Research Article |
Corresponding author: Thomas James Wood ( thomasjames.wood@umons.ac.be ) Academic editor: Thorleif Dörfel
© 2020 Thomas James Wood, Denis Michez, Diego Cejas, Patrick Lhomme, Pierre Rasmont.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Wood TJ, Michez D, Cejas D, Lhomme P, Rasmont P (2020) An update and revision of the Andrena fauna of Morocco (Hymenoptera, Apoidea, Andrenidae) with the description of eleven new North African species. ZooKeys 974: 31-92. https://doi.org/10.3897/zookeys.974.54794
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Morocco has a diverse bee fauna, but one that has also been relatively understudied in recent years. Here a revision of the species-rich genus Andrena is presented that reveals eleven new species for science and substantially improves our understanding of North African Andrena. From Morocco, Andrena (Aciandrena) semiadesus Wood, sp. nov., Andrena (Aciandrena) triangulivalvis Wood sp. nov., Andrena (Campylogaster) sparsipunctata Wood sp. nov., Andrena (Carandrena) hebescens Wood sp. nov., Andrena (Cnemidandrena) niveofacies Wood sp. nov., Andrena (incertae sedis) tenebricorpus Wood sp. nov., Andrena (Notandrena) acutidentis Wood sp. nov., Andrena (Poliandrena) breviceps Wood sp. nov., and Andrena (Poliandrena) farinosoides Wood sp. nov. are described and their ecology is discussed. Andrena (Aciandrena) astrella Warncke, 1975 is synonymised with Andrena (Aciandrena) fulica Warncke, 1974 syn. nov. The unknown female of Andrena (Nobandrena) ounifa Warncke, 1974, and the unknown male of Andrena (Poliandrena) guichardi Warncke, 1980 are described. Andrena (incertae sedis) gafsensis Wood sp. nov. from Tunisia is described due to its similarity to Andrena tenebricorpus. Andrena (Poecilandrena) nigriclypeus Wood sp. nov. from Algeria is also described as it was collected within 10 km of the Moroccan border. A further 18 species are recorded in Morocco for the first time. Andrena (Melandrena) nitida (Müller, 1776) and Andrena (Notandrena) nitidiuscula Schenck, 1853 are removed from the Moroccan list due to historic problems in the application of these names to Mediterranean taxa.
alpine, deserts, endemic, faunal list, pollen host plants, solitary bees, taxonomy
Andrena are the second most speciose genus of bees worldwide after Lasioglossum (
Inspection of 5,685 female and 2,275 male Andrena specimens from contemporary and ongoing collections in Morocco and undetermined material from the Oberösterreich Landesmusum, Linz, the Naturalis Biodiversity Center, Leiden, and the personal collection of Maximillian Schwarz, Ansfelden, comprised 155 valid species, with many new records for Morocco and undescribed taxa. These are described herein, and the identity of several problematic taxa are reviewed.
Identification was enabled through a comprehensive review of the literature in combination with visits to the Warncke collection in Linz as part of the lead author’s ongoing review of West Palearctic Andrena species (
The subgeneric classification of Andrena continues to pose problems. Because of the very large number of species, taxonomic workers have largely been restricted to only a part of the global fauna (broadly, West Palearctic, Central and Eastern Palearctic, and Nearctic falling into three groups), and therefore a co-ordinated global system of subgeneric classification has not yet been possible. In both a Moroccan and West Palearctic context, the system of
Body length was measured from the vertical plane of the front of the head to the tip of the metasoma. Morphological terminology follows
In order to investigate a possible synonymy (see section on Andrena alchata Warncke, 1974), DNA was extracted and amplified from two specimens. DNA was extracted from two legs of each individual using a NucleoSpin Tissue (Macherey-Nagel, Düren Germany) extraction kit and following manufacturer´s instructions. The region selected for amplification was the LEP fragment within the mitochondrial gene cytochrome oxidase I (cox1), a region widely used in Hymenoptera taxonomy (
MSC Maximillian Schwarz personal collection, Ansfelden, Linz, Austria
NMNL National Museum of Natural History Naturalis, Leiden, the Netherlands
OÖLM Oberösterreich Landesmusum, Linz, Austria
TJW Thomas Wood personal collection, Mons, Belgium
UMONS University of Mons collection, Mons, Belgium
Holotype : Morocco: Fès-Meknès, Laanoucer, 1456 m, 33.6166N, -4.7484W, 11–12.iv.2019, 1♂, white pan trap, leg. L. Hamroud & A. Sentil. Deposited in the OÖLM. Paratypes: Morocco: Fès-Meknès, Laanoucer, 1456 m, 11–12.iv.2019, 4♂, white and yellow pan traps, leg. L. Hamroud & A. Sentil; Drâa-Tafilalet, 20 km W Boudnib, 9.iv.1995, 2♂, 2♀, leg. Ma. Halada. Paratypes are deposited at the OÖLM, with a male and female retained in the personal collection of TJW.
The finely shagreened propodeal triangle, the narrow facial foveae, the absence of longitudinal striations on the clypeus, and the yellow-marked clypeus in the male place this bee in the Aciandrena. The classification of Aciandrena and its relationship to Micrandrena, Graecandrena, and Distandrena is somewhat in flux, and the subgenus is currently polyphyletic (
Female: Body length 7 mm (Fig.
Male. Body length 6.5–7 mm (Fig.
Central and eastern Morocco (Fig.
None recorded.
The name semi (partly or partially) + adesus (eaten, worn, eroded) was chosen to illustrate the clypeus of the male, where the central yellow marking is laterally diminished by intruding black marks.
Holotype : Morocco: Drâa-Tafilalet, 20 km W Boudnib, 9.iv.1995, 1♂, leg. Ma. Halada. Deposited in the OÖLM. Paratypes: Morocco: Drâa-Tafilalet, 20 km W Boudnib, 9.iv.1995, 3♂, leg. Ma. Halada. Paratypes are deposited at the OÖLM, with a male retained in the personal collection of TJW.
The male of A. triangulivalvis resembles other Aciandrena with a yellow polished clypeus such as A. pratincola. However, it can be instantly separated from all other Aciandrena with a yellow clypeus by the structure of the genitalia. Normal Aciandrena genitalia are simple, with a relatively narrow, unmodified penis valve (Figs
Female: Unknown.
Male. Body length 6 mm (Fig.
Eastern Morocco from the province of Drâa-Tafilalet (Fig.
None recorded.
The name trianguli (triangular) + valvis (valve) was chosen because of the remarkable male genitalia in which the penis valve is inflated and triangular in shape, strongly contrasting with other Aciandrena species.
Holotype : Morocco: Guelmim-Oued Noun, 10 km E Guelmim, 15–16.iv.1995, 1♀, leg. Ma. Halada. Deposited in the OÖLM. Paratypes: Morocco: Guelmim-Oued Noun, 10 km E Guelmim, 15–16.iv.1995, 1♂, 1♀, leg. Ma. Halada, OÖLM; Oriental, 10 km S Bouarfa, 20.v.1995, 2♂, leg. Ma. Halada, OÖLM. Paratypes are deposited in the OÖLM.
Andrena sparsipunctata can be easily placed into the Campylogaster because of the large, dense, and clear punctures on the episternum combined with its large body size. However, recent evidence shows that Campylogaster is strongly polyphyletic, and the species in northwestern Africa do not fall close to A. (Campylogaster) erberi Morawitz, 1871, the type species of Campylogaster that differs by its tormentose pilosity. A new subgenus is probably needed for the species around A. sparsipunctata (
Andrena (Campylogaster) sparsipunctata sp. nov. 13 female profile 14 female mesonotum in semi-profile 15 female head dorsal view 16 female tergites. Andrena (Campylogaster) pruinosa succinea Dours, 1872 17 female head dorsal view 18 female tergites. Andrena caroli Pérez, 1895 19 female head dorsal view 20 female tergites.
In males, A. sparsipunctata can be recognised by the shape of the labrum which is wide and rectangular (Fig.
Andrena (Campylogaster) sparsipunctata sp. nov. 21 male profile 22 male genitalia 23 male face and labrum 24 male tergites. Andrena (Campylogaster) pruinosa succinea Dours, 1872 25 male face and labrum 26 male tergites. Andrena caroli Pérez, 1895 27 male face and labrum 28 male tergites.
Female: Body length 12 mm (Fig.
Male. Body length 11–12 mm (Fig.
Southern and eastern Morocco in desert environments (Fig.
None recorded.
There is uncertainty over the exact status of A. pruinosa succinea, as the North African animals differ in colouration of the metasoma and scutal hairs (red) from A. pruinosa sensu stricto (black-brown) from Spain. However, structural differences are minor (e.g., the shape of the foveae is the same), and molecular investigation is warranted. It is however clear that the name succinea cannot apply to the bees described here as the first tergite of females is much less densely punctured than A. pruinosa s.l. from either Spain or North Africa, and
The name sparsi (sparse) + punctata (punctured) was chosen to illustrate the first tergite of females, which is much less densely punctate than the similarly coloured A. pruinosa succinea and A. caroli.
Holotype : Morocco: Guelmim-Oued Noun, 15–16.iv.1995, 1♀, leg. Ma. Halada. Deposited in the OÖLM. Paratypes: Morocco: Guelmim-Oued Noun, 15–16.iv.1995, 4♂, 6♀, leg. Ma. Halada, OÖLM; Souss-Massa, Tassademt, 50 km NE, Agadir, 19.iv.1996, 1♀, leg. M. Schwarz, OÖLM; Souss-Massa, Aoulouz-Taliouine, 19.iii.1988, 1♂, leg. H. Teunissen. Paratypes are deposited at the OÖLM and NMNL, with a male and female retained in the personal collection of TJW.
The female of A. hebescens can be placed in the Carandrena because the dorsolateral angle of the pronotum has a transverse ridge, the propodeal triangle is shagreened and weakly rugose at the base, and there is almost no punctation on the metasoma, and the head has a typical Carandrena shape, broader than long, with the inner eye margins slightly converging below (Fig.
The male of A. hebescens has a yellow marking on the clypeus, but this is reduced in size and does not cover the entire clypeal surface, with two black markings that extend in from the clypeal margins giving the marking an inverted ‘T’ shape (Fig.
Female: Body length 8 mm (Fig.
Male. Body length 8 mm (Fig.
South-western Morocco in the Souss valley (Fig.
None recorded.
The name hebescens was chosen because this member of the Carandrena, though morphologically similar to several species with metallic green colouration, is completely dark, therefore heb- (dull or blunt) + escens (becoming).
Holotype : Morocco: Marrakesh-Safi, Asni, S Imlil, 2900 m, 24.viii.1992, 1♀, leg. Warncke [see Remarks]. Deposited in the OÖLM. Paratypes: Morocco: Marrakesh-Safi, Asni, S Imlil, 3300 m, 21.viii.1992, 1♂, leg. Warncke, OÖLM.
Andrena niveofacies can easily be recognised as part of the Cnemidandrena because of the transverse ridge on the dorsolateral angle of the pronotum, the triangular hind tibiae in the female, the upturned process of the labrum in the male (Fig.
Within the Cnemidandrena, A. niveofacies females have completely brown hair on the scutum (intermixed with black in A. denticulata (Kirby, 1802)), the galea is dull (shiny in A. fuscipes (Kirby, 1802)), T5–6 are black haired (light haired in A. tridentata (Kirby, 1802)), and the face is pale haired (black at least in part in A. nigriceps (Kirby, 1802) and A. freygessneri Alfken, 1904). The female A. niveofacies is most similar to A. simillima Smith, 1851 but is can be easily separated as it has strikingly bright white hairs on the face and gena (Fig.
The male has a non-carinate gena (Fig.
Female: Body length 10 mm (Fig.
Male. Body length 10 mm (Fig.
Probably restricted to the High Atlas Mountains of Morocco. This material represents the first record of the subgenus Cnemidandrena from Morocco and more broadly the whole of North Africa (Fig.
The single female had a full scopa (Fig.
There has historically been uncertainty over the species status of taxa assigned to A. simillima. The former subspecies A. freygessneri was returned to species status by
The collection labels themselves are a small mystery, as they have the collector name Warncke struck through by hand (Warncke). They may have been used by a collector accompanying Warncke who used his spare preprinted labels, but if this is the case then their identity is not clear.
The subspecific epithet niveofacies from niveus (snow) + facies (face) was chosen to illustrate the bright white facial hair.
(Andrena simillima sischkai): Greece: Olympus, 2000–2200 m, 18.viii.1978, 1♀, leg. K. Warncke, OÖLM (holotype); Bulgaria: mer., Pirin, Popina-Lka I (1350 m), 23–27.vii.1974, 1♂, leg. Dr. A. Hoffer, OÖLM (paratype); (Andrena simillima simillima): United Kingdom: Devon, Weston Cliff NT, Branscombe, 23.vii.1992, 1♀, leg. M. Edwards, TJW.
Holotype : Tunisia: 40 km NW Gafsa, 17.iv.1994, 1♀, leg. M. Schwarz. Deposited in the OÖLM. Paratypes: Tunisia: 40 km NW Gafsa, 17.iv.1994, 3♂, 1♀, leg. M. Schwarz. Paratypes are deposited at the OÖLM, with a male and female retained in the personal collection of TJW.
Placement of this species into a subgenus is not immediately obvious. Andrena gafsensis is a small to medium sized Andrena with a strongly flattened clypeus that is extensively shiny in both the female (Fig.
Nobandrena normally contains large species in the range of 13–14 mm in length, most of which have males with domed clypei which are extensively yellow-marked, with the yellow extending onto the lower paraocular areas. However, there are two smaller species A. iliaca Warncke, 1969 (7–8 mm, Turkey, Israel, and Jordan) and A. ounifa Warncke, 1974 (7–7.5 mm, Algeria, see below for new description of female) placed into the Nobandrena by Warncke where the male clypeus is shinier and flatter, whilst retaining the same colour pattern. This flattened clypeus can be seen in the female of A. ounifa which has a strongly flattened and shining clypeus in both the female (Fig.
Given these problems, Andrena gafsensis is therefore best recognised in the female by the distinctive clypeal structure (Fig.
Female: Body length 9 mm (Fig.
Male. Body length 9–10 mm (Fig.
Known only from the locus typicus in southern Tunisia.
None recorded.
Named after the nearest indicated town Gafsa in southern Tunisia.
Holotype : Morocco: Guelmim-Oued Noun, 10 km E Guelmim, 15.iv.1995, 1♀, leg. Ma. Halada. Deposited in the OÖLM.
Andrena tenebricorpus is very similar to A. gafsensis and faces the same problems of subgeneric classification, and is therefore also not currently placed in one until molecular data are available. Both species have a flattened and shiny clypeus (Fig.
In terms of overall colouration and appearance A. tenebricorpus is extremely similar to A. guichardi Warncke, 1980 (also found only in south-western Morocco) with the same dark to dark brownish tergal colouration with contrasting white hair bands (see Fig.
Female: Body length 10 mm (Fig.
Male. Unknown.
South-western Morocco (Fig.
None recorded.
Whilst A. tenebricorpus is structurally very similar to A. gafsensis, the subtle morphological differences suggest that A. tenebricorpus is distinct. Moreover, the Souss valley to the Guelmim region is bio-climatically quite distinct to the deserts of southern Tunisia and hosts many unique bee species within Morocco itself, further suggesting a distinct specific identity.
The name tenebricorpus from tenebris (dark) + corpus (body) was chosen because of the structural similarities between A. tenebricorpus and A. gafsensis, but without the same reddish colouration.
Holotype : Morocco: Souss-Massa, 10 km SE Ait Baha, 18.iv.1996, 1♀, leg. M. Schwarz. Deposited in the OÖLM. Paratypes: Morocco: Souss-Massa, 10 km SE Ait Baha, 18.iv.1996, 2♂, 2♀, leg. M. Schwarz; 20.iv.1996, 2♂, leg. M. Schwarz; Souss-Massa, 10 km W Tiznit, 6.v.1995, 5♂, 1♀, leg. Mi. Halada, OÖLM; Souss-Massa, 30 km SE Taliouine, 17.iv.1996, 3♀, leg. M. Schwarz, OÖLM; Souss-Massa, Biougra-Tafraout, 13.ii.1987, 1♂, leg H. Teunissen, NMNL. Paratypes are deposited at the OÖLM and NMNL, with a male and female retained in the personal collection of TJW.
A small Notandrena recognised in the subgenus by the dorsolateral angle of the pronotum with a transverse ridge, the clearly punctured metasoma, and the weakly rugose (not shagreened) propodeal triangle. Because of its small size it can be placed into the nitidiuscula group, and it is most similar to A. fulvicornis Schenck, 1853 (see also species newly recorded for Morocco below). It differs by the clypeus which has a central, shining, impunctate line (Fig.
The male can easily be recognised as a Notandrena because of the greatly enlarged and carinate gena (Fig.
Female: Body length 8–8.5 mm (Fig.
Male. Body length 8 mm (Fig.
The Souss valley in south-western Morocco (Fig.
None recorded. Other members of the Notandrena are associated with Apiaceae (
The name acuti (sharp) + dentis (teeth) was chosen because of the male genitalia where the apices of the gonostyli are produced into points in contrast to other members of this group where they are rounded.
Holotype : Algeria: Tlemcen, 20 km N de Maghnia, Bab Taza, 34.968N, -1.7622W, 9.iv.1983, 1♂, leg. R. Leys & P. v. d. Hurk. Deposited in the NMNL.
The subgenus Poecilandrena is poorly defined and has been treated as a wastebasket for species with few apomorphies (
Currently only one species of Poecilandrena sensu lato is known from North Africa, the similarly red-marked A. maximiliani Scheuchl, 2009 which was described from Tunisia (see
Female: Unknown.
Male. Body length 8 mm (Fig.
North-western Algeria close to the Moroccan border (Fig.
None recorded.
The name nigri (black) + clypeus (clypeus) was chosen because of the entirely black male clypeus, which is unusual within small red-marked Andrena species.
Holotype : Morocco: Drâa-Tafilalet, 10 km N Erfoud, 10.iv.1995, 1♀, leg. Ma. Halada. Deposited in the OÖLM. Paratypes: Morocco: Drâa-Tafilalet, 10 km N Erfoud, 10.iv.1995, 2♀, leg. Ma. Halada, OÖLM; Drâa-Tafilalet, 20 km E Agdz, 20.iv.1995, 2♀, leg. Mi. Halada, OÖLM; Drâa-Tafilalet, Tagounite, 60 km S Zagora, 23.iv.1995, 1♀, leg. Ma. Halada, OÖLM. Paratypes are deposited at the OÖLM with a female retained in the personal collection of TJW.
The subgenus Poliandrena Warncke, 1968 is currently unsatisfactorily defined (females keying out in three places in
Andrena breviceps is small relative to the rest of the Poliandrena, comparable in size to A. marsae Schmiedeknecht, 1900 and A. laurivora Warncke, 1974, two of the smallest Poliandrena, but A. breviceps can easily be separated from them by the colour of the tergites which are dark brown (red in A. marsae, dark metallic green-blue in A. laurivora) and the sculpturing of the scutum which is shiny and (relatively within the Poliandrena) sparsely punctate, punctures separated by 2–3 puncture diameters (Fig.
Female: Body length 8.5–9 mm (Fig.
Male. Unknown.
The eastern Moroccan desert in the province of Drâa-Tafilalet (Fig.
None recorded.
The name brevi (short) + ceps (head) was chosen because of the particularly short and wide head of this species, even within the Poliandrena.
Holotype : Morocco: Oriental, 40 km S Guercif, 15–17.v.1995, 1♀, leg. Ma. Halada. Deposited in the OÖLM. Paratypes: Morocco: Oriental, 40 km S Guercif, 15–17.v.1995, 2♂, 24♀, leg. Ma. Halada, OÖLM; Drâa-Tafilalet, 30 km E Midelt, 13.v.1995, 1♀, leg. Mi. Halada, OÖLM; Drâa-Tafilalet, 10 km N Rich, 2♀, leg. Mi. Halada, OÖLM. Paratypes are deposited at the OÖLM with three females retained in the personal collection of TJW.
Andrena farinosoides can also be placed in the Poliandrena because of its short and broad head (Fig.
Diagnosis of the male is more difficult, but both A. farinosa and A. farinosoides are in the group with dark, densely punctate tergites and black (not yellow) clypei with an upturned fore margin (Fig.
Female: Body length 8.5–9.5 mm (Fig.
Male. Body length 7 mm (Fig.
The eastern Moroccan desert in the provinces of Oriental and Drâa-Tafilalet (Fig.
None recorded.
Given the similarity to A. farinosa, the name A. farinosoides (farinosa + oides, form or likeness) was chosen to illustrate this close link.
(Andrena farinosa): Spain: Barcelona, [no date], 1♀, designated paratype [technically paralectotype] by Warncke, Warncke Colln., OÖLM (illustrated Figs
Andrena (Aciandrena) astrella ssp. fulica Warncke, 1974: p44
Andrena (Aciandrena) astrella Warncke, 1975: 305. syn. nov.
Tunisia: Tunis, 1898, 1♂, leg. O. Schmiedeknecht (paratype A. fulica), OÖLM; Morocco: Fès-Meknès, Ifrane environs, 9.v.1997, 1♂, leg. K. Deneš, OÖLM (illustrated Figs
Morocco: Fès-Meknès, Ifrane environs, 9.v.1997, 1♂, leg. K. Deneš, OÖLM (illustrated Figs
Andrena fulica (Morocco, Algeria, and Tunisia) and A. astrella (Spain and Portugal) were both described by Warncke, with A. fulica described as A. astrella ssp. fulica (Warncke, 1974) despite being formally published before A. astrella (
The two species can be rapidly recognised within the Aciandrena because they have punctured tergites with the punctures extending onto the tergal margins, a character that is unique within this subgenus. According to
Inspection of Aciandrena males from Ifrane in Morocco (9.v.1997) revealed the two colour forms in sympatry, though unfortunately only two specimens in total (one A. fulica, one A. astrella) were collected during this sampling making an assessment of variation impossible. Additionally, specimens from Benidorm in Spain (20.iv.1982) also showed the two colour forms in sympatry. Across all Iberian material studied, 46/47 (97.8%) males showed a consistently yellow clypeus. Inspection of the genitalia showed no obvious differences between the two Ifrane specimens (Figs
Aciandrena do not always show consistent clypeal colouration. In the recently described A. (Aciandrena) abruptifovea Wood, 2020, a series of six males from the same type locality site collected on the same day vary considerably in colouration from a clypeus with 80% yellow coverage, to 40%, to entirely black (
Morocco: Drâa-Tafilalet, Errachidia, 11.iv.1995, 2♀, leg. Ma. Halada, one female deposited in the OÖLM, with one female retained in the personal collection of TJW.
Previously known only from the type locality in the western Algerian part of the Sahara Desert. The specimens from Errachidia are approximately 350 kilometres to the west. They agree with the male in size, the flattened and shiny clypeus (Figs
Female: Body length 7.5 mm (Fig.
Algeria: Beni Ounif, 6.iii.[year unknown], 1♂, leg. Weber, OÖLM (holotype), illustrated Figs
Morocco: Guelmim-Oued Noun, 15–16.iv.1995, 3♂, 1♀, leg. Ma. Halada, OÖLM; Souss-Massa, 10 km S Taroudant, 12.iv.1995, 1♀, leg. Ma. Halada, allotype male and one other male deposited in the OÖLM, with one male and one female retained in the personal collection of TJW.
Known only from south-western Morocco (
Male: Body length 9 mm (Fig.
Morocco: Sidi-Ifni, within 100 m of the sea, 31.iii.1974, 1♀, leg. K.M. Guichard & G.R. Else, OÖLM (paratype).
Andrena alchata: Algeria: Maison Carrée, Alger, 1♂, leg. Dr. J. Bequaert, OÖLM (holotype); Andrena doursana agadira: Morocco: S-Marokko, Agadir, 20.ii.1977, 1♀, OÖLM (holotype); Andrena doursana citreola Warncke, 1975: Spain: Vaciamadrid, 25.v.1919, 1♀, leg. Dusmet, OÖLM (holotype); Morocco: Azrou Ras el Ma, 30.iii.1923, 1♀, leg. Schulthess, OÖLM (paratype); Rez Dj. Zalagh, 25.iii.1923, 1♂, leg. Schulthess, OÖLM (paratype).
Andrena alchata: Morocco: Fès-Meknès, Laanoucer, 33.6167, -4.7489, 2.v.2018, 1♂, 3♀, leg P. Lhomme & A. Sentil; 33.6708, -4.8527, 2–3.v.2018, 1♀, white pan trap, leg P. Lhomme & A. Sentil; 33.6699, -4.8673, 10–11.v.2018, 1♂, 1♀, yellow pan trap, leg P. Lhomme & O. Ihsane; 33.7099, -4.8431, 15–16.v.2018, 1♀, white pan trap, leg. P. Lhomme & O. Ihsane; 7099, -4.8431, 15–16.v.2018, 1♀, white pan trap, leg. P. Lhomme; Casablanca-Settat, Oueled Sghir, 32.8230, -7.6421, 23.ii.2018, 1♂, 1♀, A. Sentil & I.E. Abdouni, all UMONS.
Andrena alchata is known from Morocco and Algeria and was only described from the male sex (
Sampling at both Laanoucer and Oueled Sghir in northern Morocco resulted in the capture of males of A. alchata and females corresponding to A. doursana agadira on the wing at the same time. Other Truncandrena species were present, with females of Andrena ferrugineicrus Dours, 1872 and A. schmiedekneckti Magretti, 1883 captured at Laanoucer and males and females of Andrena varia Pérez, 1895 at Oueled Sghir. However, no males or females of A. doursana citreola Warncke, 1975, the form occurring in this part of Morocco, were recorded. This situation raised the possibility that A. alchata and A. doursana agadira are actually synonymous.
Captured females are darker than A. d. citreola (compare Figs
Warncke did not describe male of A. d. agadira, commenting that they were indistinguishable from other A. doursana subspecies. In contrast, males of A. alchata are easy to distinguish as the white markings on the face of A. alchata are much more extensive, covering the clypeus and the lower paraocular areas (Fig.
A female corresponding to A. d. agadira and a male A. alchata were selected from the same site (Laanoucer, 2.v.2018) for molecular investigation. A cox1 fragment of 263 base pairs was obtained after sequencing. A complete homology between the sequences of the female (GENBANK SUB7440720) and the A. alchata male (SUB7440720) was found, confirming their conspecificity. However, this result raises a difficult issue, as Warncke believed the males of A. d. agadira to be identical to other A. doursana subspecies. This therefore means that either A. alchata and A. d. agadira are not synonymous, and simply very similar morphologically in the female sex, or the undescribed A. d. agadira males are incorrectly associated with the type series females. Given this uncertainty, it is not appropriate to propose synonymy between A. d. agadira and A. alchata until genetic sequences can be obtained from the locus typicus to clarify sex associations in this region. What is clear is that captured females represent A. alchata, but given the large degree of variation in the colour of Truncandrena pubescence, it is possible that they simply resemble females of A. d. agadira. As such, we do not describe females here until this situation can be clarified further.
Morocco: Guelmim-Oued Noun, 10 km E Guelmim, 15–16.iv.1995, 5♂, leg. Ma. Halada, OÖLM; Drâa-Tafilalet, 20 km W Boudnib, 9.iv.1995, 1♂, leg. Ma. Halada, OÖLM.
Previously recorded from Egypt and Tunisia (
Egypt: Min. Agr. Egypt, Dekhela, 20.ii.1917, 1♂, leg. Storey, OÖLM (paratype, illustrated Fig.
Morocco: Oriental, 70 km S Oujda, 8.iv.1995, 1♂, leg. Ma. Halada, OÖLM.
Known from Algeria, Tunisia, Egypt, and Israel (
Morocco: Fès-Meknès, Ifrane environs, 1700 m, 10.v.1997, 1♀, leg. P. Průdek; South of Azrou, 25.iv.2017, 1♂, M. Snižek, both OÖLM.
This is a predominantly central and northern European bee, but there are isolated southern populations in upland areas of Spain, Portugal, and Turkey (
Replacement name for Andrena melaleuca Friese, 1922 nec Andrena melaleuca Pérez, 1895
Morocco: Fès-Meknès, 12 km east of Ifrane, 9.v.1997, 1♀, leg. J. Halada, OÖLM; Fès-Meknès, Laanoucer, 1417 m, 3.v.2018, 1♂, leg. A. Sentil & P. Lhomme, UMONS.
Previously known from Algeria, Tunisia, and Libya (
Morocco: Fès-Meknès, Azrou, 20 km south, 23.iv.2009, 1♀, E. & P. Hajdaj, OÖLM.
A West Mediterranean species, found in Algeria, Italy, Spain, and Tunisia (
Morocco: Fès-Meknès, Bhalil, 10 km NW Sefrou, 28.v.1995, 1♀, leg. Ma. Halada; Fès-Meknès, El-Menzel, 30 km E Sefrou, 29.v.1995, 2♀, leg. Mi. Halada, all OÖLM.
Known from Algeria, Italy (Sardinia), and Tunisia (
Morocco: Drâa-Tafilalet, Er-Rich, 1253 meters, 28.ii.2019, 3♂, 2♀, Moricandia foleyi, leg. O. Ihsane; Fès-Meknès, Laanoucer, 1387 meters, 11.v.2018, 1♀, leg. P. Lhomme & O. Ihsane, all UMONS.
Previously known only from Algeria and Tunisia (
Algeria: Biskra, 3.ii.1997, 1♀, OÖLM; Tunisia: 30 km N Foum Tatahouine, 15.ii.1992, 1♀, leg. Warncke, OÖLM; 21.ii.1992, 1♂, leg. Warncke, OÖLM.
Morocco: Fès-Meknès, south of Azrou, 25.iv.2017, 1♀, leg. M. Snižek, OÖLM; Fès-Meknès, Laanoucer, 1416 m, 11–12.iv.2019, 1♀, white pan trap; 1♀, yellow pan trap, both leg. L. Hamroud & A. Sentil, UMONS; Fès-Meknès, Ain Leuh, Azrou S, 17.iii.1990, 6♀, leg. H. Teunissen, NMNL, Leiden; Fès-Meknès, Col du Zad, 1800 m, 4.iii.1989, 1♀, 2100 m, 10.iii.1989, 2♀, all leg. H. Teunissen, NMNL, Leiden.
Previously known only from central Israel (
Whilst overall the Moroccan material has more extensive reddish tergal margins and darker pubescence, the lack of any major structural differences mean that we consider this material to be conspecific with that from Israel, despite the large degree of geographic separation. This record extends the range of A. menahemella some 3,600 km to the west, giving a disjunct distribution of Morocco and Israel with no records from Algeria, Tunisia, Libya, or Egypt. However, this situation is not unprecedented amongst Andrena, with Andrena aegyptiaca Friese, 1899 showing a disjunct distribution being absent from much of the central part of North Africa (
As the two known specimens were caught in pan traps, no information on floral preferences is available. This reflects the situation in Israel, where females are known only from pan traps (
Israel: Netiv Italamed He, 16.ii.2010, 1♀, leg. G. Pisanty, OÖLM (paratype, illustrated Figs
Morocco: Fès-Meknès, Laanoucer, 1474 meters, 2–3.v.2018, 1♀, yellow pan trap; 33.6234, -4.9002, 11.v.2018, 2♀, all leg. P. Lhomme & O. Ihsane, UMONS.
Described from Algeria (
Algeria: Teniet, 10.v.1895, 1♀, OÖLM (holotype).
Morocco: Tangier-Tétouan-Al Hoceima, Issaguen, 150 km SE Tanger, 1550 m, 12.v.2015, 1♀, leg. Mucska, OÖLM; Fès-Meknès, Ifrane environs, 1700 m, 10.v.1997, 1♀, leg. P. Průdek, OÖLM.
Known from central Europe south into Spain and Greece (
Morocco: Souss-Massa, 10 km W Tiznit, 6.v.1995, 2♀, leg. Mi. Halada, OÖKM; Fès-Meknès, Laanoucer, 33.6302, -4.8847, 2–3.v.2018, 1♂; 10–11.v.2018, 2♀; 15–16.v.2018, 2♀, all caught in yellow pan traps, leg. P. Lhomme & O. Ihsane; Fès-Meknès, Laanoucer, 33.6150, -4.7752, 11–12.iv.2019, 1♀, white pan trap, leg. L. Hamroud & A. Sentil, UMONS.
Originally described from central Spain (
Spain: Montarco, 10.v.1933, leg. Dusmet, 1♀, OÖLM (holotype).
Morocco: Fès-Meknès, 12 km east of Ifrane, 9.v.1997, 1♂, 20♀, leg. J. Halada; Fès-Meknès, Ifrane environs, 9.v.1997, 2♂, 18♀, leg. K. Deneš; Fès-Meknès, Tissa environs, 8.v.1997, 1♀, leg. K. Deneš, all OÖLM.
This taxon has been poorly recorded and documented, leading to nomenclatural confusion. It is part of the Pallandrena subgenus that is characterised by females with plumose scopa on the ventral side of the tibiae (Fig.
The bee is similar to Andrena braunsiana Friese, 1887 which is found in central Europe eastwards to Greece, Turkey and the Caucasus (
Confusion exists over this taxon because the location of the type of Schmiedeknecht collected from Tunisia is unclear and it may be lost (
There are no flower records associated with these specimens, but the unusual modified tibial scopa of the Pallandrena suggests some kind of floral specialisation. None of the specimens had full pollen loads, but fragments could be removed from four females (Fig.
Morocco: Souss-Massa, 30 km north of Tafraoute (Tafraut), 7.v.1995, 1♀, leg. Ma. Halada, OÖLM; Béni Mellal-Khénifra, Khenifra env., 11.v.1997, 1♀, leg. K. Deneš, OÖLM; Casablanca-Settat, Oueled Sghir, 23.ii.2018, 8♂, 11♀; 27.iii.2018, 18♀; 4–5.iv.2018, 1♀, yellow pan trap, all leg. A. Sentil & El Abdouni, UMONS; Casablanca-Settat, Mzamza, Janoubia, 32.9524, -7.5142, 2♀, 6.v.2019, Sinapis arvensis, leg. A. Sentil, UMONS; Fès-Meknès, Sidi Youssef Ben Ahmed, 29.iii.2018, 1♀, leg. P. Lhomme & O. Ihsane, UMONS; Fès-Meknès, Laanoucer, 33.6150, -4.7752, 25.iv.2019, 1♀, leg. L. Hamroud & P. Lhomme, UMONS.
This species was previously known only from a small region of Tunisia and eastern Algeria (
Algeria: Tébessa, Ouenza, 24.iii.2009, 1♀, OÖLM (holotype).
Morocco: Fès-Meknès, Aghbalou, Akourar, 9.v.2019, 1♀, Marrubium vulgare, leg. L. Hamroud & A. Sentil, UMONS.
Known from Algeria, Tunisia, and Libya (
Morocco: Oriental, 40 km south of Guercif, 15–17.v.1995, 100♀, leg. Ma. Halada, OÖLM; Fès-Meknès, Ifkern, 25 km E Boulemane, 24.v.1995, 2♀, leg. Ma. Halada, OÖLM
Previously known only from Spain (
Spain: Aragon, Albarracín, 5.vi.1925, 1♀, OÖLM (holotype); Albacete, Almansa, 25.v.1983, 1♀, leg. H. Teunissen, NMNL; Granada, Pantano de Cubillas, 27.v.1982, 1♀, leg. R. Leys, NMNL; Granada, Pantano de los Bermejales, 26.v.1982, 1♀, leg. R. Leys, NMNL; Almería, Sierra de Maria, 25 km W Lorca, 10.v.2003, 1♂, 6♀, leg. J. Halada, OÖLM; Granada, S-Sierra Nevada, env. Lanjaron, 4.v.2003, 3♀, leg. J. Halada, OÖLM; Zaragoza, Vera de Moncayo, 15.v.1995, 1♂, 1♀, leg. H. & J.E. Wiering, NMNL.
Morocco: Souss-Massa, 20 km north Foum-Zguid, 29–30.iv.1995, 1♀, leg. Ma. Halada, OÖLM; Drâa-Tafilalet, 5 km south of Zagora, 25.iv.1995, 1♀, leg. Ma. Halada, OÖLM.
Described from desert regions of eastern Algeria, Tunisia, and Egypt (Sinai,
Morocco: Fès-Meknès, 20 km north of Missour, 14.v.1995, 1♀, leg. Ma. Halada, OÖLM.
Previously known from Syria, Jordan, and Israel through North Africa to Algeria (
Andrena nitida was recorded from Morocco by
Warncke Collection, OÖLM (Andrena nitida mixtura): Morocco: Ifrane, 18.vii.1931, 1♀, leg. A. Nadig; Koudia, 19.iii.1969, 1♀, leg. J.N. Tasei; Portugal: Carcavelhos, 29.iv.1956, 1♀, leg. N.F. d’Andrade (holotype); Coimbra, Ponte da Portela, 30.iii.1968, 1♂, leg. M.A. Diniz; Spain: Catalonia, Arenys, 15.iv.1929, 1♀, leg. Zariquiey (paratype); Playa de Aro, Gerona, 1♀, leg. H. Pochon (paratype); Catalonia, Beceite, 16.vii.1923, 1♀, leg. Zariquiey (paratype); Alicante, Orihuela, 30.v.1925, 1♀, leg. Andréu (paratype); 8.iv.1925, 1♀, leg. Andréu; 16.vi.1949, 1♂, leg. Andréu; (illegible), 8.vi.1912, 1♂, leg. J.M. Dusmet y Alonso; Barcelona, 7.vii.1898, 1♂; Tunisia: 2 km E Menzel Bourguiba, 28.iii.1976, 1♀, leg. P. Robinson.
Morocco: Souss-Massa, 10 km S Taroudant, 12.iv.1995, 1♀, leg. Mi. Halada, OÖLM; Tangier-Tétouan-Al Hoceima, 3 km Wm Bni Hadifa, 800 m, 15.v.1995, 1♀, leg. Aßmuth, Sanetra & Schulz, OÖLM; Fès-Meknès, 5 km SE Azrou, 31.v.1995, 2♂, leg. Ma. Halada, OÖLM; Drâa-Tafilalet, Ait sais, 23–26.v.2019, 5♀, leg. O. Ihsane, Y. Bencharki, UMONS; Béni Mellal-Khénifra, Aoulou env., 17.v.1997, 1♀, leg. J. Halada, OÖLM; Fès-Meknès, Bhalil, 10 km NW Sefrou, 28.v.1995, 12♂, 24♀, leg. Ma. Halada, OÖLM; Rabat-Salé-Kénitra, Bouknadel, 28.v.2019, 4♀, leg. I. El Abdouni & P. Lhomme, UMONS; Fès-Meknès, Fes, 23.v.1930, 1♂, leg. Werner, OÖLM; Rabat-Salé-Kénitra, Haddada, 12.vi.2018, 1♀; 8.iv-3.vi.2019, 2♀, leg. I. El Abdouni, P. Lhomme & A. Sentil, UMONS; Fès-Meknès, Ifrane, 1670 m, 11.v.2015, 1♀; leg. K. Deneš, OÖLM; Rabat-Salé-Kénitra, Kenitra, 22.vi.1987, 1♂, leg. M. Schwarz, MSC; Fès-Meknès, Laanoucer, 15.v.2018, 1♂, leg. P. Lhomme & O. Ihsane, UMONS; Lot Journu, Abjelil, 8.v.1997, 1♂, leg. P. Průdek, OÖLM; Casablanca-Settat, Mzamza Janoubia, 6–31.v.2019, 12♂, 15♀, leg. A. Sentil, UMONS; Drâa-Tafilalet, Mzizl, 22.v.2019, 1♀, leg. O. Ihsane & Y. Bencharki, UMONS; Fès-Meknès, Oued Sebou, riv, near El-Menzel, 24–27.v.1999, 1♂, 30♀, leg. P. & V. Průdek, OÖLM; Casablanca-Settat, Oueled Sghir, 23.ii-20.vi.2018, 1♂, 2♀, 10.iii-29.vi.2019, 7♂, 10♀, leg. A. Sentil, I. El Abdouni & M. Chokri, UMONS; Drâa-Tafilalet, Sidi Boukil, 24.iv-22.v.2019, 1♂, 18♀, leg. O. Ihsane & Y. Bencharki, UMONS; Drâa-Tafilalet, Tabia, 24.v.2019, 3♀, O. Ihsane & Y. Bencharki, UMONS; Fès-Meknès, Tazzeka N.P., Bab-Bou-Idir env., 28.v.1999, 1♀, leg. P. Průdek, OÖLM.
Andrena fulvicornis was described by Schenck in the same publication as A. nitidiuscula (
At Linz, all examined material in the Warncke Collection from Algeria, Egypt, Morocco, Portugal, Spain, and Tunisia (variably identified by Warncke as A. nitidiuscula or A. nitidiuscula nigellata depending on sampling location) conformed to Andrena fulvicornis sensu Schmid-Egger and Doczkal. Overall, examination of 273 specimens of this species pair from Iberia and North Africa revealed 272 A. fulvicornis [Algeria (2), Egypt (1), Morocco (161), Portugal (57), Spain (16), Tunisia (35)] and a single specimen of A. nitidiuscula from Portugal (
The identity of taxa from this complex described from North Africa previously considered as A. nitidiuscula nigellata sensu
True A. nitidiuscula is almost certainly absent from North Africa, and indeed may be rare in hot areas of Mediterranean Europe, being restricted to areas with a cooler microclimate such as the coastline of northern Portugal (
Senckenberg, Frankfurt (Andrena fulvicornis): no collection details, 1♀ (Neotype, designated Schwenninger 2012); (Andrena nitidiuscula): no collection details, 1♀ (Lectotype, designated Schwenninger 2013); Warncke Collection, OÖLM (all conforming to Andrena fulvicornis): Algeria: Algiers, 2.v.1913, 1♀; Oran, 1895, 1♂, leg. Schmiedeknecht; Egypt: Kerdasa [Kirdasah], 19.v.1929, 1♂, leg. H. Priesner; Portugal: Carcavelos, 13.vi.1953, 1♀, leg. N.F. d’Andrade; Évora, 3.vii.1953, 1♀, leg. N.F. d’Andrade; Sintra, 31.v.1953, ♀, leg. N.F. d’Andrade; Spain: Aranjuez, 26.v.1912, 1♀, leg. Dusmet; Huesca, Benasque, 12.vii.1907, 1♀; Segovia, Madrona, 30.vii.1968, 1♀, leg. K. Warncke; Sierra de Arecena, Rio Odiel dei Calanas, 25.iv.1981, 1♀, M. Kühbander; Tunisia: Tunis, 1898, 1♂, leg. Schmiedeknecht; other collections (Andrena fulvicornis): Tunisia: 30 km N Gabes, 10.iv.1994, 33♀, leg. M. Schwarz, M. Schwarz Colln.; Zana, 6.iv.1965, 1♀, leg. R.T. Simon Thomas, NMNL, ZMA.INS.5087405
This work increases the number of Andrena species known from Morocco by 26, a richness increase of almost 17%. The presence of species previously known only from varying combinations of Spain, Algeria, Tunisia, Libya, Egypt, and Israel, including recently described species like A. menahemella, A. selena, and A. tebessana, previously undescribed species, and species more typically known from Europe such as A. synadelpha indicates that our knowledge of North African Andrena remains incomplete.
Morocco has previously been identified not only as an area supporting a rich bee fauna but also as a hotspot of bee diversity and endemism (
The distribution of sampling locations for species newly described for science (Fig.
Distribution of sampling locations of newly described species A Andrena hebescens (circles), Andrena semiadesus (squares) B Andrena tenebricorpus (diamond), Andrena niveofacies (circle), Andrena triangulivalvis (triangle) C Andrena sparsipunctata (circles), Andrena breviceps (diamonds) D Andrena acutidentis (squares), Andrena farinosoides (triangles), Andrena nigriclypeus (circle). Relief is indicated by shading, measured in meters above sea level.
There are many outstanding problems in North African Andrena arising from the large number of subspecies described or erected by Warncke such as in A. pandosa Warncke, 1968 and A. medeninensis Pérez, 1895 that require focused taxonomic attention to resolve satisfactorily, and which are beyond the scope of this paper. Dealing with these species complexes should form the base for future taxonomic research on Andrena in this region.
We would like to thank Ahlam Sentil, Insafe El Abdouni, Laila Hamroud, Oumayma Ihsane, and Youssef Ben Charki for their significant contribution towards the contemporary insect collections, and to Dr. Stefanie Christmann, leader of the ICARDA Pollination project in Morocco that made all recent sampling possible. Our thanks to Fritz Gusenleitner, Esther Ockermüller, Martin Schwarz for access to the Warncke Collection and hospitality at Linz, to Frederique Bakker for access to Andrena material from Naturalis, and to Maximillian Schwarz for the loan of material from his collection. Our thanks also go to Christophe Praz and Romain Le Divelec for helpful discussion on Mediterranean Andrena, and to Christian Schmid-Egger, Gideon Pisanty, and an anonymous reviewer whose comments greatly improved the manuscript. This research was funded by the Federal German Ministry for the Environment, Nature Conservation and Nuclear Safety (BMU) through the International Climate Initiative (IKI). It was also partly supported by the “Fonds de la Recherche Scientifique – FNRS” and the “Research Foundation of Flanders – FWO” under EOS Project (n°3094785).