermanshahan, Dubitzky 2006).
Material examined. LEBANON: N Lebanon, Arz
Bcharre Forest of the Cedars of God Reforestation Area,
1933 m, 18.IV.2018, 2♂, 2♀, leg. Boustani, TCFNR/
TJW, Geranium libanoticum; Arz Bcharre, Forest of the
Cedars of God, Forest Limit, 1897 m, 20.V.2019, 1♂, leg.
Boustani. Allotype male deposited at the OÖLM.
Description of male. Body length 11 mm (Figure 51).
24 T. J. Wood et al.

Head. Black, clearly wider than long. Clypeus black, broad,
strongly domed, evenly reticulate with reticulation forming
weak lateral carinae, underlying surface impunctate, weakly
shining. Process of labrum twice as broad as long, only very
shallowly raised above the rest of the labrum but smooth and
shiny and therefore well differentiated from the clearly
punctured remaining surface area. Face, gena, vertex, and
scape with long white hairs, those on the lower part of the
gena clearly exceeding the length of the scape. Gena slightly
wider than the width of the compound eye. Antennae uni-
formly dark, segment 3 equalling 4 + 5. Ocelloccipital
distance wide, twice width of posterior ocellus.
Mesosoma. Scutum and scutellum weakly shagreened,
weakly shining, moderately but very shallowly punctate,
punctures separated by 1–2 puncture diameters. Pronotum
non-carinate. Episternum and propodeum shagreened
with raised moderately strong reticulation, propodeal tri-
angle clearly differentiated from the rest of the propo-
deum by a change in surface sculpture from moderate to
shallow reticulation. Legs dark, only tarsal claws slightly
lightened to red, pubescence white. Wings hyaline, vena-
tion dark brown to amber apically, nervulus interfurcal.
Metasoma. Tergites dark, margins lightened orange to
yellow, clearly hyaline apically (Figure 52). Tergal inte-
gument very subtly shagreened, predominantly shining,
weakly punctured throughout, punctures separated by 1–2
puncture diameters. S8 long and narrow, apically nar-
rowly emarginate, therefore clearly forked. Genitalia
long (Figure 53), generally reminiscent of members of
the Ulandrena due to the widely parting gonocoxites and
the large penis valve with an apical blister (Figure 54),
though without apical gonocoxal teeth. Gonostyli narrow,
forming projecting points that stand out in prole
(Figure 53).
Notes. Described only from the female, A. christineae was
known from high-elevation mountain sites in eastern Turkey
and western Iran. Females can be easily recognised as
Pallandrena because of the deep emargination in the
labrum (Figure 49) and the plumose scopa (Figure 50).
Andrena christineae females can be recognised easily
because of the red coloration of the tergites (Figures 47,
48, Dubitzky 2006). Where altitudes were recorded,
specimens were collected from 2050–2450 m (Turkey,
Hakkari, Mt. Sat, 10.VI.1981), 1600 m (Turkey, Ağrı,
10 km N Tutak, 28.V.1980), and 1900–2000 m (Iran,
Kermanshahan, Buchan, 19.V.1975, Dubitzky 2006). Sites
in northern Lebanon are therefore at an ecologically
consistent elevation. The bee may also be present in
Turkish mountains closer to Lebanon, linking the
populations together, and given this specialised ecological
niche, the species is probably under-recorded.
Lebanese material represents the earliest recorded
ight date of 18.IV for this species, but even this early
in the season male material shows extensive wing wear
(Figure 51), perhaps explaining why males were not
captured by previous collectors. Andrena christineae, as
well as other Pallandrena species such as A. byrsicola
Schmiedeknecht, 1900, are likely to be a specialised on
Geranium or other member of the Geraniaceae, in part
because of the extraordinary plumose scopa that denes
the subgenus Pallandrena (Figure 50, TJW unpublished
data). This scopa type is reminiscent of the plumose
scopa of the specialist A. hattorana (Fabricius, 1775)
that is presumably an adaptation to collecting the simi-
larly large grains of Knautia (Dipsacaceae) that can reach
100 μm in diameter. The scopal hairs of specialised bees
often reect the characteristics of the particular pollen
type that they specialise on, with long simple hairs in
bees specialised on large grains from Onagraceae or
Cucurbitaceae or dense plumose or branched hairs in
bees specialised on small grains from Asteraceae
(Linsley 1958; Thorp 1979; Portman & Tepedino 2017).
To date, morphological adaptations to pollen harvesting
in Andrena have received little attention, though see
LaBerge (1987) and Dubitzky et al. (2010) for the sub-
genera Scoliandrena and Hamandrena, respectively.
Andrena (Pallandrena) pallidicincta Brullé, 1832 *
Distribution. From the Balkans to Turkey and Israel
(Gusenleitner & Schwarz 2002).
Material examined. LEBANON: N Lebanon, Ehden,
Horch Ehden, Gate area, 1424 m, 2.V.2017, 6♂, leg.
Boustani, TCFNR/TJW, Geranium spp.; N Lebanon,
Donnieh, 23.V.2012, 1♀, leg. M. Kasparek, TCFNR;
SYRIA: 50 km NE Tartus, Banyas, 1.V.2000, 1♀, leg.
F. Kantner, OÖLM.
Notes. Probably a specialist on Geranium like A. christi-
neae based on the oral associations and the remarkable
plumose scopa that are typical of the Pallandrena.
Reported as new for Syria.
Andrena (Plastandrena) pilipes Fabricius, 1781
Distribution. Taking a broad interpretation, A. pilipes is
found around the Mediterranean into Central Asia
(Gusenleitner & Schwarz 2002).
Literature. Grace (2010): Lebanon.
Material examined. LEBANON: Bekaa, Der el Ahmar,
3.VII.2019, 1♂, leg. G. Ghisbain, UMONS.
Notes. Andrena pilipes may contain additional taxa such
as A. spectabilis Smith, 1853 which was reported from
Annales de la Société entomologique de France (N.S.) 25

Jordan (Al-Ghzawi et al. 2006). For now, we take a broad
approach for this taxon.
Andrena (Poecilandrena) bytinskii Warncke, 1969 *
Distribution. Israel and (unpublished) Turkey (Pisanty
et al. 2018)
Material examined. JORDAN: Aljun environs, 1.
V.2006, 1♀, leg. F. Kantner, OÖLM; Ajlun S of Anjara,
27.IV.2002, 1♀, leg. F. M. Snižek, OÖLM; 15 km W
Jerash, Dibbin, 2.V.2006, 5♀, K. Deneš, OÖLM/TJW;
LEBANON: N Lebanon, Fehta, El Biara, 1632 m, 24.
IV.2018, 1♂, leg. Boustani, TJW.
Notes. Not restricted to high altitude sites in Israel
(Pisanty et al. 2018) or Jordan, so the single Lebanese
site is probably not representative of its ecological niche.
Newly recorded for Jordan.
Andrena (Poecilandrena) freidbergi Pisanty &
Scheuchl, 2018 *
Distribution. Previously known only from Israel (Mount
Hermon, Pisanty et al. 2018).
Material examined. LEBANON: N Lebanon, Arz Bcharre,
Forest of the Cedars of God, Forest Limit, 1873 m, 20.
V.2019, 1♀, leg. Boustani, TJW, Veronica polifolia.
Notes. No previous ower records (Pisanty et al. 2018),
but the association with Veronica would be consistent
with the subgenus, many members of which are
specialists or have strong association with this plant
genus. The species seems to be restricted to high
elevation sites in both Israel and Lebanon.
Andrena (Poecilandrena) labiata regina Friese, 1921 *
Distribution. This subspecies is known from Israel and
Turkey (Pisanty et al. 2018).
Material examined. LEBANON: N Lebanon, Arz
Tannourine Reserve Trail 4, 1800 m, 6.V.2017, 1♀, leg.
Boustani, TCFNR; N Lebanon, Hadath El Jebbeh, Border
of Cedar Forest, 1681 m, 19.IV.2018, leg. Boustani, 1♀,
TCFNR, Bellevalia exuosa; N Lebanon, Bcharre, Dahr
El Adib, 2306 m, 27.V.2017, 1♂, 2♀, leg. Boustani,
TCFNR, Gagea spp.; N Lebanon, Bcharre, Bcharre
Reforestation Area, 2347 m, 27.VI.2017, Boustani, 1♀,
TCFNR, Veronica polifolia.
Notes. The nominate subspecies of A. labiata is
polylectic with no clear preferences (Wood & Roberts
2017), so this may also be true for this subspecies as
well given the range of ower visits.
Andrena (Poecilandrena) rusticola Warncke, 1975 *
Distribution. Known from Israel and Turkey (Pisanty
et al. 2018).
Material examined. JORDAN: Aljoun, 28.IV.2012, 2♀,
leg. M. Kafka, OÖLM/TJW; LEBANON: N Lebanon,
Horch Ehden, Nabeh Jouit, 1336 m, 24.IV.2019, 1♂,
leg. Bous & Rasm, TJW, Veronica syriaca; SYRIA:
Latakia, Qaranjah, 750 m, 3.IV.1988, 1♂, leg. L. Blank,
MSC; Tartus, Sata, 10 km E, 300 m, 3.IV.1988, 1♀, leg.
L. Blank, MSC.
Notes. The association with Veronica is consistent with
data from Israel (Veronica leiocarpa, Pisanty et al. 2018).
This species is reported as new for Jordan and Syria.
Andrena (Poecilandrena) sphecodimorpha mediterranea
Scheuchl & Pisanty, 2016 *
Distribution. This subspecies is known from Israel
(Pisanty et al. 2018).
Material examined. JORDAN: Ajlun env [Aljoun],
840 m, 1.V.2006, 1♀, leg. K. Deneš, OÖLM;
LEBANON: N Lebanon, Tannourine Reserve, Trail 4,
1781 m, 13.V.2019, 1♂, leg. Boustani, TJW, Thlaspi
spp.; N Lebanon, Horch Ehden, Nabeh Jouit, 1336 m,
24.IV.2019, 1♂, leg. Bous & Rasm, TJW, Veronica
syriaca; N Lebanon, south of Jairoun, 23.V.2012, 1♀,
leg. M. Kasparek, TCFNR; SYRIA: Crac des
Chevaliers, 30.V.1995, 1♀, leg. K. Deneš, OÖLM.
Notes. Most of the records reported here occur a little
later in the season than the February–April ight period
reported by Pisanty et al. (2016, 2018). However, many
do occur further north and at a higher altitude, possibly
explaining this discrepancy. This species is reported as
new for Jordan and Syria.
Andrena (Poliandrena) unicincta Friese, 1899 *
Distribution. Known from Israel (Gusenleitner & Schwarz
2002), but the identication of this group of red-marked
Poliandrena has historically been difcult and confused,
see comments on Andrena unifasciata below.
Material examined. ISRAEL: 10 km E of Kiryat Gat,
13.V.2019, 6♀, leg. M. Halada, OÖLM; Bet Shemesh,
Britannia Park, 23.IV.2018, 13♀, leg. M. Halada, OÖLM;
R70, 10 km NNE Nahariya, 28.IV.2018, 1♀, leg. M.
Halada, OÖLM; Ramot Naftali, 10 km S of Kiryat
Shmona, 27.IV.2018, 13♀, leg. M. Halada, OÖLM;
JORDAN: 30 km NW Aljun, 600 m, 29.IV.2006, 1♀,
leg. K. Deneš, OÖLM; 10 km W Jarash, 1.V.1996, 1♀,
leg. Ma. Halada, OÖLM; Irbid, Alkfarat, 28–29.IV.2012,
26 T. J. Wood et al.

1♀, leg. M. Kafka, OÖLM; 15 km W Jerash, Dibbin, 2.
V.2006, 2♀, leg. K. Deneš, OÖLM; LEBANON: S
Lebanon, Saidoun, 5.III.2018, 1♀, leg. A. Baghdadi,
SOILS; 24.III.2019, 1♂, leg. A. Baghdadi, SOILS; N
Lebanon, Tannourine El Tahta, Al Mahbase, 893 m, 5.
V.2019, 1♀, leg. Boustani, TCFNR, Sonchus oleraceus.
Notes. See comments on Andrena unifasciata. This spe-
cies is reported as new for Jordan.
Andrena (Poliandrena) unifasciata Friese, 1899
stat. rev.
Distribution. Jordan, Lebanon, and the West Bank.
Literature. Friese (1899): Beirut, 28.IV.1889, 1♂, 1♀,
leg. F. Morice.
Material examined (Andrena unifasciata). WEST
BANK: [specimen labelled ‘Israel’] Wadi el Kelt [Wadi
Qelt], 5.III.1954, 1♀, (no collector information), OÖLM;
JORDAN: 15 km W Jerash, Dibbin, 2.V.2006, 4♀, leg. K.
Deneš, OÖLM; (Andrena caspica Morawitz, 1886):
ISRAEL: 10 km E of Kiryat Gat, 13.V.2019, 5♀, leg.
M. Halada, OÖLM; JORDAN: 20 km S of North Shuna,
Tall Al Arbatin, 19.IV.1996, 3♀, leg. Ma. Halada,
OÖLM; Aljoun, 28.IV.2012, 1♂, leg. M. Kafka, OÖLM;
SYRIA: Latakia, Saladinburg [Citadel of Saladin], 900 m,
4.IV.1988, 1♂, leg. L. Blank, MSC; Tartus, Sata, 10 km
E, 300 m, 3.IV.1988, 1♂, leg. L. Blank, MSC.
Notes. The situation within Near Eastern Poliandrena is
complex and has historically been greatly confused. As
summarised in Gusenleitner & Schwarz (2002), there are
six taxa (A. basimacula Alfken, 1929; A. caspica Morawitz,
1886; A. kriechbaumeri Schmiedeknecht, 1883; A. polita
Smith, 1847; A. unicincta; and A. westensis Warncke, 1965)
that form a group with the same genital construction.
Andrena basimacula is found in Libya and can be easily
separated due to its dark pubescence and A. kriechbaumeri
(Balkans to Turkey) can also be separated because of the
very strongly postfurcal position of the nervulus. This leaves
four taxa occurring in the Eastern Mediterranean that have
been dealt with confusingly and in various combinations
with each other.
For example, Andrena westensis was originally
described as a subspecies of A. unicincta from Greece
(Warncke 1965a), then later he suggested that it was instead
a subspecies of A. caspica, also including A. unicincta and
A. unifasciata as junior synonyms (Warncke 1967).
However, he later listed both A. caspica and A. unicincta
as good species (Warncke 1969a), but then moved A. wes-
tensis and A. caspica into synonymy with A. polita
(Warncke 1975a). As a result, the true distribution of all
these data is unclear; the map presented in Gusenleitner and
Schwarz (2002) is possibly the broadest possible interpreta-
tion of A. polita containing all six taxa.
Within a Lebanese context, it is best to identify the true
taxa occurring in the Levant. Friese described two similar
species from the Near East. Andrena unicincta from
Jerusalem, 7–10.IV.1899, 2♂, 2♀, leg. F. Morice, and
Andrena unifasciata Friese, 1899 from Beirut, 28.IV.1899,
also leg. F. Morice. According to Friese, the two taxa are
very similar but A. unifasciata differs in the female sex by
the clearly shiny mesonotum (“sonst wie unicincta und nur
an dem glänzenden, sparsam und tief punctirten
Mesonotum”, sculpturing clearly dull in unicincta) and the
black clypeus in the male (yellow in A. unicincta).
Andrena unifasciata was synonymised with A. caspica
by Warncke (1967) without any justication or written
reasoning, as was often the case (e.g. Schmid-Egger &
Doczkal 1995). It is noted by Gusenleitner and Schwarz
(2002) that A. caspica are variable in size, sometimes con-
taining smaller individuals (p. 164) and that sometimes the
colour of the clypeus varies from yellow to black (p. 165).
Examination of material from the Levant shows that the
original position of Friese was correct, and A. unifasciata
is a valid taxon that can be separated from other similar
Poliandrena in the Levant in the female sex by the degree of
shagreenation on the scutum, the shape of the labral process,
the overall body size, and the position of the nervulus and
additionally in the male sex by the coloration of the clypeus
(Table 1). Andrena kriechbaumeri is not included in this
determination table because it can be easily separated in
both sexes by the position of the nervulus which is strongly
postfurcal, and it is probably absent from the Levant. There
are therefore three red-marked Poliandrena in the Levant,
all of which probably have overlapping distributions.
Museum material should be carefully re-examined to clarify
their exact distributional ranges.
Grace (2010) reports A. caspica from Lebanon, but
given the confusion over A. caspica and A. unifasciata
this could refer to either taxon, and possibly refers simply
to the type locality of A. unifasciata, and so the presence
of true A. caspica in Lebanon is for now considered
unproven. The true identity of Levantine material identi-
ed as A. caspica should be reviewed. For now, we
consider that only A. unicincta and A. unifasciata are
present in Lebanon.
Andrena (Simandrena) thomsoni Ducke, 1898 *
Distribution. Southern Europe from France to Turkey
and Central Asia (Gusenleitner & Schwarz 2002).
Material examined. LEBANON: Bekaa, Kefraya,
1101 m, 27.IV.2019, 2♂, leg. Boustani, TCFNR; Bekaa,
West Bekaa, Ammiq, 871 m, 27.IV.2019, 1♂, leg.
Boustani, TJW, Sonchus oleraceus; N Lebanon,
Annales de la Société entomologique de France (N.S.) 27

Tannourine El Tahta, Wadi Ain El Raha, 900 m, 24.III.2017,
1♀, leg. Boustani, TJW; N Lebanon, Arz Tannourine,
Tannourine Forest Reserve Outskirts, 1739 m, 25.V.2017,
1♂, leg. Boustani, TCFNR, Crepis reuteriana.
Notes. Andrena thomsoni was reported from the western
and southern coasts of Turkey in high elevation areas such
as the Taurus mountains (Warncke 1975a), making this a
similar situation to that of A. rufula. Lebanese specimens
conform closely to comparative A. thomsoni material from
southern France. Andrena gasparella Patiny, 1998 was
described from a single locality in western Turkey
(Akşehir, Konya) and is similar to A. thomsoni, but
Lebanese material have a scutellum that is dull, not shiny
(Patiny 1998), in line with true A. thomsoni.
Andrena (Simandrena) transitoria Morawitz, 1871
Distribution. Central Europe and Italy eastwards to
Turkey, the Near East, and Central Asia (Gusenleitner &
Schwarz 2002).
Literature. Mavromoustakis (1962): Mount Lebanon,
Hammana, 16.V.1953, 1♀; Mount Lebanon, Ein el Arar,
25.V.1953, 1♀.
Material examined. JORDAN: 30 km N Tala [At-
Talah], 2.V.1996, 1♀, leg. Ma. Halada, OÖLM; 16 km
WWN Aljun [Aljoun], 600 m, 21.V.2007, 3♂, leg. Z.
Kejval, OÖLM; LEBANON: Beirut, American University
of Beirut, 1.V.1980, 1♀, leg. O.K. Rima, AUB.
Notes. This species is reported as new for Jordan.
Andrena (Simandrena) vetula Lepeletier, 1841
Distribution. Circum-Mediterranean to Central Asia
(Gusenleitner & Schwarz 2002).
Literature. Mavromoustakis (1962): Mount Lebanon,
Ein el Arar (near Baabdate), 25.V.1953, 1♀; S Lebanon
Djezzine [Jezzine], 2.VI.1953, 1♀.
Material examined. LEBANON: S Lebanon, Saidoun,
15.IV.2018, 1♂, Rapistrum rugosum; 22.IV.2018, 1♀; 5.
V.2018, 1♀, all leg. A. Baghdadi, SOILS; Mount
Lebanon, Daichouniyé, Nahr Beirut Climbing Site,
133 m, 27.IV.2017, 3♂, 3♀, leg. Boustani, TCFNR.
Notes. Originally placed in the Ptilandrena, A. vetula
clearly falls within with Simandrena (Pisanty et al. 2020).
Andrena (Suandrena) cyanomicans mirna Warncke,
1969 *
Distribution. Iberia, north-western Africa, and the Near
East (Gusenleitner & Schwarz 2002).
Table 1. Determination table for Poliandrena that have historically been confused in the Near East region. Characters require direct comparison across different taxa.
A. caspica Morawitz, 1886
A. polita Smith,
1847 A. unicincta Friese, 1899
A. unifasciata
Friese, 1899
A. westensis
Warncke, 1965
♀ labral process Clearly and deeply emarginate Rounded with tiny
emargination
Rounded with tiny emargination Slightly
emarginate
Rounded with tiny
emargination
♀ scutum structure Slightly shiny Slightly shiny Dull Shiny Dull
♀ nervulus Interfurcal Postfurcal Postfurcal Interfurcal Postfurcal
♀ size Larger, 12–13 mm Larger, 12–13 mm Larger, 11–12 mm Smaller,
10–11 mm
Larger, 12–13 mm
♀♂ abdomen colour Partly red on tergal discs Black, tergal
margins
lightened
Partly red on tergal discs Partly red on
tergal discs
Black, tergal
margins
lightened
♂ clypeus colour Completely yellow, sometimes also with small
yellow marks on lower paraocular area
Completely black Yellow front margin, occupying 1/3
of the total clypeal area
Completely black Completely yellow
28 T. J. Wood et al.

Material examined. LEBANON: Mount Lebanon, Fanar,
11.III.1965, 1♀, (no collector information, from the R.
Traboulsi Collection), USEK; Bekaa, American
University of Beirut farm, 15.III.1965, 1♂, 2♀, leg. M.
A. Sawsan, TCFNR.
Notes. The Suandrena are in need of revision. The
subspecies A. c. fratella Warncke, 1968 (Morocco) is
highly likely to be a valid species more closely related
to the Andrena maderensis Cockerell, 1922 group
(Kratochwil et al. 2014), and it is also likely that A. c.
mirna is a valid species.
Andrena (Taeniandrena) hova Warncke, 1975 *
Distribution. Central and eastern Turkey (Gusenleitner &
Schwarz 2002).
Material examined. LEBANON: N Lebanon, Fehta, El
Biara, 1632 m, 25.V.2018, 1♀, leg. Boustani, TCFNR, Vicia
tenuifolia; N Lebanon, Hadath El Jebbe, Al Fouar, 1529 m,
25.V.2018, 1♀, leg. Boustani, TCFNR, Vicia tenuifolia; N
Lebanon, Arz Tannourine, Gate area, 1754 m, 29.VI–6.
VII.2019, 1♀, leg. Boustani & G. Ghisbain, TCFNR, Vicia
tenuifolia; N Lebanon, Arz Bcharre, Forest of the Cedars of
God, 1883 m, 29.VI.2019, 2♂, 2♀, leg. Boustani & Jabbour,
TCFNR, Vicia tenuifolia and Vicia canescens; N Lebanon,
Arz Bcharre, Forest of the Cedars of God, Forest Limit,
1873 m, 5.VI.2019, 7♀, leg. Boustani, TCFNR, Vicia
tenuifolia; N Lebanon, Arz Bcharre, Bcharre Reforestation
Area, 1993 m, 27.VI.2017, 4♂, 3♀, leg. Boustani, TCFNR,
Vicia tenuifolia and Vicia canescens; N Lebanon, Bcharre
Reforestation Area, 2216 m, 2.VII.2019, 3♂, 5♀, leg.
Boustani, TCFNR, Vicia canescens.
Notes. Andrena hova was described from central (Ankara,
Madenşehri) and eastern (Erzurum) Turkey (Warncke
1975a, 1975b). Like other members of the Taeniandrena,
A. hova is probably oligolectic on Fabaceae (Wood &
Roberts 2017). All females were collected from Vicia
species, so far the only conrmed pollen host.
Andrena (Taeniandrena) ovatula (Kirby, 1802)
aggregate *
Distribution. Following a broad interpretation, A. ovatula
is a circum-Mediterranean species extending eastwards
into Central Asia (Gusenleitner & Schwarz 2002).
Material examined. LEBANON: N Lebanon, Horch
Ehden, Ain Al Bayada Gate, 1597 m, 5.VII.2019, 3♀,
leg. Boustani & G. Ghisbain, TJW, Medicago spp. &
Coronilla varia; N Lebanon, Arz Bcharre, Forest of the
Cedars of God, Forest Limit, 1897 m, 2.VII.2019, 1♂,
leg. Boustani, TJW, Anarrhinum orientale.
Notes. This taxon contains several cryptic species that are
identiable using COI barcoding throughout the Western
Palearctic, even in well-studied regions such as central and
western Europe, and the group therefore requires a deep
revision (Praz & Wood, in prep.). It is not at all clear
which name should be applied to Lebanese material, and
so these specimens are reported as A. ovatula in the broadest
possible sense (e.g. Warncke 1969a records from Israel) as
A. ovatula sensu stricto, a western taxon with a strong
Atlantic distribution (Praz & Wood, in prep.), is almost
certainly not found in the Eastern Mediterranean.
Andrena (Taeniandrena) similis Smith, 1849
Distribution. There is currently some taxonomic
uncertainty surrounding animals from North Africa; we
favour a single circum-Mediterranean taxon. Andrena
similis is otherwise found across Europe to Turkey, the
Caucasus, and the Near East (Gusenleitner & Schwarz 2002).
Literature. Grace (2010): Lebanon.
Material examined. JORDAN: North Shuna environs,
29–30.IV.1996, 1♂, leg. Mi. Halada, OÖLM; Ajlun S of
Anjara, 27.IV.2002, 1♀, leg. M. Snižek, OÖLM; 15 km
W of Jerash, 2.V.2006, 1♀, leg. K. Deneš, OÖLM; South
of Irbid, 13.IV.2009, 1♂, leg. M. Snižek, OÖLM;
LEBANON: S Lebanon, Sidon [Saida], 14.III.1973, 1♀,
(no collector information), AUB; Mount Lebanon,
Ksaibe, 518 m, 18.IV.2018, 1♂, leg. Boustani, TCFNR,
Sonchus spp.; Mount Lebanon, Bhamdoun, 1.V.1975, 1♀,
(no collector information), AUB; Mount Lebanon,
Bentael, 337 m, 23.IV.2019, 1♀, leg. Bous, Rasm,
Neme, TCFNR, Trifolium spp.; N Lebanon, Tannourine
El Tahta, Wadi Ain El Raha, 900 m, 27.III.2017, 1♀, leg.
Boustani, TCFNR; N Lebanon, Tannourine El Tahta,
Wadi El Fouar, 1187 m, 19.IV.2018, 1♂, leg. Boustani,
TCFNR, Anchusa hybrida; N Lebanon, Tannourine El
Tahta, Mar Boutrous, 1207 m, 25.IV.2019, 1♀, leg.
Boustani, TCFNR, Hymenocarpos circinnatus; N
Lebanon, Hadath El Jebbeh, Chemin Wadi Ain El Raha,
1519 m, 29.VI.2017, 2♀, leg. Boustani, TCFNR, Ononis
natrix; N Lebanon, Arz Tannourine, Tannourine Forest
Reserve Outskirts, 1766 m, 6.V.2017, 1♂, leg. Boustani,
TCFNR; N Lebanon, Arz Tannourine Reserve Trail 4,
1800 m, 6.V.2017, 1♀, leg. Boustani, TCFNR; N
Lebanon, Ehden, Jord, 1983 m, 5.VII.2019, 1♀, leg. G.
Ghisbain, UMONS, Onobrychis cornuta; N Lebanon,
Tannourine, Jord Tannourine, 2296 m, 30.VI.2019, 1♀,
leg. A. Gekiere, UMONS, Onobrychis cornuta; Arz
Bcharre, Forest of the Cedars of God, Forest Limit,
1897 m, 5.VI.2019, 3♀, leg. Boustani, TCFNR,
Onobrychis cornuta; N Lebanon, south of Jairoun, 23.
V.2012, 1♀, leg. M. Kasparek, TCFNR.
Notes. This species is reported as new for Jordan.
Annales de la Société entomologique de France (N.S.) 29

Andrena (Truncandrena) doursana mizorhina
Warncke, 1975 *
Distribution. Andrena doursana s.l. is found from
Morocco across North Africa and north to south-east
Turkey (Gusenleitner & Schwarz 2002). The subspecies
A. d. mizorhina is found in Adana, Turkey (Warncke
1975a) and probably Israel (Warncke 1969a, see Notes
for explanation).
Material examined. LEBANON: Bekaa, Kefraya, 27.
IV.2019, 1♂, 1♀, leg. Boustani, TCFNR.
Notes. Andrena doursana is a highly variable species,
with many described subspecies, several of which may
be valid taxa. The form occurring in the Levant was
originally called A. doursana derbentina Morawitz,
1886 (Warncke 1969a) after they were synonymised by
Warncke (1967). Andrena derbentina was described from
the Caucasus whereas A. doursana is a Mediterranean
taxon. Writing later, Warncke (1975b) described the
form in south-eastern Turkey as A. d. mizorhina, writing
that this was the bee formerly referred to as A. d.
derbentina (presumably referring to his 1969
publication) and that A. derbentina was a different, and
valid, species. Andrena d. mizorhina has a male clypeus
that is entirely dark, in contrast with North African and
Iberian forms of A. doursana, and examination of
paratype males of A. derbentina show that they have a
yellow tip to the end of the clypeus, at least in some
specimens (Gusenleitner & Schwarz 2002). The
Lebanese specimen has an entirely dark clypeus, and we
therefore take the position that this material is consistent
with A. d. mizorhina sensu Warncke (1975b).
Andrena (Truncandrena) fuligula Warncke, 1965 *
Distribution. Israel and Turkey (Gusenleitner & Schwarz
2002).
Material examined. LEBANON: Beka’a [Bekaa],
Qaraoun Lake, 24.III.2013, 1♂, 1♀, leg. M. Kasparek,
TJW; SYRIA: 8 km N of Shaykh Miskin [Al Sheikh
Maskin], 28.III.1994, 1♂, leg. S. Becvar, OÖLM; An
Nasrah [Al-Nasrah], 8-13.IV.2005, 1♂, leg. S. Jaki, OÖLM.
Notes. This species is reported as new for Syria.
Andrena (Truncandrena) medeninensis usura
Warncke, 1967 *
Distribution. Iberia, North Africa, Cyprus, and Turkey
(Gusenleitner & Schwarz 2002). The subspecies A. m.
usura is found in Cyprus and Turkey.
Material examined. LEBANON: Mount Lebanon, Chouf
Biosphere Reserve, Barouk Trails crossing, 1772 m, 16.
V.2019, 1♂, leg. Boustani, TCFNR, Thlaspi spp.; N
Lebanon, Tannourine Reserve, Trail 4, 1781 m, 13.
V.2019, 2♀, leg. Boustani, TCFNR/TJW, Thlaspi spp.;
N Lebanon, Arz Tannourine, Gate, 1796 m, 11.V.2019,
1♀, leg. Boustani, TCFNR; Mount Lebanon, Mazraat al
Daher, 529 m, 19.VI.2019, 1♀, leg. Boustani, TCFNR.
Notes. Another Truncandrena species that shows a huge
degree of variation over its range. This taxon and
Truncandrena species in general would benet from
molecular investigation.
Andrena (Truncandrena) optata Warncke, 1975
Distribution. The exact distribution of A. optata is
slightly unclear because it was originally described from
Greece as a subspecies of A. rufomaculata (below). It is
probably found in Eastern Europe, the Balkans, and
Turkey, but the southern limit of its range is unclear
(see Gusenleitner & Schwarz 2002 for combined map).
Literature. Grace (2010): Lebanon.
Material examined. LEBANON: Mount Lebanon, Chouf
Biosphere Reserve, Barouk-Maaser dirt road, 1769 m, 16.
V.2019, 1♂, leg. Boustani, TCFNR, Cynoglossum
nebrodense; N Lebanon, Tannourine El Tahta, Mar
Boutrous, 1207 m, 1♂, leg. Boustani, TJW.
Notes. Subsequent authors have accepted A. optata as a
valid species from A. rufomaculata as it is larger, has
signicantly longer mouthparts, and there are consistent
differences in the male genitalia. The species is
widespread in Turkey (Hazir et al. 2014).
Andrena (Truncandrena) rotundilabris rila Warncke,
1969
Literature. Warncke (1969a): N Lebanon, Cedars near
Bscharré [Bcharre], 1900 m, 12-19.VI.1931, leg. W.
Zerny (paratype); Grace (2010): Lebanon.
Distribution. Turkey and the Caucasus to Lebanon and
Israel (Gusenleitner & Schwarz 2002).
Andrena (Truncandrena) rufomaculata Friese, 1921
Distribution. For the same reasons as A. optata, the true
distribution of A. rufomaculata is unclear. It is probably
found in Turkey and the Levant (Gusenleitner & Schwarz
2002; Hazir et al. 2014).
Literature. Grace (2010): Lebanon.
Material examined. LEBANON: S Lebanon, Hilaliyah,
8.IV.1975, 1♂, (no collector information, from the R.
Traboulsi Collection), AUB; Beyrouth, (no date), 1♂,
leg. E. Baram, AUB; S Lebanon, Saidoun, 18.III.2018,
1♂, leg. A. Baghdadi, TJW; 3.IV.2018, 1♂, leg. A.
Baghdadi, SOILS; Mount Lebanon, Bchetfîne, 6.V.1995,
30 T. J. Wood et al.

1♂, leg. A. Fayad, AUB; Mount Lebanon, Fanar, 15.
III.1965, 1♂, (no collector information), USEK; Mount
Lebanon, Daichouniyé, Nahr Beirut Climbing Site,
133 m, 3.IV.2017, 2♂, 1♀, leg. Boustani, TCFNR/TJW;
Mount Lebanon, Jubayl [Byblos], 17.IV.2017, 1♂, leg. O.
Murr, M. Fakhry, USEK; N Lebanon, Tannourine El
Tahta, Wadi Ain El Raha, 900 m, 27.III.2017, 1♂, leg.
Boustani, TCFNR; Bekaa, American University of Beirut
farm, 17.IV.1961, 1♂, (no collector information), AUB.
Andrena (Truncandrena) serraticornis Warncke, 1965 *
Distribution. Greece, Turkey and Israel (Gusenleitner &
Schwarz 2002).
Material examined. JORDAN: Jarash environs, 1.
V.1996, 1♀, leg. Ma. Halada, OÖLM; 15 km W Jerash,
Dibbin, 2.V.2006, 1♀, leg. K. Deneš, OÖLM;
LEBANON: Mount Lebanon, Chammis, Wadi Cheber,
377 m, 5.IV.2019, 1♂, 2♀, leg. Boustani, TCFNR/TJW,
Sonchus tenerrimus; Mount Lebanon, Falougha, Cedar
woods, 1480 m, 27.IV.2019, 3♀, leg. Boustani, TCFNR,
Myagrum perfoliatum; SYRIA: Tartus, St. Georg-Kloster,
250 m, 3.IV.1988, 1♂, leg. L. Blank, TJW.
Notes. This species is reported as new for Jordan and
Syria.
Andrena (Truncandrena) tscheki tritica Warncke, 1965 *
Distribution. Central Europe to Turkey and the Near
East. Subspecies tritica is found in the Eastern
Mediterranean (Gusenleitner & Schwarz 2002).
Material examined. LEBANON: S Lebanon, Saidoun, 18.
III.2018, 1♀; 24.III.2019, 1♀; 15.IV.2019, 1♀, all leg. A.
Baghdadi, SOILS/TJW; S Lebanon, Saidoun, Joura, 25.
III.2018, 1♂, 1♀, leg. A. Baghdadi, SOILS, Raphanus
raphanistrum; N Lebanon, Tannourine El Tahta, Wadi Ain
El Raha, 900 m, 24–27.III.2017, 2♀, leg. Boustani, TCFNR.
Andrena (Ulandrena) dauma Warncke, 1969 *
Distribution. Israel and Turkey (Gusenleitner & Schwarz
2002).
Material examined. JORDAN: 20 km N of Karak,
1000 m, 27.IV.2006, 1♀, leg. K. Deneš, OÖLM; NW of
Ajlun, 850 m, 20.V.2007, 4♀, leg. Z. Kejval, OÖLM;
Ajlun S of Anjara, 27.IV.2002, 1♀, leg. M. Snižek,
OÖLM; 17 km SW of Amman, 23.IV.2006, 2♀, leg. F.
Kantner, OÖLM; LEBANON: N Lebanon, Arz
Tannourine, Tannourine Forest Reserve Outskirts,
1739 m, 6.V.2017, 1♂, 1♀, leg. Boustani, TCFNR/TJW;
N Lebanon, Arz Lubnan, Bcharre Reforestation Area,
1967 m, 12.V.2017, 1♀, leg. Boustani, TCFNR; SYRIA:
25 km W Damascus, Burqush environs [near Baddoûaa],
5.V.2000, 1♀, leg. F. Kantner, OÖLM.
Notes. This species is reported as new for Jordan and
Syria.
Andrena (Ulandrena) fulvitarsis Brullé, 1832 *
Distribution. Italy, Ukraine, and the Balkans to Turkey
and Israel (Gusenleitner & Schwarz 2002).
Material examined. JORDAN: 20 km S of North Shuna,
Tall Al Arbatin, 19.IV.1996, 2♀, leg. Ma. Halada,
OÖLM/TJW; LEBANON: Arz Bcharre, Forest of the
Cedars of God, Forest Limit, 1873 m, 20.V.2019, 1♀,
leg. Boustani, TJW.
Notes. This species is reported as new for Jordan.
Andrena (Ulandrena) isabellina Warncke, 1969 *
Distribution. Israel and Turkey (Gusenleitner & Schwarz
2002).
Material examined. JORDAN: Pella env. [Tabaqat Fahl],
−80 m, 29.IV.2006, 1♀, leg. K. Deneš, OÖLM; 30 km NW
Aljun, 600 m, 29.IV.2006, 1♀, leg. K. Deneš, OÖLM;
30 km WWN of Aljun, 30.IV.2006, 1♀, leg. K. Deneš,
OÖLM; Aljoun, 28.IV.2012, 1♀, leg. M. Kafka, OÖLM;
Aljun environs, 1.V.2006, 4♀, leg. K. Deneš, OÖLM; Irbid,
Saham village, 19–25.IV.2003, 1♂, 1♀, leg. I. Pljushtch,
OÖLM; Aman environs., Hisban villiage, 9.IV.2003, 1♀,
leg. I. Pljushtch, OÖLM; 15 km W Jerash, Dibbin, 2.
V.1996, 1♀, leg. K. Deneš, OÖLM; 10 km N, NE of
Jarash [Jerash], 20.IV.2002, 1♂, 2♀, leg. M. Snižek,
OÖLM/TJW; LEBANON: Bekaa, Kefraya, 1009 m, 27.
IV.2019, 2♀, leg. Boustani, TCFNR/TJW, Crepis spp.; N
Lebanon, Hadath El Jebbe, Border of the Cedar forest,
1646 m, 13.V.2019, 2♀, leg. Boustani, TCFNR, Crepis cf
sancta; SYRIA: Tartus, St. Georg-Kloster, 250 m, 3.
IV.1988, 1♂, leg. L. Blank, TJW; Tartus, Sata, 10 km E,
300 m, 1♂, leg. L. Blank, MSC; 60 km S Damascus, 14.
V.1996, 1♀, leg. Ma. Halada, OÖLM.
Notes. This species is reported as new for Jordan and Syria.
Andrena (Zonandrena) avipes Panzer, 1799
Distribution. The most widespread taxon in the West Palearctic,
found throughout Europe, North Africa, the Middle East, and into
Central Asia (Gusenleitner & Schwarz 2002).
Literature. Mavromoustakis (1962) S Lebanon, Djezzine
[Jezzine], 2.VI.1952, 4♀; Mount Lebanon, Hammana, 16.
V.1953, 1♀; Mount Lebanon, Ein el Arar, 25.VI.1953,
1♀; Kadisha river, 21.VI.1960, 1♀.
Material examined and notes. LEBANON: 4♂ and 29♀
constituting 23 records between 1975 and 2019 from
across Lebanon, this taxon is common and widespread.
Annales de la Société entomologique de France (N.S.) 31

Andrena (Zonandrena) sigiella Gusenleitner, 1998 *
Distribution. Israel (Gusenleitner & Schwarz 2002).
Material examined. JORDAN: Pella env. [Tabaqat Fahl],
−80 m, 29.IV.2006, 1♀, leg. K. Deneš, OÖLM; S of At Tala,
27–30.III.2013, 1♀, leg. M. Snižek, OÖLM; Al Karak env
[Kerak], 6.IV.2013, 1♀, leg. M. Snižek, OÖLM; 20 km NNW
Al Karak, 1–30.IV.2006, 1♀, leg. F. Kantner, OÖLM;
LEBANON: Mount Lebanon, Barja, Terbe, 326 m, 17.
IV.2017, 2♀, leg. Boustani, TCFNR; Bekaa, American
University of Beirut farm, 13.IV.1962, 1♀, leg. C.
Christian; 2.VI.1962, 1♀, leg. Z. Rai, both AUB; SYRIA:
Salkhad, 6.V.1996, 1♀, leg. Ma. Halada, OÖLM.
Notes. This species is reported as new for Jordan and
Syria.
Discussion
Altogether, the critical review of the literature and
museum specimens combined with contemporary survey
efforts have produced a total of 86 Andrena species from
Lebanon, with 53 species reported for the rst time
(including newly described species), as well as 25 and
23 species reported for the rst time from Jordan and
Syria, respectively. In-depth studies of other insect groups
are scarce for Lebanon, but the revisionary works that
have been compiled follow the same trend; high diversity
with several new records for Lebanon, and new species
for science described from mountainous areas (Larsen
1974; Németh 2019).
Discussion of the biogeography of Lebanese Andrena
species is difcult because most of the knowledge of their
distribution in neighbouring areas comes from Israel and
Turkey, and what we know about their faunas themselves
continues to change, with 35 species newly reported for
Israel very recently, a faunal richness increase of around
20% (Pisanty et al. 2018). The knowledge of the faunas
of Jordan and Syria is also highly incomplete, as evi-
denced by the new records presented here. That said,
three interesting groups of species can be identied in
the Lebanese Andrena fauna presented here.
The largest is the surprising collection of species more
typically found in temperate Europe, specically A. alf-
kenella (average site 1618 m above sea level), A. humilis
(1789 m), A. labialis (1728 m), A. minutula (1529 m), A.
minutuloides (1598 m), A. rufula (1553 m), and A. thom-
soni (1152 m) which were all found at altitude.
Altogether, these species were found at an average of
1567 m above sea level, illustrating their isolation. In
this regard, the mountains of northern Lebanon show a
strong afnity with the Taurus mountains in south-east
Turkey where these species can also be found at what was
thought to be their south-eastern range limit (Warncke
1975a). This broadly mirrors the situation in bumble
bees (Bombus) where B. melanurus Lepeletier, 1836 and
B. niveatus Kriechbaumer, 1870 show isolated popula-
tions at altitude in the Lebanese mountains, with the latter
only extending further south to mountains in northern
Israel (Rasmont & Iserbyt 2014; Boustani et al. 2020).
The second group is more Levantine in nature, and
contains species previously endemic to Israel, specically
A. aruana, A. freidbergi, A. pinkeunia, A. sigiella, A. sphe-
codimorpha mediterranea, and A. stenofovea. This group is
particularly rich in Poecilandrena, and the Lebanese has
gone from zero to ve conrmed species of
Poecilandrena, including two species or subspecies (A.
friedburgi and A. sphecodimorpha mediterranea) described
since 2016. Because of the small geographic area of the
countries of the Levant, it was highly likely that
Poecilandrena species known only from Israel would be
found more widely when the opportunity for scientic study
became available (Pisanty et al. 2018). Finally, the smallest
group is for species previously restricted to mountains in
Turkey and Armenia, specically A. gallinula and A. hova,
and in the case of A. christineae, mountains in Iran as well.
The smaller number of more typically ‘Turkish’ species
suggests that the relationship between the montane faunas
of Lebanon and Turkey may be relatively weak. However,
as evidenced by these new discoveries the level of sampling
in this region is low and it is difcult to draw strong
conclusions at the present time.
This problem is further illustrated by the limitations
placed on the program of sampling underlying the current
work, as little to no material was available for study from
the most southerly or northerly parts of Lebanon
(Figure 1), specically the governorates of Akkar,
Nabatieh, and South governorate, as these areas are dif-
cult to access safely and have not been the subject of
historical collections. These governorates contain some of
the most low-lying parts of the country with the most
extensive areas of thermomediterranean and eumediterra-
nean habitat (Abi-Saleh & Sa 1988), suggesting that
additional sampling here is likely to detect many of the
typical faunal elements from low-lying parts of the Near
East such as A. iliaca Warncke, 1969, A. nisoria Warncke,
1969, A. toelgiana Friese, 1921, and A. venerabilis
Alfken, 1935, as well as more recently described species
to date known only from Mediterranean Israel (Pisanty
et al. 2016, 2018).
There remains substantial uncertainty over several East
Mediterranean Andrena groups, most notably in
Carandrena, Euandrena, and Micrandrena (Pisanty et al.
2018; Praz et al. 2019). There are at least four additional
possibly undescribed Euandrena species in Lebanon that are
not listed here and which are the subject of molecular
investigation, including some of the cryptic members of
the Andrena bicolor complex in southern Europe identied
32 T. J. Wood et al.

by Praz et al. (2019), as well as several Micrandrena species
that cannot yet be condently assigned a name. Combined
with the high species richness of Andrena in neighbouring
countries in the Levant, the large number of unconrmed
species records for the country (online supplementary mate-
rial), and the need for increased study in undersampled
areas, it is highly likely that the true number of Andrena
species in Lebanon will be well over 100. The new species
described here, combined with other recent work (Pisanty
et al. 2016, 2018), support the position that studies of
Andrena in the Levant are far from complete.
Acknowledgements
Our thanks to Fritz Gusenleitner, Esther Ockermüller and Martin
Schwarz for access to the Warncke Collection and hospitality at
Linz, to Maximillian Schwarz (Ansfelden, Austria) for access to
his collection, and to Christophe Praz (University of Neuchâtel,
Switzerland) for valuable discussion of Mediterranean Andrena.
Maps were made using raster les prepared by Jonathan de Ferranti
(Scotland), interactive coverage maps supplied by Christoph
Hormann. We would also like to thank the Tannourine Cedar
Forest Nature Reserve (Lebanon), especially scientic expert Dr
Nabil Nemer for his advice and help in planning and carrying out
the eld work, and director Challita Tanios for his logistic support,
Sandra Koussa Saba (Director of Horch Ehden Nature Reserve)
and Charbel Tawk (Committee of the Cedar Forest Friends,
Bcharre) for their kind facilitation of eld work. We thank the
American University of Beirut (Lebanon), University of the Holy
Spirit Kaslik (Lebanon), Dr Max Kasparek (Heidelberg,
Germany), and SOILS Permaculture Association (Saidoun,
Lebanon) for the loan of specimens, and to all the people who
participated in collecting the specimens; Antoine Gekière,
Guillaume Ghisbain, Jad Jabbour, Mariam Fakhry, Nancy Abi
Khalil, Wael Yammine, and Xavier Van Achter.
Supplementary material
The supplemental data for this article can be accessed here.
References
Abi-Saleh B, Sa S. 1988. Carte de la végétation du Liban au 1/
500 000 + Notice explicative. Ecologia Mediterranea.
14:123–142. doi:10.3406/ecmed.1988.1209.
Al-Ghzawi A, Zaitoun S, Mazary S, Schindler M, Wittmann D.
2006. Diversity of bees (Hymenoptera: Apiforms) in extensive
orchards in the highlands of Jordan. Arxius de Miscellania
Zoologica. 4:42–48. doi:10.32800/amz.2006.04.0042.
Amiet F, Herrmann M, Müller A, Neumeyer R. 2010. Apidae 6:
Andrena, Melitturga, Panurginus, Panurgus. Vol. 26, Fauna
Helvetica. Neuchâtel (Switzerland): Centre Suisse de
Cartographie de la Faune (CSCF)/Schweizerische
Entomologische Gesellschaft (SEG).
Ascher JS, Pickering J. 2020. Discover life bee species guide and
world checklist (Hymenoptera: Apoidea: Anthophila). http://
www.discoverlife.org/mp/20q?guide=Apoidea_species.
Barbier Y, Rasmont P, Dufrêne M, Sibert J-M. 2000. Data
Fauna-Flora 1.0. Guide d’utilisation. Mons (Belgium):
Université de Mons-Hainaut; p. 106.
Bou Dagher-Kharrat M, El Zeina H, Rouhan G. 2018. Setting
conservation priorities for Lebanese ora. Identication of
important plant areas. Journal for Nature Conservation.
43:85–94. doi:10.1016/j.jnc.2017.11.004.
Boustani M, Yammine W, Nemer N, Abou Fakhr Hammad E,
Michez D, Rasmont P. 2020. Distribution and ower visita-
tion records of Bumblebees in Lebanon. Annales de la
Société entomologique de France. 56:115–124.
doi:10.1080/00379271.2020.1749885.
Dubitzky A. 2006. New palearctic species of the bee genus
Andrena (Insecta: Hymenoptera: Andrenidae). Zootaxa.
1284:1–27. doi:10.11646/zootaxa.1284.1.1.
Dubitzky A, Plant J, Schönitzer K. 2010. Phylogeny of the bee
genus Andrena Fabricius based on morphology.
Mitteilungen Muenchener Entomologischen Gesellschaft.
100:137–202.
Friese H. 1899. Neue palaearktische Sammelbienen.
Entomologische Nachrichten Berlin. 25:321–346.
Friese H. 1922. Neue Formen der Bienengattung Andrena
(Hym.). Konowia. 1:209–217.
Grace A. 2010. Introductory Biogeography to Bees of the
Eastern Mediterranean and Near East. Sussex (UK):
Bexhill Museum; p. 285.
Grünwaldt W, Osytshnjuk AZ, Scheuchl E. 2005. Neue
Andrena-Arten aus der Paläarktis (Hymenoptera: Apidae:
Andreninae). Entomofauna. 26:349–368.
Gusenleitner F. 1998. Neue westpaläarktische Andrena-Arten
(Hymenoptera: Apidae: Andreninae). Entomofauna.
19:109–144.
Gusenleitner F, Schwarz M. 2002. Weltweite Checkliste der
Bienengattung Andrena mit Bemerkungen und Ergänzungen
zu paläarktischen Arten (Hymenoptera, Apidae, Andreninae,
Andrena). Entomofauna, Supplement. 10:1–1280.
Hazir C, Keskin N, Scheuchl E. 2014. Faunistic, geographical
and biological contributions to the bee genus Andrena
(Hymenoptera, Andrenidae, Andreninae) from Turkey.
Zookeys. 38:59–133. doi:10.3897/jhr.38.7288.
Kratochwil A. 2015. Revision of the Andrena (Micrandrena)
tiaretta group: redescription of A. tiaretta WARNCKE,
1974 and description of two new species (A. cyrenaica
nov.sp. and A. orientalis nov.sp.) demarcating the central
and eastern part of the range (Libya, Israel, Syria). Linzer
biologische Beitrage. 47:1403–1437.
Kratochwil A, Schwabe A, Scheuchl E. 2014. Andrena
(Suandrena) portosanctana Cockerell, 1922 and A.
(Suandrena) maderensis Cockerell, 1922 – new taxonomical
and ecological data for two closely related endemic bee
species of the Madeira Archipelago, Portugal. Linzer biolo-
gische Beiträge. 46:1535–1567.
LaBerge WE. 1987. A revision of the bees of the genus Andrena of
the Western Hemisphere. Part XII. Subgenera Leucandrena,
Ptilandrena, Scoliandrena and Melandrena. Transactions of
the Entomological Society of America. 112:191–248.
Larsen TB. 1974. Butteries of Lebanon. Beirut (Lebanon):
National Council for Scientic Research; p. xv +256 + 16 pls.
Linsley EG. 1958. The ecology of solitary bees. Hilgardia.
27:543–585. doi:10.3733/hilg.v27n19p543.
Mavromoustakis GA. 1962. On the Bees (Hymenoptera:
Apoidea) of Lebanon Part III. Annals and Magazine of
Natural History. 5(13):647–655. doi:10.1080/
00222936208651300.
Michener CD. 2007. The bees of the world. 2nd ed. Baltimore
(USA): Johns Hopkins University Press.
Németh T. 2019. Elateridae (Coleoptera) of Lebanon. Zootaxa.
4614:127–159. doi:10.11646/zootaxa.4614.1.6.
Annales de la Société entomologique de France (N.S.) 33

Patiny S. 1998. Description d’une nouvelle espèce turque de
Simandrena Pérez, 1890 (Hymenoptera, Apoidea,
Andrenidae, Andreninae). Bembix. 10:34–36.
Pisanty G, Richter R, Martin T, Dettman J, Cardinal S. 2020.
Molecular phylogeny and historical biogeography of andre-
nine bees (Hymenoptera: Andrenidae). bioRxiv.
doi:10.1101/2020.06.09.103556.
Pisanty G, Scheuchl E, Dorchin N. 2016. Eight new species of
Andrena Fabricius (Hymenoptera: Apoidea: Andrenidae)
from Israel—a Mediterranean hotspot for wild bees.
Zootaxa. 4189:485–515. doi:10.11646/zootaxa.4189.3.3.
Pisanty G, Scheuchl E, Dorchin N. 2018. Taxonomic review of
the subgenus Andrena (Poecilandrena) (Hymenoptera:
Andrenidae) in Israel and the Levant. Zootaxa. 4374:151–
188. doi:10.11646/zootaxa.4374.2.1.
Portman ZM, Tepedino VJ. 2017. Convergent evolution of
pollen transport mode in two distantly related bee genera
(Hymenoptera: Andrenidae and Melittidae). Apidologie.
48:461–472. doi:10.1007/s13592-016-0489-8.
Praz C, Müller A, Genoud D. 2019. Hidden diversity in
European bees: Andrena amieti sp. n., a new Alpine bee
species related to Andrena bicolor (Fabricius, 1775)
(Hymenoptera, Apoidea, Andrenidae). Alpine Entomology.
3:11–38. doi:10.3897/alpento.3.29675.
Rasmont P, Devalez J, Pauly A, Michez D, Radchenko V.
2017. Addition to the checklist of IUCN European wild
bees (Hymenoptera: Apoidea). Annales de la Société
entomologique de France. 53:17–32. doi:10.1080/
00379271.2017.1307696.
Rasmont P, Iserbyt S. 2014. Atlas of the European Bees: genus
Bombus. 3rd ed. Mons (Gembloux): STEP Project, Atlas
Hymenoptera. http://www.atlashymenoptera.net/page.aspx?
ID=169.
Scheuchl E, Benarfa N, Louadi K. 2011. Description of a new
Andrena species from Algeria (Hymenoptera: Apoidea:
Andrenidae). Entomofauna. 32:221–232.
Scheuchl E, Hazir C. 2012. Description of three new Andrena
species (Hymenoptera: Apoidea: Andrenidae) from Turkey.
Florida Entomologist. 95:831–838. doi:10.1653/
024.095.0403.
Scheuchl E, Schindler M, Al-Ghawi AM. 2004. Andrena
(Holandrena) mbriatoides Scheuchl n. sp. (Hymenoptera:
Apoidea, Andrenidae), a new bee species from Jordan.
Zoology in the Middle East. 32:85–90. doi:10.1080/
09397140.2004.10638049.
Schmid-Egger C, Doczkal D. 1995. Der taxonomische Status
von Andrena fulvicornis Schenck, 1853 (Hymenoptera:
Apidae). Entomofauna. 16:1–12.
Schmid-Egger C, Scheuchl E. 1997. Illustrierte
Bestimmungstabellen der Wildbienen Deutschlands und
Österreichs und Berücksichtigung der Arten der Schweiz.
Band III Andrenidae. Velden: Eigenverlag; p. 180.
Schuberth J. 1995. Eine als neu erkannte Sandbienenart aus
Südosteuropa: andrena wilhelmi n. sp. (Hymenoptera,
Apoidea, Andrenidae). Linzer biologische Beiträge. 27:807–
821.
Schuberth J, Grünwaldt W, Schönitzer K. 2001. Klärung and
Neubeschreibung der Sandbiene Andrena asiatica Friese,
1921. Beiträge zur Entomologie. 51:65–71.
Schwenninger HR. 2015. Revision of the Western Palaearctic
species of the Andrena taraxaci-group with description of
four new species (Hymenoptera: Andrenidae). Stuttgarter
Beiträge zur Naturkunde A, Neue Serie. 8:251–270.
Thorp RW. 1979. Structural, behavioral, and physiological adapta-
tions of bees (Apoidea) for collecting pollen. Annals of the
Missouri Botanical Garden. 66:788–812. doi:10.2307/2398919.
Traboulsi F. 2012. A history of modern Lebanon. 2nd ed.
London: Pluto Press; p. 321.
Warncke K. 1965a. Beitrag zur Kenntnis der Bienengattung
Andrena Fabricius in Griechenland. Beiträge zur
Entomologie. 15:27–76.
Warncke K. 1965b. Beitrag zur Kenntnis der Bienengattung Andrena
F. in der Türkei (Hymenoptera, Apoidea). Mitteilungen der
Münchner Entomologischen Gesellschaft. 55:244–273.
Warncke K. 1967. Beitrag zur Klärung paläarktischer Andrena-
Arten. Eos. 43:171–318.
Warncke K. 1968. Die Untergattungen der
westpaläarktischen Bienengattung Andrena F. Memórias
e Estudos do Museu Zoológico da Universidade de
Coimbra. 307:1–110.
Warncke K. 1969a. A contribution to the knowledge of the
genus Andrena (Apoidea) in Israel. Israel Journal of
Entomology. 4:377–408.
Warncke K. 1969b. Bienen der Gattung Andrena F. aus der
Türkei und dem Balkan (Hymenoptera, Apoidea,
Andrenidae). Bulletin des Recherches agronomiques de
Gembloux. 4:302–305.
Warncke K. 1972. Zwei neue Sandbienen aus der Ukraine und
aus Ungarn (Hym. Apoidea). Nachrichtenblatt der
Bayerischen Entomologen. 21:123–127.
Warncke K. 1975a. Die Sandbienen der Türkei (Hymenoptera,
Apoidea, Andrena), Teil A. Mitteilungen der Münchner
Entomologischen Gesellschaft. 64:81–116.
Warncke K. 1975b. Die Sandbienen der Türkei (Hymenoptera,
Apoidea, Andrena), Teil B. Mitteilungen der Münchner
Entomologischen Gesellschaft. 65:29–102.
Wood TJ, Roberts SPM. 2017. An assessment of historical and
contemporary diet breadth in polylectic Andrena bee spe-
cies. Biological Conservation. 215:72–80. doi:10.1016/j.
biocon.2017.09.009.
34 T. J. Wood et al.