Monograph |
Corresponding author: Thomas J. Wood ( thomasjames.wood@umons.ac.be ) Academic editor: Jack Neff
© 2023 Thomas J. Wood.
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 (2023) The genus Andrena Fabricius, 1775 in the Iberian Peninsula (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 96: 241-484. https://doi.org/10.3897/jhr.96.101873
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The Iberian Peninsula is a global hotspot for bee diversity due to its large number of different habitats, particularly Mediterranean scrubland, mountains, and hot and cold steppe. In line with its status as a hotspot of bee diversity, the peninsula hosts a very large Andrena fauna, which despite progress in recent years remains incompletely studied, particularly with reference to genetic investigation. Here the Iberian Andrena fauna is comprehensively revised, resulting in a total of 228 recorded species. Numerous taxonomic changes are necessary following inspection of museum specimens, type material, and genetic investigation. The following subgenera are described: Pruinosandrena subgen. nov., containing six taxa previously placed in the subgenus Campylogaster Dours, 1873, and Blandandrena subgen. nov., Bryandrena subgen. nov., Limbandrena subgen. nov., and Ovandrena subgen. nov., containing one, one, one, and four taxa previously placed in the subgenus Poliandrena Warncke, 1968. Andrena (Limbandrena) toelgiana Friese, 1921 syn. nov. is synonymised with A. (Limbandrena) limbata Eversmann, 1852. The current lectotype of A. (Micrandrena) obsoleta Pérez, 1895 was incorrectly designated by Warncke; the taxon differs from A. obsoleta sensu Warncke, belonging instead to a taxon within the A. mariana Warncke, 1968 complex. A new lectotype is designated for A. obsoleta sp. resurr. from Algeria, and A. mariana solda Warncke, 1974 syn. nov. is synonymised with it; A. (Micrandrena) alma Warncke, 1975 stat. nov., A. (Micrandrena) mica Warncke, 1974 stat. nov., and A. (Micrandrena) tenostra Warncke, 1975 stat. nov. are raised to species status. Andrena (Truncandrena) abunda Warncke, 1974 stat. nov., A. (Micrandrena) lecana Warncke, 1975 stat. nov., A. (Pruinosandrena) parata Warncke, 1967 stat. nov., A. (Micrandrena) pauxilla Stöckhert, 1935 sp. resurr., A. (Pruinosandrena) succinea Dours, 1872 sp. resurr., and A. (Notandrena) varuga Warncke, 1975 stat. nov. are also returned or elevated to species status. A lectotype is designated for A. (Euandrena) lavandulae Pérez, 1902 sp. resurr. which is returned to species status, and A. (Euandrena) impressa Warncke, 1967 syn. nov. is synonymised with it. Andrena (Truncandrena) nigropilosa Warncke, 1967 stat. nov. is elevated to species status, and A. (Truncandrena) truncatilabris espanola Warncke, 1967 syn. nov. is synonymised with it as a junior subjective synonym. A lectotype is designated for A. (Melandrena) vachali Pérez, 1895; A. (Melandrena) creberrima Pérez, 1895 syn. nov. and A. (Melandrena) vachali syn. nov. are synonymised with A. (Melandrena) discors Erichson, 1841, and Andrena (Melandrena) hispania Warncke, 1967 syn. nov. is synonymised with A. (Melandrena) morio Brullé, 1832. Andrena (Pruinosandrena) mayeti Pérez, 1895 syn. nov. is newly synonymised with A. (Pruinosandrena) caroli Pérez, 1895 and A. (incertae sedis) setosa Pérez, 1903 syn. nov. is newly synonymised with A. (incertae sedis) ranunculorum Morawitz, 1877. Andrena (Simandrena) cilissaeformis Pérez, 1895 sp. resurr. is returned to species status, and is the correct name for A. (Simandrena) breviscopa auctorum. Andrena (incertae sedis) breviscopa Pérez, 1895 is returned to synonymy with A. (incertae sedis) numida Lepeletier, 1841, and A. (incertae sedis) inconspicua Morawitz, 1871 is newly synonymised syn. nov. with A. numida. Andrena (Euandrena) isolata sp. nov. and A. (Micrandrena) ortizi sp. nov. are described from the Sierra Nevada (Granada), A. (Truncandrena) ghisbaini sp. nov. is described from Málaga province, and A. (Avandrena) juliae sp. nov. is described from Cádiz province. The males of A. (Micrandrena) alma and A. (?Euandrena) ramosa Wood, 2022 are described. Additional lectotypes are designated for A. (Plastandrena) asperrima Pérez, 1895, A. (Plastandrena) atricapilla Pérez, 1895, A. (Aenandrena) hystrix Schmiedeknecht, 1883, A. (Pruinosandrena) lanuginosa Spinola, 1843, A. (Notandrena) ranunculi Schmiedeknecht, 1883, and A. (Euandrena) symphyti Schmiedeknecht, 1883. Neotypes are designated for A. (Chlorandrena) boyerella Dours, 1872, A. (Notandrena) griseobalteata Dours, 1872, A. (Taeniandrena) poupillieri Dours, 1872, A. (Pruinosandrena) succinea Dours, 1872, and A. (incertae sedis) numida Lepeletier, 1841. Type photographs and diagnostic characters are presented in each case, as well as new dietary information for understudied species. Finally, an identification key is presented in order to facilitate future research on this hyper-diverse genus in one of their global diversity hotspots, and current and future research perspectives for Iberian Andrena are discussed.
Cryptic species, DNA barcoding, Iberian endemic species, solitary bees, taxonomy
The Iberian Peninsula is one of the best places to find and study wild bees globally, with more than 1,000 species present due to its dry and warm climate, abundance of open seasonal habitats, status as a glacial refugium, and rich diversity of flowering plants (
Despite the great number of species present in Iberia, very few authors have worked on this fauna in any detail, certainly compared to that of north-western Africa (e.g.
As a result of these collective efforts, our understanding of Iberian Andrena is as great as it has ever been. However, commencing ecological or taxonomic work on this fauna remains highly challenging due to the lack of accessible identification resources. Valuable information is available in original descriptions, group revisions (e.g.
It is important to briefly discuss species concepts in the context of this work. Because Andrena taxonomy in the West Palaearctic region was dominated by Klaus Warncke in the second half of the 20th Century, it is his classification system that has largely been followed by subsequent workers (e.g.
To date, there has not been a deep discussion of species concepts in Andrena compared to better studied bee groups such as bumble bees (Bombus, e.g.
As such, the present work builds on the morphological species concepts developed by previous workers and integrates genetic and ecological lines of evidence in order to ensure evidence-based species delineation. In this context, subspecies are used pragmatically, following the position of
Andrena specimens were sampled in Iberia and Morocco, predominantly during May–July 2021 in Spain and March–July 2022 in Morocco, but also using specimens collected in previous years. For genetic barcoding, a single midleg was removed from pinned specimens and sent to the Canadian Center for DNA barcoding (CCDB) in Guelph, Canada, for DNA extraction and sequencing (
Phylogenetic trees were supplemented with additional published sequences (e.g.
Taxonomic decisions were informed by analysis of the COI gene. However, decisions were not taken exclusively on the basis of these analyses, as there are a number of inherent limitations when using this fragment to inform taxonomy. The COI fragment itself only represents a single locus of mitochondrial DNA which is inherited matrilineally, can introgress across species boundaries, can transfer to the nuclear genome, cannot detect hybridisation, and can produce topologies that do not represent species trees (e.g.
More broadly, COI analysis is most useful when dealing with species-specific alpha taxonomic decisions. Due to the rate at which this fragment accumulates mutations, whilst closely related species are grouped together, more distantly related species often show no clear structuring and the true evolutionary relationships are not captured. For this, more conserved genes must be sampled, such as by using UCEs. In this work, several new Andrena subgenera are described and illustrative phylogenetic trees are presented based on analysis of the COI fragment; the description of these subgenera is based on the work of
Finally, there are several cases presented here where morphological and genetic data produce ambiguous results concerning the status of certain taxa. Several of these taxa are widespread, and are usually described from outside of the Iberian Peninsula (e.g. see problems with paraphyly of A. hedikae Jäger, 1934). In these cases, no taxonomic action is taken, as it is preferable to have topotypic genetic samples and to consider these taxa across their entire nominal range. Where multiple valid species are potentially present, these species are referred to using the phrase ‘aggregate’ to reflect this situation. In contrast, when species are described from Iberia or have ranges that are restricted to the peninsula or to the West Mediterranean region, taxonomic decisions can be and are made with a greater degree of confidence which reflects the stronger and more complete evidence base available here.
For updating the Iberian Andrena species total, the checklist of
For the identification keys, the female key is partly based on an unpublished key to Iberian Andrena written by Klaus Warncke (in German) that was kindly shared with me by Erwin Scheuchl (Ergolding, Germany). This key contained around 170 species, so considerable modifications were needed to account for the substantially larger faunal total recorded here, as well as accounting for newly described species and other taxonomic changes. The male key is novel, but both the female and male keys have been strongly inspired by the keys of
The geographic scope of the key is limited to the Iberian Peninsula. It cannot be used in North Africa due to the many different or endemic faunal elements found there; for example, it only covers 114 of the 201 (56.7%) Andrena species known from Morocco (Wood in prep.). In a European context, the key can be used in the Balearic Islands, though only a fraction of the species covered in this work occur there. It can be generally used in southern France up to (but not including) the Maritime Alps, but some taxa are missing, such as endemic species (e.g. Andrena (Taeniandrena) vocifera Warncke, 1975) or widespread European species that do not cross the Pyrenees into Iberia (e.g. Andrena (Micrandrena) pusilla Pérez, 1903). However, the existing keys of
No distribution maps are presented as part of this work, as they are for other revisions such as that of
Pollen was removed from female Andrena specimens in order to quantify the pollen foraging niche of understudied species. Specimens were selected from Iberia and other Mediterranean countries when the species’ range extends beyond the peninsula. Pollen was removed, processed, and identified following the methodology of
Morphological terminology follows
The following abbreviations are used in the species descriptions: A = antennal segments, S = metasomal sterna, and T = metasomal terga. Subgeneric concepts follow
AMC Personal collection of Andreas Müller, Wädenswil, Switzerland;
CMHC Carlos M. Herrera collection, Estación Biológica de Doñana, Seville, Spain;
FJOS Personal collection of Francisco Javier Ortiz-Sánchez, El Ejido, Spain;
FLOW FLOWer lab collection, University of Coimbra, Coimbra, Portugal;
MSC Personal collection of Maximillian Schwarz, Ansfelden, Austria;
MZUR Zoological Museum of Sapienza University of Rome, Rome, Italy;
OÖLM Oberösterreiches Landesmusum, Linz, Austria;
SMFD Naturmuseum Senckenberg, Frankfurt am Main, Germany;
TJWC Personal collection of Thomas J. Wood, Mons, Belgium;
UMONS Laboratory of Zoology collection, University of Mons, Mons, Belgium;
Genetic study of Iberian Andrena resulted in a number of important necessary changes to species concepts, as well as further supporting recent decisions. These results are presented here by subgenus; not all Iberian Andrena subgenera are treated here, as no taxonomic problems were detected for the majority of species.
In Iberia, this is a species-poor subgenus, consisting solely of A. fulica Warncke, 1974 and A. vacella Warncke, 1975. Warncke described A. astrella Warncke, 1975 from Iberia, and used it in combination with A. fulica as a subspecies. The two taxa were synonymised by
Phylogenetic tree (maximum likelihood) of Andrena from the subgenera Aciandrena Warncke, 1968, Aenandrena Warncke, 1968 sensu lato, Graecandrena Warncke, 1968, Poecilandrena Hedicke, 1933, and the numida-group based on the mitochondrial COI gene. Andrena (incertae sedis) relata Warncke, 1975 is used as an outgroup. Numbers adjacent to branches represent bootstrap support (values of <75 are omitted).
This subgenus was found to be paraphyletic by
In the true Aenandrena, four species are currently recognised, of which two are widely distributed, A. (Aenandrena) aeneiventris Morawitz, 1872 that was described from Italy and A. (Aenandrena) hedikae Jäger, 1934 that was described from the western Balkans. Both of the widespread species are nominally distributed from Iberia and Morocco to Central Asia in dry and warm parts of the Palaearctic (
This subgenus is clearly supported genetically (
Andrena (Chlorandrena) livens Pérez, 1895 was described from north-eastern Spain. Warncke described A. livens algeria Warncke, 1967 from Tunisia on the basis of slight differences in the genital capsule and sternum eight. He also described A. livens gruenwaldti Warncke, 1967 from Sicily. Based on the sequences here, a Moroccan specimen conforming to A. livens algeria shows relatively low differentiation from Portuguese and Spanish specimens, being separated by an average genetic distance of 2.37% (range 2.16–2.47%). Additional samples are required to investigate the status of these subspecies, and also to sample A. (Chlorandrena) agnata Warncke, 1967 which is a poorly understood species in an Iberian context; I have seen no Iberian specimens, and it is included on the list on the basis of the single female paratype from Madrid, collected 6.vi.1946 by Dusmet (
The status of A. (Chlorandrena) boyerella Dours, 1872 and A. (Chlorandrena) leucolippa Pérez, 1895 and the relationship of these taxa to each other has been confused.
However, this classification is not immediately stable due to the fact that the type series of A. boyerella is lost, as is the case for all of Dours’ types.
Andrena (Chlorandrena) humilis Imhoff, 1832 is the most widespread West Palaearctic Chlorandrena, and it is quite variable over its range. A broad species concept has been used as no consistent morphological differences can be found because of this variability. Sequences from Austria, Belgium, Finland, Germany, Portugal, and Spain formed a broad A. humilis clade (Fig.
Finally, the taraxaci-group (see
Material examined. Andrena leucolippa: France: Riscle [43.6564°N, -0.0894°W], 1♀,
Andrena (Didonia) mucida Kriechbaumer, 1873 is a highly unusual species. It is bivoltine, with the first generation seemingly specialising on Muscari (Asparagaceae) and the second generation specialising on genera from the former Dipsacaceae (now Caprifoliaceae) such as Scabiosa. This specialisation is associated with a morphological change, with females of the first generation possessing tibial scopae composed of simple hairs, and females of the second generation possessing tibial scopae composed of plumose hairs. There is the possibility that these generations may actually represent distinct species, as for putatively bivoltine taxa like A. (Holandrena) decipiens Schenck, 1861 that was found to consist of two taxa (Mandery et al. 2008). However, barcodes from females from the first and second generations from central and southern Spain show that this is not the case, with an average intraspecific genetic distance of 0.30% (range 0.15–0.46%; Fig.
Phylogenetic tree (maximum likelihood) of Andrena from the subgenera Didonia Gribodo, 1894 and Simandrena Pérez, 1890 based on the mitochondrial COI gene. Andrena (Nobandrena) funerea Warncke, 1975 is used as an outgroup. Numbers adjacent to branches represent bootstrap support (values of <75 are omitted).
Additionally, A. (Euandrena) solenopalpa Benoist, 1945 was previously placed in the subgenus Didonia (
Material examined. Andrena mucida: Spain: Guadalajara, Veguillas, 2 km N, Barranco de la Isa, 17.v.2021, 1♀, leg. T.J. Wood, TJWC [BOLD accession number WPATW185-21], on Muscari spp.; Málaga, Benaoján, Cueva del Hundidero, 3.vi.2021, 1♀, leg. T.J. Wood, TJWC [BOLD accession number WPATW257-21], on Scabiosa atropurpurea; Guadalajara, Veguillas, CM-1006, 9.vii.2021, 1♀, leg. T.J. Wood, TJWC [BOLD accession number WPATW326-21], on Scabiosa atropurpurea.
In comparison to the situation in the Eastern Mediterranean (
Euandrena specimens with black and orange pubescence can be found infrequently across north-western Africa. They have typically been referred to as A. bicolor in the literature, but two distinct taxa are present. One corresponds to A. bicolor s.l., but the other is clearly distinct genetically; morphologically it can be distinguished by the structure of the clypeus, with A. bicolor s.l. with the clypeus shiny between the punctures, whereas in the second taxon the puncture interspaces are shagreened and dull, and there is a weak longitudinal furrow, similar to what can be seen in species like A. (Euandrena) angustior, though more apically situated and occupying a shorter distance.
In the original description of A. oraniensis, Lepeletier (1841: 245) draws attention to the colour of the hairs on the hind legs. Specifically, he states that: “cuisses des deux postérieures garnies des poils ferrugineux pales ; leurs jambes et leurs tarses à poils noirs en dessus, ferrugineux en dessous”. This bicoloured tibial scopa (dark dorsally, ferruginous ventrally) does not correspond at all to members of the A. bicolor s.l. group which have uniformly orange tibial scopae. It does however correspond very well to the concept of A. florentina which has a distinctive bicoloured scopa, one of the characters that allows it separation from its sister taxon A. bicolorata (Rossi, 1790). Based on the identifications made by Pérez, who probably saw the original specimen in Lepeletier’s collection and Lepeletier’s original description, the position is taken that A. oraniensis cannot be a Euandrena taxon displaying the colour pattern of A. bicolor s.l. However, without a type, it is undesirable to make A. oraniensis the senior synonym for A. florentina. Andrena oraniensis is therefore declared a nomen dubium until such time as the original syntypic series can be located.
The next oldest available name from North Africa is A. bicolor agraria.
Material examined. Andrena bicolor: Algeria: Theniet El Had [35.8727°N, 2.0007°E] (1♀,
Andrena (Euandrena) lavandulae Pérez, 1902: 156 ♀♂ [France, lectotype by present designation:
Andrena (Euandrena) angustior impressa Warncke, 1967: 234, ♀♂ [Morocco: OÖLM, examined] syn. nov.
Remarks.
In the
Material examined. France: Banyuls [Banyuls-sur-Mer, 42.5658°N, 2.8658°E], 1♂, 1♀,
This subgenus is also species-poor in Iberia, containing only A. impunctata Pérez, 1895, A. montarca Warncke, 1975, A. nebularia Warncke, 1975, and A. verticalis Pérez, 1895. Two taxa are uncommonly collected in Iberia (A. impunctata and A. montarca). Andrena nebularia was considered to be endemic to Spain, but new collections in Morocco have demonstrated its presence in a small part of the Middle Atlas. Genetically, there is almost no differentiation, with the Moroccan specimens separated by 0.26% and 0.52% (Fig.
Material examined. Andrena nebularia: Morocco: Fès-Meknès, Boulemane, 5 km SE, junction of R503 and N4, 1900 m, 19.v.2022, 2♀, leg. T.J. Wood, TJWC; Fès-Meknès, Boulemane, R503, 7 km SE of Boulemane, 1900 m, 1♂, 6♀, 22.v.2022, leg. T.J. Wood, TJWC; Fès-Meknès, Boulemane, R503, SE of Ait Karmosse, 1750 m, 22.v.2022, 1♂, leg. T.J. Wood, TJWC.
Andrena fuscosa was described from southern Spain by
Phylogenetic tree (maximum likelihood) of Andrena from the subgenera Melanapis Cameron, 1902, Plastandrena Hedicke, 1933, and Suandrena Warncke, 1968 based on the mitochondrial COI gene. Andrena (incertae sedis) innesi Gribodo, 1894 is used as an outgroup. Numbers adjacent to branches represent bootstrap support (values of <75 are omitted).
This subgenus is strongly derived within Andrena, but it shows substantial morphological variation that has led to the description of the subgenera Hyperandrena Pittioni, 1948 and Zonandrena Hedicke, 1933, both of which are now placed within an expanded Melandrena. There are several problems within this subgenus in an Iberian context.
Andrena (Melandrena) morio Brullé, 1832: 353, ♀♂ [Greece:
Andrena (Melandrena) hispania Warncke, 1967: 212, ♀♂ [Spain: OÖLM, examined] syn. nov.
Remarks. Andrena hispania Warncke, 1967 was described from Algeciras in southern Spain. The identification characters given by Warncke are comparatively weak and rely on hair colour and the degree of infuscation of the wings, without mentioning definitive structural characters. Genetically, the three A. hispania sequences from Spain and Portugal mixed with A. morio sequences from Israel, Morocco, Portugal, Spain, Tunisia, and Turkey without forming a cluster, this group having bootstrap support of 100 (Fig.
Remarks. The specific differences between these taxa is unclear across southern Europe. In some northern countries, only two taxa are present (A. nitida and A. thoracica, e.g. the United Kingdom), with no introgression observed. Andrena nitida flies only in the spring, whereas A. thoracica is bivoltine, flying in both the spring and the summer. In Central Europe, A. limata can be found, this taxon also being bivoltine. Differentiation between the three taxa in Central Europe has often utilised hair colouration characters, as in this region the three taxa are separable with reference to the hairs on the sides of the mesosoma (light in A. nitida and A. limata, dark in A. thoracica) and the hairs of the face and the tibial scopal (light and dark in A. nitida, uniformly dark in A. limata and A. thoracica). There are additional characters such as the colour of the hind tibial spur and the density of the punctures T1, but these are less commonly referred to; both A. nitida and A. limata have dense punctures on T1 (separated by up to 1 puncture diameter), whereas they are more clearly spaced in A. thoracica (punctures separated by 1–2 puncture diameters). It is important to note that A. limata is a replacement name for A. lucida Lepeletier, 1841 nec. A. lucida Panzer, 1798 which is nominally a synonym of A. bicolor Fabricius, 1775 but this must be established (see section on subgenus Euandrena). Andrena lucida Lepeletier was described from ‘France’, without further details. I have not been able to inspect the type which should be in the
The situation in southern Europe is much more challenging. In south-western Europe, A. limata becomes much darker, and therefore closely resembles the colour form of A. thoracica, with extensive black pubescence on the mesosoma laterally. This colour form was described as A. limata mixtura Warncke, 1967 (illustrated by
Analysis of barcodes does not provide clarity (Fig.
Andrena (Melandrena) discors Erichson, 1841: 192, ♀ [Algeria:
Andrena (Melandrena) creberrima Pérez, 1895: 46, ♀♂ [Algeria:
Andrena (Melandrena) vachali Pérez, 1895: ♀ [Tunisia, lectotype by present designation:
Andrena (Melandrena) bodemeyeri Benoist, 1969: 246, ♀ [Tunisia:
Remarks. There has been extensive confusion between A. discors and A. creberrima.
The situation is confused due to the status of two additional taxa described by Pérez, A. (Melandrena) creberrima Pérez, 1895 and A. (Melandrena) vachali Pérez, 1895. Andrena creberrima was described from Algeria like A. discors.
Grouping A. creberrima under A. discors is straightforward. The problem comes with A. vachali which was classically considered to be distinct from A. discors and with a greater range extending from the Canary Islands to southern Israel (
Taking a step back, the differentiation between A. creberrima, A. discors, and A. vachali has almost always been based on colouration, with A. discors the darkest, A. vachali the lightest, and A. creberrima somewhat intermediate. Males are generally rarer in collections; taken together, I have examined 215 female specimens, but only 73 male specimens of this discors-creberrima-vachali group. However, examination of males has led me to the conclusion that there are no apparent differences in the males of A. creberrima, A. discors, and A. vachali, and indeed the male of only one of these nominal taxa was actually originally described. All have white hairs over the majority of the face with clear black hairs laterally along the inner margins of the compound eyes, a genital capsule that is typical of the former Zonandrena with the dorsal surface of the gonocoxae with granular shagreen, with the penis valves moderately broad, and without an emargination in the outer margins of the gonostyli (Fig.
Based on this colour variation, the lack of variation in the male genital capsule, the unclear and overlapping distributions given by Warncke (A. creberrima and A. vachali both reported from Crete, A. creberrima and A. discors both reported from Algeria), and the very low genetic distance between Portuguese and Israeli specimens, both A. creberrima syn. nov. and A. vachali syn. nov. are synonymised with A. discors. This resolves this long-running confusion as to the correct name and identity of this taxon (
Distribution. Portugal, Spain (Canary Islands, mainland), Morocco, Algeria, Tunisia, Italy, Malta, Libya, Greece (Crete), Egypt, Israel, Jordan.
Material examined (illustrative). Algeria: label information illegible, possibly ‘Bone’ [= Annaba], 1♀,
This is by far the most species-rich subgenus in Iberia, containing 37 species.
The treatment of A. obsoleta has a long and confused history that has caused many problems.
Warncke described A. mariana s. str. from the island of Fuerteventura in the Canary Islands, stating that the species could potentially be found in Morocco (
Examination of A. mariana solda material reveals that it is conspecific with the newly designated lectotype of A. obsoleta and is synonymised with it syn. nov. As identified by Warncke, the taxon has a wide distribution across Mediterranean parts of Morocco, Algeria, and Tunisia (
As A. mariana solda is actually a synonym of a valid species that was described prior to A. (Micrandrena) mariana s. str., this has implications for the species-status of the other taxa lumped under A. mariana by Warncke. Genetic sequences were available for A. mariana mica Warncke, 1974 from southern Morocco, A. mariana s. str. from south-western Morocco, and A. mariana alma Warncke, 1975 from Portugal (locus typicus near Córdoba, southern Spain; Fig.
Warncke described two further subspecies of A. mariana: A. mariana leptura from Egypt and A. mariana tenostra from south-eastern Spain, the latter specifically from a single female specimen from Villajoyosa in the province of Alicante, then listing additional specimens from Almería and Murcia (
Finally, a number of specimens were found in the very south of Spain (Málaga, Sevilla) which show a morphology very close to that of A. mica. However, ecologically this does not make sense as A. mica is not known from the more humid and Mediterranean areas north of the Atlas Mountains. Examination of these specimens shows that A3 is much longer than A4+5, whereas it is as long as A4+5 in A. mica. These specimens probably represent an additional undescribed species in the A. mariana complex, but they are not described at this time, as it would be beneficial to have barcoded specimens to confirm this differentiation. They are therefore referred to as aff mica in the identification key.
It is important to note that this material does not belong to A. abjecta, the status of which in Iberia is unclear.
Material examined. Andrena alma: Portugal: Albandeira, near Lagoa, 20.iv.2005, 1♀, leg. D.W. Baldock, TJWC; Algarve, Casaqueimada (7 km N of Silves), 20.iii.1995, 1♀, leg. T. & M. Simon Thomas,
Andrena mariana s. str.: Algeria: Saida, 15 km S of Sfissifa, Ben Ikhou, st. 6, 6.iv.1983, 6♀, leg. R. Leys & P. v. d. Hurk,
Andrena mica: Algeria: Ghardaia [32.5047°N, 3.6419°E], 1♀, OÖLM (holotype); Morocco: 30 km E Midelt, 13.v.1995, 1♀, leg. Mi. Halada, OÖLM; Drâa-Tafilalet, Ouarzazate, P1506, Telouet, Adaha, 1700 m, 18.iv.2022, 6♀, leg. T.J. Wood, TJWC; Drâa-Tafilalet, Ouarzazate, 2 km W Agouim, 1800 m, 13.iv.2022, 3♀, leg. T.J. Wood, TJWC; Ifkern, 25 km E Boulemane, 24.v.1995, 1♀, leg. Mi. Halada, OÖLM; M’rirt (30 km N), 11.iii.1989, 1♀, leg. H. Teunissen,
Andrena aff mica: Spain: 40 km W Málaga, Yunquera, 800 m, 29.iv.2003, 5♀, leg. J. Halada, OÖLM/TJWC; La Corchuela (Dos Hermanas, Sevilla), 35 m, 27.iii.2009, 2♂, 2♀, leg. F.J. Ortiz-Sánchez, FJOS; Río Blanco, Aguadulce (Sevilla), 300 m, 17.v.2008, 1♀, leg. F.J. Ortiz-Sánchez, FJOS.
Andrena nitidula: Algeria: Babor, 1♀,
Andrena obsoleta: Algeria: Bône [=Annaba, 36.9092°N, 7.7264°E], 1♀,
Andrena tenostra: Spain: Villajoyosa [38.5097°N, -0.2299°E], 11.v.1936, 1♀, leg. Andréu, OÖLM (holotype); Salobreña, Granada, 8.v.1983, 3♀, leg. W. Perrandin, OÖLM/TJWC.
The status of these two species has been extensively argued over, and despite much attention the position remains somewhat unclear.
Andrena distinguenda was described from western Germany, and A. nitidula was described from south-western France (lectotype from Bordeaux [though this is not indicated on the specimen, it bears the number ‘675’ which refers to the entry for A. nitidula in the catalogue of Pérez], designated by
Barcode analysis complicates this matter further (Fig.
Morphologically, both Spanish and Moroccan specimens falling into the A. distinguenda clade conform to the concept of A. nitidula using the criteria specified by
Inspection of Micrandrena specimens from high altitude in the Sierra Nevada revealed the presence of a species that morphologically resembles A. (Micrandrena) rugulosa Stöckhert, 1935 due to its head that is only slightly shorter than wide rather than clearly shorter than wide, an unusual character in Micrandrena. This finding is remarkable, because although widely distributed in Central and Eastern Europe, A. rugulosa has a western limit in the Swiss Alps, and has not been previously recorded from France, Spain, or Portugal (
The Sierra Nevada sequences were separated from A. atlantea by an average genetic distance of 6.04% (range 5.71–6.38%) and from the undescribed Micrandrena by an average genetic distance of 6.34% (range 6.31–6.38%). These three species would therefore seem to represent an isolated Micrandrena lineage that is restricted to the Sierra Nevada and the High and Middle Atlas Mountains in Morocco, with consequent genetic and morphological divergence. This is the same pattern as observed in the subgenus Euandrena, suggesting that the Sierra Nevada hosts the remnants of a fauna that was presumably once more widespread across Iberia and North Africa. Additional sampling and genetic analysis is required to determine if this pattern holds true for other bee groups. The new Micrandrena species from the Sierra Nevada is described below, and the undescribed Micrandrena species from Morocco will be described in an upcoming publication.
Andrena niveata was described from Germany and Hungary, without further information (
Examination of material from Spain shows that the two taxa are distinct, but morphological separation is challenging, may not be possible in all cases when old, abraded, or dirty specimens are available, and is best made with reference to the male genital capsule. Taking male specimens, the shape of the gonostyli are distinctive. In A. niveata s. str., the gonostyli are elongate, with the inner margins of the gonostyli only weakly produced towards the penis valves (Fig.
Andrena (Micrandrena) niveata Friese, 1887 A male genital capsule C male terga, dorsal view E female propodeal triangle G female terga, dorsal view; Andrena (Micrandrena) lecana Warncke, 1975 B male genital capsule D male terga, dorsal view F female propodeal triangle H female terga, dorsal view.
The position is taken here that the strength of tergal shagreenation in A. niveata s. str. females is variable across Europe, but that A. lecana stat. nov. is a valid species based on the overlapping range in combination with the consistent difference in the shape of the male genital capsule. It has a distribution across steppic parts of central Spain, extending into mountainous areas in south-eastern Spain. Females displaying any level of shagreenation on the tergal discs are considered to represent Iberian populations of A. niveata s. str., and females with completely polished tergal discs without a trace of shagreenation represent A. lecana (see identification key). Future genetic investigation using more targeted primers will be necessary to confirm this position. Finally, specimens of A. lecana from high altitude in the Sierra Nevada show slightly different antennal ratios in the male sex, though the male genital capsule is otherwise identical; this requires further investigation.
Material examined. Andrena lecana: Spain: Ribas [Rivas-Vaciamadrid, 40.3503°N, -3.5390°E], 6.v.1908, 1♀, leg. Dusmet, OÖLM (holotype); Carboneras de Guadazaón (Cuenca), 1030 m, 16.v.2009, 1♀, leg. F.J. Ortiz-Sánchez, FJOS; Guadalajara, Alcolea del Pinar, 12.v.2021, 1♀, leg. T.J. Wood, TJWC; Guadalajara, Lupiana, 12.v.2021, 1♀, leg. T.J. Wood, TJWC; Huéscar (Granada), 1900, 1♂, leg. Escalera, OÖLM (paratype); La Cabrilla, Sierra Cazorla (Jaén), 1600 m, 1–3.vi.2022, 1♀, leg. C.M. Hererra, CMHC; Madrid, Rivas-Vaciamadrid, Canal de Manzanares to Camino de Uclés, 19.v.2021, 2♀, leg. T.J. Wood, TJWC; Órgiva, N, 1300 m, Sierra Nevada, 26.vi.1988, 1♀, leg. M. Schwarz, OÖLM; Pozuelo, La Fuente, 1♂, OÖLM (paratype); Segovia, Madrona, 500 m NE, Arroyo del Hocino, 15.v.2021, 2♀, leg. T.J. Wood, TJWC; Sierra Nevada, Trevélez, Refugio La Campiñuela, 2400 m, 14.vi.2021, 7♂, 2♀, leg. T.J. Wood, TJWC.
Andrena niveata s. str.: Spain: Cáceres, Cuacos de Yuste, 500 m, 11.v.1999, 1♀, leg. H. & J.E. Wiering,
Members of this species group are challenging to identify and have been inconsistently treated in the literature, with variable species concepts. Andrena spreta was described from Algeria, with
This position was not adopted by subsequent authors.
Analysis of barcodes provides unambiguous support for the existence of three distinct species (Fig.
A single barcode was available from Moroccan specimens from the Middle Atlas tentatively identified as A. pauxilla which showed an average genetic distance to European A. pauxilla specimens of 2.00% (range 1.48%–2.29%). This is considered to represent only separation by distance, and thus A. pauxilla is recorded for the first time in North Africa and unambiguously recorded in Spain. Within Spain, A. pauxilla appears to be principally recorded from mountain ranges such as the Sierra de las Nieves, the Sierra Nevada (Fig.
Andrena (Micrandrena) pauxilla Stöckhert, 1935 A habitat, Granada, Sierra Nevada, Mirador Monte Ahí de Cara, 2100 m, 12.vi.2021 B female collecting pollen from Vella spinosa (Brassicaceae) C habitat, Cuenca, Mirador Valle de Valdecabras, 21.vi.2021 D female collecting pollen from Sedum spp. (Crassulaceae).
Material examined. Andrena curtula: Spain: Barcelona [41.4028°N, 2.1332°E], 1♀,
Andrena pauxilla: France: Bischenberg, 28.vi.1936, 1♀, leg. M. Klein, det. E. Stöckhert, OÖLM; Hausbergen, 29.vi.1930, 1♀, leg. M. Klein, det. H.R. Schwenninger, OÖLM; Morocco: Fès-Meknès, Boulemane, R503, 7 km SE of Boulemane, 1900 m, 22.v.2022, 1♂, 1♀, leg. T.J. Wood, TJWC; Spain: Canet de Mar, 26.iii.1963, 1♀, leg. F. Vergés, det. H.R. Schwenninger, OÖLM; Cuenca, Huerta del Marquesado, environs north of town, 26.vi.2021, 3♀, leg. T.J. Wood, TJWC; Granada, Sierra Nevada, Jardín Botánico Hoya de Pedraza environs, 1900 m, 9.vi.2021, 1♀, leg. T.J. Wood, TJWC; Granada, Sierra Nevada, Mirador Monte Ahí de Cara, 2100 m, 12.vi.2021, 1♀, leg. T.J. Wood, TJWC; Guadalajara, Aldeanueva de Atienza, 9.vii.2021, 1♀, leg. T.J. Wood, TJWC; Málaga, PN Sierra de las Nieves, mountain peak S of Pinsapo Escalereta, 30.v.2021, 1♀, leg. T.J. Wood, TJWC; La Cabrilla, Sierra Cazorla (Jaén), 1600 m, 3.vi.2022, leg. C.M. Herrera, CMHC; Sierra Cazorla, Puerto Llano, 1800 m, 11.vi.2022, 1♂, 3♀, leg. J. Valverde, CMHC; Teruel, Guadalaviar, Rambla de los Ojos, 27.vi.2021, 1♀, leg, T.J. Wood, TJWC; Teruel, Villar del Cobo, Barranco de los Oncenachos, 27.vi.2021, 6♀, leg, T.J. Wood, TJWC; Cuenca, Mirador Valle de Valdecabras, 21.vi.2021, 2♀, leg, T.J. Wood, TJWC.
Andrena pusilla: France: Nantes [47.2233°N, -1.5542°W], 1♀,
Andrena spreta: Algeria: Biskra [34.8600°N, 5.6995°E], 1♀,
Andrena strohmella was described from southern Germany and is a typically early spring species in the Central European Andrena fauna, with records extending south to the High and Maritime Alps in France, and west to the Bordeaux region; it has not previously been reported from the Pyrenees (
More broadly, morphological differences between A. strohmella and A. icterina are slight, though they do not appear to introgress based on examined specimens. Some characters such as the strength of the carinae on the dorsolateral corners of the first tergum are not completely consistent, because some specimens in Central Europe can be found in which these are very weakly produced. Genetic data will hopefully clarify the status of A. icterina, but unfortunately, like A. lecana, this taxon appears to be challenging to barcode, as all seven Iberian specimens sent for genetic analysis failed or returned corrupted sequences.
Material examined. Andrena icterina: Portugal: Bragança, Serapicos, 16.v.2021, 1♀, leg. A. Soares, A. Soares Coll.; Spain: Ávila, Hoyocasero, El Pinar de Hoyocasero, 16.v.2021, 1♀, leg. T.J. Wood, TJWC; Ávila, Navalsauz, 1 km E, Alberche stream, 16.v.2021, 1♀, leg. T.J. Wood, TJWC; Cádiz prov., Vent. L. Canillas HozgargantaTal b. Jimena 250 m, 14.iv.1985, 3♀, leg. W. Schacht, OÖLM; Campamento Alfaguara (Alfacar, Granada), 1420 m, 13.v.2007, 1♂, 2♀, leg. F.J. Ortiz-Sánchez, FJOS; Cortijo Tortas, Paterna del Madera (Albacete), 1310 m, 30.iv.2022, 2♂, leg. F.J. Ortiz-Sánchez, FJOS; Granada, Sierra de Baza, Prados del Rey, 2000 m, 19.vi.2021, 1♀, leg. T.J. Wood, TJWC; Granada, Sierra Nevada, Capileira to La Cebadilla, 1500 m, 8.vi.2021, 1♀, leg. T.J. Wood, TJWC; Granada, Sierra Nevada, Jardín Botánico Hoya de Pedraza environs, 1900 m, 2♀, leg. T.J. Wood, TJWC; Granada, Venta de los Alazores, 25.v.1982, 1♀, leg. R. Leys,
The definition of this subgenus was expanded by
Andrena (incertae sedis) urdula Warncke, 1965 was described from Greece and is a rare and poorly understood taxon known only from a small number of specimens. It is reliably known only from Greece (type series), Spain (central Spain), and Morocco, as the distribution map presented by
Warncke described two similar taxa in the group of small metallic green Notandrena, A. (Notandrena) reperta Warncke, 1974 and A. (Notandrena) reperta varuga Warncke, 1975. Warncke actually described A. reperta as a subspecies of A. varuga, but due to the order of publication, A. reperta has priority. Both species can be recognised because the hind tibial spur is apically curved, though this character is more pronounced in Iberian specimens. Genetically, specimens of A. reperta from Morocco and A. reperta varuga from central Spain formed a clade with a specimen of A. (Notandrena) nigroviridula Dours, 1873 from Morocco. All three taxa were well separated; A. reperta and A. reperta varuga by 12.84%, A. reperta and A. nigroviridula by 11.67%, and A. reperta varuga and A. nigroviridula by 10.12% (Fig.
Andrena (Notandrena) fulvicornis Schenck, 1861 has been recognised as distinct from A. (Notandrena) nitidiuscula Schenck, 1853 (
Material examined. Andrena nigrospina: Portugal: Minho, Ruivães, N103, 12.v.2019, 1♂, 1♀, leg. Wood, TJWC; Spain: Cuenca, Pajaroncillo, 3 km SW, Arroyo de Peña Quebrada, 26.vi.2021, 4♀, leg. T.J. Wood, TJWC (barcoded); Guadalajara, Bustares, 2 km N, Alto Rey, 1780 m, 1♀, 9.vii.2021, leg. T.J. Wood, TJWC (barcoded).
Andrena agilissima is a widespread West Palaearctic species that is well-known in Central and Southern Europe to north-western Africa. In contrast, A. asperrima is much less well known, having a more Mediterranean distribution in France, Spain, Morocco, Algeria, and Tunisia. Unlike A. agilissima, A. asperrima is bivoltine and is exceptionally variable in the density, size, and strength of the integumental punctation. In the female sex, the typical form has strong and dense punctures on the terga, allowing easy separation from A. agilissima in which the terga have small and subtle punctures. However, many specimens of A. asperrima can be found which have greatly reduced tergal punctation and which are therefore extremely similar to A. agilissima; they can be separated by the smaller body size and the sparser punctation of the scutum. This sparsely punctate form is more common in the south-west and was described from Morocco by Warncke as A. asperrima alascana Warncke, 1974. Overall, the two species are clearly separable by their genital capsules in the male sex.
Because of this variation, it is important to ensure that Iberian material is conspecific with North African material, since the oldest names of
Andrena atricapilla Pérez, 1895 was also described from Algeria, but only in the male sex.
Genetic analysis of specimens of A. agilissima and A. asperrima from Croatia, Morocco, Portugal, and Spain showed two clear clades (Fig.
However, what can be concluded is that Iberian material of A. asperrima is conspecific with North African material, including the weakly punctate form that dominates in Morocco. In combination with the new lectotype designations, the invariant genital capsule, and these genetic results, the concept of
Andrena (Plastandrena) asperrima Pérez, 1895: 33, ♀♂ [Algeria, lectotype by present designation:
Andrena (Plastandrena) atricapilla Pérez, 1895: 33, ♂ [Algeria, lectotype by present designation:
Andrena (Plastandrena) flessae var. elcheensis Friese, 1922: 211, ♀ [Spain:
Andrena (Plastandrena) hemicyanea Cockerell, 1930: 112, ♀ [Tunisia: type lost?]
Andrena (Plastandrena) asperrima alascana Warncke, 1974: 36, ♀♂ [Morocco: OÖLM, examined].
Distribution. Spain, France, Morocco, Algeria, Tunisia.
Material examined. Algeria: Constantine [36.3645°N, 6.6409°E], 1♀,
This subgenus is strongly polyphyletic (
Collection of material from the Sierra Nevada produced red-marked Poecilandrena females [WPATW281-21] that morphologically resemble A. potentillae in the reduced punctation density at the edge of the clypeus. No ‘potentillae’ males with their distinctive genital capsule could be found. A female sequence clearly fell into a clade with an A. labiata sequence from Belgium, the two specimens separated by 2.87% (Fig.
Finally,
Barcode analysis returned Simandrena as paraphyletic (Fig.
Andrena (Simandrena) vetula Lepeletier, 1841 was recently placed in the Simandrena, as its unusual male morphology had led to confused previous placement (
The species pair of A. (Simandrena) confinis Stöckhert, 1930 and A. (Simandrena) congruens Schmiedeknecht, 1884 continues to pose problems.
Andrena (Simandrena) cilissaeformis Pérez, 1895: 42, ♀ [Spain, lectotype by present designation:
Andrena breviscopa auctorum.
Remarks. Andrena breviscopa Pérez, 1895 was described in the female and male sexes from North Africa. Warncke’s treatment of A. breviscopa is curious, because he designated a lectotype (Fig.
The use of the name A. breviscopa to apply to the taxon present in Spain, Morocco, and Algeria is therefore incorrect. The correct name is A. cilissaeformis Pérez, 1895 sp. resurr. Andrena cilissaeformis was described from Spain, not Algeria as stated in
Distribution. Spain, Morocco, Algeria.
Material examined. Andrena breviscopa: Algeria: Ghardaia [32.5047°N, 3.6419°E], 1♀,
Large parts of this subgenus have been revised recently by
Two further issues require discussion. The first is the identity of A. poupillieri.
Warncke changed his mind about the status of A. poupillieri – in
Material examined. Spain: Málaga, Estepona, 24.iii.1986, 1♂, leg. J. van Oosterhout,
Material examined. France: Hautes-Pyrénées, Eget Cité, 4.v.2017, 1♂, leg. R. Rudelle, R. Rudelle Colln.; Spain: Huesca, San Juan de la Peña, 14.v.1995, 1♂, leg. H. & J.E. Wiering,
This subgenus contains species that often vary extensively in the colouration of their pubescence, sometimes display minimal variation in structural characteristics in the female sex, and can sometimes only be reliably identified in the male sex. These identification difficulties have led to a large number of subspecific concepts in the literature, the integrity of which must be examined using molecular data. There are a number of taxonomic changes to make which affect the Iberian and more broadly West Mediterranean fauna.
This nominal species is highly variable across its range which was previously considered to be from Morocco and Iberia to Turkey and the Levant. Andrena doursana was originally described from Algeria, and
The differentiation between these remaining subspecies relies on the colour of the female pubescence, as there are no structural differences in the males; indeed, the subspecies A. d. agadira and A. d. bengasia were described only from the female sex. Andrena d. citreola is bright, and has predominantly white hairs on the face and a light brown terminal fringe with scattered white hairs laterally. Andrena d. agadira is much darker, with dark facial hairs and a uniformly dark terminal fringe.
Analysis of barcodes from southern and northern Morocco and Iberia shows that female specimens identified as A. doursana s. str., A. d. agadira, and A. d. citreola did not form differentiated clades (Fig.
More broadly, this A. doursana clade had bootstrap support of 78, and was sister to the A. alchata sequences generated by
Andrena medeninensis was described from Tunisia, and like A. doursana, it nominally displays great variation across its range from Morocco and Iberia to Turkey and the Levant.
Sequences of A. medeninensis s. str. and A. m. tiznita formed a clade with bootstrap support of 91 (Fig.
As it was unfortunately not possible to sample the Iberian subspecies A. m. donata, and no genetic sequences are available from Tunisia, the locus typicus for A. medeninensis s. str., no further taxonomic action is taken here. Given the large genetic difference displayed by A. abunda despite almost no morphological differentiation (at least in the female sex), it is difficult to comment on the Iberian subspecies which simply appears to be a colour variant of this nominally widespread species.
Andrena truncatilabris is a widespread species that was originally described from the Caucasus from what is today Armenia (
Sequences from specimens from Spain and Morocco showed almost no genetic differentiation, with an average intraspecific distance of 0.99% (range 0.00–1.85%; Fig.
Given this genetic difference, it is clear that specimens from Iberia and north-western Africa are both conspecific and distinct from A. truncatilabris s. str. Given this distribution, the use of the name A. t. espanola is undesirable, and so A. nigropilosa stat. nov. is elevated to species status and A. t. espanola syn. nov. is synonymised with it as a subjective junior synonym, as the two names were described in the same publication. The updated synonymy is therefore as follows:
Andrena (Truncandrena) truncatilabris nigropilosa Warncke, 1967: 225, ♀♂ [Algeria: OÖLM, examined].
Andrena (Truncandrena) truncatilabris espanola Warncke, 1967: 224, ♀♂ [Spain: OÖLM, examined] syn. nov.
Distribution. Portugal, Spain, France, Morocco, Algeria, Tunisia (newly recorded). Material from south-eastern France and northern Italy must be carefully revised, but the position is taken here that the Maritime Alps represent a barrier between A. nigropilosa and A. truncatilabris s. str. This should be confirmed with genetic evidence.
Material examined. Algeria: S. Algeria, Laghouat [33.8082°N, 2.8316°E], iii.–iv.1929, 1♀, leg. Meyer, OÖLM (holotype of A. t. nigropilosa); Tlemcen, 20.iv.1910, 1♂, leg. de Bergeoin, OÖLM (paratype of A. t. nigropilosa); Spain: Sierra Nevada [37.0732°N, -3.3948°E], vi.1891, 1♀, leg. Handl., OÖLM (holotype of A. t. espanola); Montarco, 28.iv.1924, 1♂, leg. J.M. Dusmet y Alonso, OÖLM (paratype of A. t. espanola); Tunisia: Kef, 5 km SW Touiref, 28.iv.2012, 41, leg. C. Sevidy & A. Müller, AMC/TJWC.
Material examined. Spain: Barcelona [41.4028°N, 2.1332°E], 1♀,
As a result of the ground-breaking analysis of
In Iberia, representatives of all five subgenera can be found. These can be broadly summarised as the blanda-group, the florea-group, the limbata-group, the oviventris-group, and the relata-group. Four of these five lineages are represented in the analysis of
No action is currently taken for members of the relata-group, as the status of morphologically similar species in the Eastern Mediterranean to Central Asia is unclear, and it must be genetically demonstrated if they belong to the relata-group or not. The members of the newly described subgenera are detailed below; in an Iberian context, the following species can be considered to be part of the relata-group: A. corax, A. laurivora Warncke, 1974, A. macroptera Warncke, 1974, A. murana, and A. relata.
Members of this group of species have been placed in the subgenus Campylogaster Dours, 1873 (
Phylogenetic tree (maximum likelihood) of Andrena from the subgenera Brachyandrena Pittioni, 1948, Lepidandrena Hedicke, 1933, and currently undescribed subgenera based on the mitochondrial COI gene. Andrena (incertae sedis) relata Warncke, 1975 is used as an outgroup. Numbers adjacent to branches represent bootstrap support (values of <75 are omitted).
Finally, clarity is required for the status of taxa lumped under a broad concept of A. pruinosa Erichson, 1835, specifically A. pruinosa succinea Dours, 1872 and A. pruinosa parata Warncke, 1967.
Genetic analysis of members of the Pruinosandrena demonstrates that the broad concept of A. pruinosa used by Warncke was overly conservative (Fig.
Andrena pruinosa succinea was strongly separated from A. pruinosa s. str. by an average genetic distance of 9.45% (range 8.97–10.33%). Sequences of A. pruinosa s. str. were identical, which is not surprising as they all came from a small part of the province of Madrid. Andrena pruinosa succinea samples came from a large geographic area from south-western Morocco to Israel, but still showed low average intraspecific variation of 2.06% (range 0.14–3.80%). The two clades were not sister, being separated by A. parata and A. caroli, and were supported by bootstrap support of 99 and 95, respectively. Andrena succinea sp. resurr. is therefore considered to be a valid species, distinct from A. pruinosa s. str. Morphologically, separation of males is straightforward, and it is unclear why Warncke considered the difference unclear. Andrena succinea males have a yellow marked clypeus (see illustrations in
There are also ecological differences. Andrena succinea can be found in dry desert-edge steppe habitats, as opposed to A. pruinosa which in Iberia is found in grasslands and cold steppe that are lightly more lush, at least during the spring. For example, in Morocco, A. succinea can be found in stipa steppe habitat around Bou Rached (Oriental region, south of Guercif) on the eastern edge of the Middle Atlas as it transitions into the desert (Fig.
Andrena (Pruinosandrena) succinea Dours, 1872 A habitat, Oriental, Guercif, P5427, 2 km SW of Bou Rached, 950 m, 13.v.2022 B female collecting pollen from Brassicaceae spp.; Andrena (Pruinosandrena) pruinosa Erichson, 1835 C habitat, Madrid, Chinchón, 6 km N, M-311, 14.v.2021 D male, in hand.
Although the type of A. succinea is lost, and the type for a more recently described taxon is preserved in the
Andrena pruinosa parata Warncke, 1967: 233, ♀♂ [Spain: OÖLM, examined].
Remarks. Though described from south-eastern Spain,
Distribution. Spain.
Material examined. Spain: Alicante [38.3628°N, -0.5093°W], 1♂, leg. G. Mercet, OÖLM (holotype); Benidorm, 2.vi.1952, 1♀, leg. J. de Beaumont, OÖLM (paratype); Fortuna [Murcia], v.1928, 1♂, leg. J. M. Dusmet y Alonso, OÖLM (paratype); Madrid, Rivas-Vaciamadrid, Canal de Manzanares to Camino de Uclés, 19.v.2021, 1♀, leg. T.J. Wood, TJWC [BOLD accession number WPATW192-21].
Andrena pruinosa Erichson, 1835: 104, ♀ [Spain:
Andrena lanuginosa Spinola, 1843: 137, ♀ [Spain, lectotype by present designation:
Remarks. Examination of the type material of both A. pruinosa and A. lanuginosa (Fig.
Distribution. Spain.
Material examined. Spain: Andalusia, 1♀, leg. Waltl,
Andrena succinea Dours, 1872: 424, ♀ [Algeria: type lost, neotype designated below, OÖLM].
Andrena chrysopyga Dours, 1872: 423, ♀ (nec. Andrena chrysopyga Schenck, 1853) [Algeria: type lost].
Andrena commixta Dalla Torre & Friese, 1895: 43. nom. nov. for Andrena chrysopyga Dours, 1872
Andrena sitifensis Pérez, 1895: 46, ♀ [Algeria:
Andrena fulvisquama Popov, 1940: 260, ♀ [Algeria:
Remarks. The synonymy of A. mayeti Pérez, 1895 (described from Tunisia) with A. succinea reported by
Distribution. Morocco, Algeria, Tunisia, Libya, Egypt, Israel and the West Bank, Jordan, Syria, Saudi Arabia, Iran (
Material examined. Algeria: Setif [36.2059°N, 5.3965°E], 1♀,
Andrena (Pruinosandrena) caroli Pérez, 1895: 47, ♀ [Algeria:
Andrena (Pruinosandrena) mayeti Pérez, 1895: 47, ♀ [Tunisia:
Distribution. Morocco, Algeria, Tunisia, Egypt, Israel.
Material examined. Algeria: Biskra [34.8600°N, 5.6995°E], 1♀,
This group of Palaearctic species was previously placed in the subgenus Thysandrena Lanham, 1949 by
The status of A. hypopolia (described from southern France) has been somewhat unclear, as no major morphological differences from A. numida (described from Algeria) are apparent. Warncke used A. hypopolia in combination with A. numida as the subspecies for south-western Europe, using several other taxa as subspecies for populations in Central and Eastern Europe (ssp. ? holosericea Bramson, 1879, considered a nomen dubium by
The situation is further complicated because the type of A. hypopolia is lost (and could not be found amongst undesignated Schmiedeknecht type material located in the
Genetically, barcoded specimens from Iberia showed almost no differentiation from specimens from Germany (average genetic distance 0.26%; Fig.
This action largely maintains the status quo of
The status of A. numida albiscopa is unclear, but based on its distribution and morphology (T3 is clearly punctured), it is transferred to A. hypopolia albiscopa comb. nov. The punctures of T3 are sparser than in A. hypopolia s. str., and the interspaces are shinier. This is true also of A. hypopolia material from Central Asia (Kyrgyzstan). For now, a conservative position is taken that A. hypopolia ranges from Iberia to Central Asia and western Siberia, though the eastern limit and the status of material from Turkey requires validation through genetic analysis across this range.
Finally, examination of the lectotype of A. setosa Pérez, 1903 (Fig.
Material examined. Andrena ranunculorum: France: Arreau [42.9064°N, 0.3557°E], 1♀,
Andrena numida f. syracusae: Italy: Campania, Is. Iscia, Panza, 9.iv.1991, 6♂, leg. J. Gusenleitner, OÖLM/TJWC.
Andrena fumida: Italy: Kampanien, Salerno, Monti Alburini (NP), SE Petina, 1100 m, 8.vi.2003, 1♀, leg. H. & R. Rausch, OÖLM; Mondello [Palermo, Sicily], 10.iv.1979, 1♂, leg. J.A.W. Lucas, OÖLM; Monte Faito (Campania), 13.v.1976, 1♂, leg. Pagliano,
Andrena blanda Pérez, 1895.
Blandandrena is monotypic, and hence diagnosis of A. blanda (Fig.
Andrena blanda males can be separated by their black clypeus (Fig.
Medium-sized bees (9–10 mm) with dark integument. Head broad, 1.4 times broader than long. Gena slightly exceeding width of compound eye; ocelloccipital distance 1.5–2 times diameter of lateral ocellus, slightly broader in male sex. Facial fovea broad, occupying almost entire distance between lateral ocellus and inner margin of compound eye. Mesosoma dorsally with moderately long light brown hairs, laterally with white hairs. Pronotum laterally with humeral angle. Dorsolateral surface of propodeum with obscure and finely raised rugosity; propodeal triangle broad, poorly delineated laterally, surface with fine granular reticulation, basally with weakly raised rugosity, propodeal triangle thus defined by change in surface sculpture. Forewing with nervulus antefurcal. Hind tibial spurs simple, not broadened basally or medially. Terga weakly and obscurely punctate, punctures separated by 1–2 puncture diameters. Male genital capsule rounded, more or less circular in outline, gonocoxae with inner margins apically diverging, produced into apically projecting short pointed teeth. Gonostyli apically broadened and flattened, apical disc slightly broader than long. Penis valves relatively narrow, occupying less than half space between gonostyli.
The name is taken from the name of the type species A. blanda, with blanda being the feminine singular of the adjective blandus which can mean pleasant, agreeable, smooth. It can be used to refer to the generally unremarkable nature of the species which has made it hard to assign to a particular group of species. The gender is feminine.
Andrena blanda (Spain, including mainland Spain and newly recorded for Fuerteventura), Morocco, Algeria, Tunisia;
Algeria: Biskra [34.8600°N, 5.6995°E], 1♀,
Andrena florea Fabricius, 1793.
Bryandrena is monotypic, and hence diagnosis of A. florea is de facto diagnosis of the subgenus. The combination of broad head (Fig.
Medium-sized bees (11–13 mm), integument predominantly dark, with red markings on at least one tergum, sometimes all terga extensively red marked. Head broad, 1.3–1.4 times wider than long, inner margins of compound eyes diverging ventrally. Gena slightly exceeding width of compound eye in females, clearly exceeding width of compound eye in males; ocelloccipital distance 1.5–2 times diameter of lateral ocellus. Facial fovea broad, occupying ¾ of distance between lateral ocellus and inner margin of compound eye. Pronotum laterally with humeral angle, more pronounced in male sex. Dorsolateral surface of propodeum microreticulate, with weakly raised reticulation; propodeal triangle poorly defined laterally, comparatively smooth and lacking microreticulation, basally with raised rugosity covering variable extent, never entire propodeal triangle. Forewing with nervulus interstitial. Terga regularly and densely punctate, punctures separated by 1 puncture diameter. Male genital capsule strongly elongate, gonocoxae essentially truncate with inner margin rounded, gonostyli apically produced, elongate, strongly flattened and spatulate, 3 times longer than broad; penis valves basally broad, strongly narrowing medially to become elongate and acutely pointed apically.
The name is taken from the pollen host plant Bryonia (Cucurbitaceae) which ultimately derives from the Greek βρυωνία [bruōnía]. Andrena florea can be found frequently almost wherever Bryonia species are in flower. The gender is feminine.
Andrena florea (West Palaearctic, from Morocco and Iberia to Iran and the Ural Mountains;
Andrena limbata Eversmann, 1852 (illustrated by
Historically, A. toelgiana Friese, 1921 has been considered the sister species to A. limbata, differing by the yellow clypeus in the female sex (see
Andrena (Limbandrena) limbata Eversmann, 1852, female A scutum, dorsal view B head, dorso-frontal view; Andrena limbata dusmeti Warncke, 1967 female C profile D terga, dorsal view; A. limbata s. str. female, Bulgarian specimen E profile F terga, dorsal view; A. limbata s. str. female, Turkish specimen G profile H terga, dorsal view.
Limbandrena (and, de facto, A. limbata) can be recognised in the female sex due to the combination of squamous brown hairs on the scutum, scutellum, and metanotum (Fig.
Males can be recognised by most of the same characters: the yellow clypeus (Fig.
Medium-sized bees (11–14 mm) with dark integument with exception of yellow maculations on female (sometimes) and male clypeus (always). Head 1.1–1.2 times broader than long, compound eyes with inner margins weakly converging apically. Gena slightly exceeding width of compound eye; ocelloccipital distance long, 3 times diameter of lateral ocellus. Facial fovea moderate, occupying ½ space between lateral ocellus and compound eye. Female scutum, scutellum, and metanotum covered with short brown squamous hairs. Pronotum laterally with weak humeral angle. Dorsolateral surface of propodeum weakly and shallowly but regularly punctate, punctures separated by 0.5–1 puncture diameter. Propodeal triangle clearly delineated laterally by raised carinae, internal surface with clear pattern of fine rugosity medially, not extending over entire area. Forewing with nervulus strongly postfurcal. Hind tibial spurs simple, not broadened basally or medially, apically weakly bent. Terga densely and finely punctate, punctures separated by 0.5 puncture diameters. Male genital capsule slightly elongate, with produced and weakly rounded gonocoxal teeth. Gonostyli with weakly raised and rounded projection on inner margin. Penis valves produced into rounded hyaline extensions laterally, occupying majority of space between gonostyli.
The name is taken from the name of the type species A. limbata, with limbata being the feminine singular of the adjective limbatus which means edged or fringed, probably in reference to the distinct squamous hairs on the female scutum and scutellum. The gender is feminine.
Andrena limbata (Europe from Portugal and Spain to Turkey, Israel, northern Iran, and the Ural Mountains;
Albania: Lopan [Lapanj], 14.vi.2018, 2♀, leg. Kobe Janssen collection (Belgium); Bulgaria: Lozenec [Lozenets, Лозенец]/Mičurin, 24.vi.1988, 5♀, leg. B. & O. Tkalců, OÖLM; Croatia: Istrien, Rovinjsko Selo, 8–9.vi.2012, 1♀, leg. Holzmann, OÖLM; France: B. d. R., Fontvieille, 28.v.1993, 4♀, leg. H. & J.E. Wiering,
Andrena oviventris Pérez, 1895.
Through the combination of slightly upturned fore margin of the clypeus, broad fovea occupying at least ½ the space between the lateral ocellus and the inner margin of the compound eye, weak but distinct humeral angle, unmodified posterior face of the hind femur (without teeth, carinae, or spines), simple hind tibial spur (not broadened basally or medially), dark integument, black male clypeus, and essential absence of defining features it falls very close to members of the relata-group and to Blandandrena that were formerly lumped together under the subgenus Poliandrena (see above).
In the female sex, Ovandrena species differ from these groups by only a single major character: the structure of the propodeal triangle (Fig.
Andrena (Ovandrena) subgen. nov. characters. Andrena (Ovandrena) marsae Schmiedeknecht, 1900, female A propodeal triangle; Andrena (Ovandrena) oviventris Pérez, 1895 B female propodeal triangle C female scutal hairs, profile view F male genital capsule; Andrena (Ovandrena) farinosa Pérez, 1895 D female scutal hairs, profile view E male genital capsule.
Males can be recognised through their combination of dark clypeus with upturned fore margin, distinctive propodeal triangle as in the female sex, pronotum with weak or strong humeral angle, A3 exceeding A4 but shorter than A4+5, and genital capsule which is compact with pronounced and rounded gonocoxal teeth (Fig.
Small to moderately sized bees (7–11 mm) typically with dark integument, one species with red tergal markings; male clypeus always dark. Head broad, 1.3–1.4 times broader than long, compound eyes with inner margins weakly converging apically. Gena slightly exceeding width of compound eye; ocelloccipital distance short, slightly less than to slightly more than diameter of lateral ocellus. Facial fovea moderately broad, occupying ½ distance between lateral ocellus and compound eye. Female scutum, scutellum, and metanotum covered with shortish light brown to whitish semi-squamous hairs (Fig.
The name is taken from the type species for the genus, A. oviventris. It derives from the Latin ovum meaning egg, in reference to the egg-shaped metasoma. The gender is feminine.
Andrena farinosa Pérez, 1895 (Spain and France), Andrena farinosoides Wood, 2020 (Morocco), Andrena marsae Schmiedeknecht, 1900 (Morocco, Algeria, Tunisia), and Andrena oviventris (Morocco, Algeria, Portugal, Spain, France). The subgenus is therefore currently restricted to the Western Mediterranean, and the centre of diversity is Morocco. The status of A. (incertae sedis) inusitata Pisanty, 2022 must be resolved through genetic analysis (see
Andrena oviventris: Algeria: Teniet el Had [35.8727°N, 2.0007°E], 1♀,
Andrena farinosa: Spain: Murcie [Murcia], 1♀,
1 | Females | 2 |
– | Males |
5 |
2 | Terga almost entirely red-marked, with at most slight black marks basally on T1 and two black spots laterally on T2 (north-western Africa) | marsae Schmiedeknecht |
– | Terga dark, without red markings | 3 |
3 | Scutum less densely punctate, punctures separated by at least 1 puncture diameter, surface clearly smooth and shiny between punctures (Morocco) | farinosoides Wood |
– | Scutum densely punctate, punctures separated by 0.5 puncture diameters to confluent, narrow interspaces shiny | 4 |
4 | Tergal discs glabrous, with hairs restricted to marginal areas. Larger, 10–11 mm (north-western Africa and south-western Europe) | oviventris Pérez |
– | Tergal discs extensively covered with extremely short hairs, forming a velvety pubescence in addition to denser and longer hairs on tergal margins. Smaller, 8–9 mm (Spain and France only) | farinosa Pérez |
5 | Larger, 9–10 mm. Tongue with outer surface of galea clearly punctate, punctures separated by 1–2 puncture diameters. Sterna with weak and sparse fringes on apical margins. Tergal punctation comparatively larger and coarser (north-western Africa and south-western Europe) | oviventris Pérez |
– | Smaller, 7–8 mm. Tongue with outer surface of galea more or less smooth and shiny, without obvious punctures. Sterna with strong and dense fringes on apical margins. Tergal punctation comparatively fine | 6 |
6 | Clypeus comparatively less densely punctate, punctures separated by 0.5–1 puncture diameters, with shiny interspaces, thus appearing shiny. Pronotum with humeral angle comparatively strong (Spain and France only) | farinosa Pérez |
– | Clypeus comparatively more densely punctate, punctures separated by 0.5 puncture diameters, interspaces dull, thus appearing dull (north-western Africa) | 7 |
7 | Viewed laterally and ventrally, tergal margins, ventrolateral parts of terga, and sternal margins usually lightened reddish orange-brown (north-western Africa) |
marsae |
– | Tergal and sternal margins dark to hyaline, never extensively lightened reddish orange-brown (Morocco only) |
farinosoides |
Andrena pruinosa Erichson, 1835.
This group of species was formerly placed in the subgenus Campylogaster due to the unusual character of the strongly and densely punctate mesepisternum and dorsolateral parts of the propodeum (punctures separated by <0.5 puncture diameters). However, as discussed above, Campylogaster sensu Warncke is polyphyletic and can be broken into three distinct clades (Campylogaster s. str., Pruinosandrena, and the incisa-group). All three share the distinctly punctate mesepisternum and also an extremely long ocelloccipital distance exceeding three times the diameter of the lateral ocellus. However, separation is straightforward. The true Campylogaster have the marginal area of the terga clearly and distinctly impressed with the apical margin reflexed; the impression therefore forms a latitudinal depressed furrow. In Pruinosandrena, the marginal areas of the terga are flat, without any kind of depression. In the incisa-group, the tergal margins are flat and the mesepisternum is densely punctate, but the dorsolateral parts of the propodeum have only raised reticulation, without punctures. The propodeal triangle is also clearly delineated by raised lateral carinae, whereas these are absent in Pruinosandrena. The combination of extremely long ocelloccipital distance, dense punctures on the mesepisternum and dorsolateral parts of the propodeum, and flat tergal marginal areas is therefore unique and characterises Pruinosandrena.
Medium-sized bees (10–14 mm). Integument variable, from dark with at most tergal margins lightened hyaline-yellow to entirety of metasoma and legs red-marked; male clypeus yellow-marked in one species. Head moderately broad, 1.2 times broader than long. Gena broad, exceeding width of compound eye; ocelloccipital distance extremely long, at least 3 times diameter of lateral ocellus. Facial fovea variable, from narrow to occupying entirety of distance between lateral ocellus and inner margin of compound eye. Female scutum, scutellum, and metanotum with pubescence variable, in some species with extremely short squamous hairs, hairs longer and non-squamous in other species. Pronotum laterally with humeral angle. Mesepisternum and dorsolateral parts of propodeum densely and clearly punctate, punctures confluent to separated by <0.5 puncture diameters. Propodeal triangle without lateral carinae, internal surface with dense network of irregularly raised rugosity, thus contrasting punctate dorsolateral surface. Forewing with nervulus interstitial. Hind tibial spurs simple, not broadened basally or medially. Terga typically densely and finely punctate, punctures separated by 1 puncture diameter, at least on T2–5. Male genital capsule simple, compact, with gonocoxae apically truncate to produced into weak rounded teeth. Gonostyli apically spatulate, penis valves more or less narrow, parallel-sided, occupying less than ½ space between gonostyli.
The name is taken from the type species for the subgenus, A. pruinosa. The Latin word pruinosa is the feminine singular of pruinosus which means ‘frosty’, in reference to the squamous hairs of the mesosoma. The gender is feminine.
Andrena caroli (Morocco to Israel); Andrena nilotica Warncke, 1967 (Spain); Andrena parata (Spain); Andrena pruinosa (Spain); Andrena sparsipunctata Wood, 2020 (Morocco); Andrena succinea (Morocco to Iran and Saudi Arabia). The centre of diversity is therefore Spain + Morocco, with all six species occurring here.
Note, the male of A. sparsipunctata is unknown. The males described by
1 | Females | 2 |
– | Males | 7 |
2 | In dorsal view, fovea broad, clearly occupying more than half of the distance between the lateral ocellus and the compound eye (Fig. |
caroli Pérez |
– | Fovea narrow, occupying at most half the distance between the lateral ocellus and the compound eye (Figs |
3 |
3 | T1 with extremely sparse punctures, punctures separated by 2–4 puncture diameters. Facial fovea along its entire length separated from the inner margin of the compound eye by a distance equal to its own diameter (south-western Morocco only) | sparsipunctata Wood |
– | T1 with dense punctures, punctures separated at most by 2 puncture diameters, usually by 1 puncture diameter (Fig. |
4 |
4 | Pubescence of scutum weakly squamous, anterior dorsolateral corners of scutum with pubescence longer, clearly exceeding width of antennae in length. Terga always predominantly red. Terga with clear apical hair bands. Punctation of T1 slightly spaced, punctures separated by 1–2 puncture diameters (eastern and south-eastern Spain only) | nilotica Warncke |
– | Pubescence of scutum strongly squamous and short, anterior dorsolateral corners of scutum with pubescence short, clearly shorter than width of antennae (Figs |
5 |
5 | A3 exceeding A4+5 in length. Terga always extensively red-marked (central and south-eastern Spain) | parata Warncke |
– | A3 equalling A4+5 in length. Terga variable, from almost entirely black to entirely red-marked | 6 |
6 | Terga usually predominantly dark (Figs |
pruinosa Erichson |
– | Terga always extensively or entirely red-marked (Figs |
succinea Dours |
7 | Clypeus at least partly yellow-marked (North Africa to the Middle East) | succinea Dours |
– | Clypeus uniformly dark | 8 |
8 | A3 very short, shorter than A4 (North Africa to Israel) | caroli Pérez |
– | A3 at least slightly longer than A4 | 9 |
9 | A3 exceeding A4+5 in length (central and south-eastern Spain) | parata Warncke |
– | A3 not exceeding A4+5 | 10 |
10 | Genital capsule without clear kink in the inner margins of the gonostyli (Fig. |
nilotica Warncke |
– | Genital capsule with clear kink in the inner margins of the gonostyli (Fig. |
pruinosa Erichson |
Holotype. Spain: Cádiz, Parque Natural Los Alcornocales, Las Algamitas, Finca Murtas, 36.3273°N, -5.5986°W, 18.iii.2023, 1♀, leg. T.J. Wood, OÖLM.
Paratypes. Spain: Cádiz, Parque Natural Los Alcornocales, Las Algamitas, Finca Murtas, 18.iii.2023, 1♀, leg. T.J. Wood, TJWC; Cádiz, Tarifa, 1 km N, grazing fields, 23.iii.2023, 1♀, leg. T.J. Wood, OÖLM.
Female. Body length: 9 mm (Fig.
Male. Unknown.
Andrena juliae can be recognised as belonging to the subgenus Avandrena due to its moderate to small body size (9 mm), short and wide head (clearly wider than long), and short and wide foveae that are only slightly longer than wide, as well as the behavioural observation that it is strongly associated with Erodium (Geranicaceae, see Remarks). The posterior face of the hind femora lacks spines, separating the species from A. avara Warncke, 1967 and A. panurgina De Steffani, 1889 and placing it close to A. melacana Warncke, 1967 and A. erodiorum Wood & Ortiz-Sánchez, 2022.
Andrena juliae can be separated from both species by the structure of the propodeum which has the dorsolateral parts with fine granular shagreenation that is overlain by a fine network of raised rugosity, this rugosity not forming a linked network. The propodeal triangle itself is clearly differentiated, slightly depressed below the level of the surrounding parts of the propodeum and delineated by fine carinae, the surface with fine granular shagreenation and with a network of raised longitudinal carinae covering the basal 2/3rds. In contrast, A. erodiorum has the dorsolateral parts of the propodeum shiny, overlain with a fine network of raised rugosity that joins together to form a clear network (Fig.
The propodeum of A. juliae therefore sits between both comparison species and is distinct from both. It can be further separated by the pubescence of the mesepisternum which is predominantly composed of pale hairs, with approximately 30% of these hairs black (in A. melacana with 50–60% of the hairs of the mesepisternum black; in A. erodiorum with only 10% of these hairs black), by the facial foveae which occupy half of the space between the compound eye and a lateral ocellus (occupying ¾ of this space in A. erodiorum), by the colour of the hairs of the apical fringe of T5 and those flanking the pygidial plate which are dark brown (golden-brown in A. erodiorum, dark brown in A. melacana), and by the pubescence of the terga which are covered in sparse erect white hairs, T2–4 with dense apical hair bands of white hairs that obscure the underlying surface (in A. melacana terga with sparse short pubescence, only forming weak apical tergal hair bands that do not obscure the underlying surface; pubescence very similar in A. erodiorum).
All specimens were collected from Erodium spp. The two females from Las Algamitas were collecting pollen from this genus (Erodium pollen can be seen in Fig.
Dedicated to my friend and colleague Julia Jones (University College Dublin, Ireland) who invited me on the University field course during which this new species was discovered.
Spain (Cádiz province).
Andrena avara s. str.: Spain: Cádiz, Parque Natural Los Alcornocales, Las Algamitas, Finca Murtas, 18.iii.2023, 1♀, leg. T.J. Wood, TJWC; Cádiz, Bolonia, El Lentiscal, 24.iii.2023, 1♀, leg. T.J. Wood, TJWC.
Andrena melacana: Spain: Cádiz, Parque Natural Los Alcornocales, Las Algamitas, Finca Murtas, 18.iii.2023, 3♂, 2♀, leg. T.J. Wood, TJWC; Cádiz, Tarifa, 1 km N, grazing fields, 19.iii.2023, 4♂, 6♀, leg. T.J. Wood, TJWC.
Andrena panurgina: Spain: Cádiz, Parque Natural Los Alcornocales, Las Algamitas, Finca Murtas, 18.iii.2023, 4♂, 3♀, leg. T.J. Wood, TJWC; Cádiz, Tarifa, 1 km N, grazing fields, 19–23.iii.2023, 6♂, 2♀, leg. T.J. Wood, TJWC.
Holotype. Spain: Granada, Sierra Nevada, Trevélez to Refugio La Campiñuela, 37.0239°N, -3.2656°W, 1700–2400 m, 14.vi.2021, 1♀, leg. T.J. Wood, OÖLM [BOLD accession number WPATW368-21].
Female. Body length: 9.5 mm (Fig.
Male. Unknown.
Andrena isolata can be quickly recognised as a Euandrena due to the narrow facial foveae (dorsally occupying ⅓ of space between the lateral ocellus and the compound eye) which narrow further ventrally combined with the long A3 (slightly exceeding length of A4+5) and the simple, non-plumose hairs of the tibial scopae. Its generally dark appearance with orange-brown hairs dorsally on the mesosoma and tibial scopa plus at least some black hairs on the mesepisternum place it immediately close to A. bicolor and allied taxa. As discussed above, the subgenus Euandrena is taxonomically complex, and multiple genetically distinct taxa have been lumped under A. bicolor. In an Iberian context, A. isolata is best diagnosed against A. bicolor s.l., A. fortipunctata Wood, 2021, and the distinct and probably undescribed taxon in north-western Africa identified above. Extreme care should be taken when identifying specimens morphologically, and barcodes should be used whenever possible.
In direct comparison to barcoded A. bicolor s.l. specimens, the only consistent character that can be identified is the structure of the clypeus. Andrena isolata has the clypeus densely punctate, with punctures separated by <0.5 puncture diameters, interspaces shiny but overall the clypeus only weakly shiny due to the small size of the interspaces (Fig.
Andrena isolata probably represents a relictual species that has become isolated on the Sierra Nevada from what is now a remaining North African population. Additional genetic sampling is needed to establish whether it is found away from the Sierra Nevada, but a specimen from the nearby Sierra de Baza collected at an altitude of 2000 m barcoded as A. bicolor s.l. [WPATW297-21]. Andrena isolata may well be restricted to the Sierra Nevada. Additional sampling is required to establish its ecology, including its voltinism. Its capture on Campanula implies that it has a similar ecology to A. bicolor s.l. (see
Derived from the Latin insulatus, to be made into an island, isolata (feminine form) thus means to be isolated, in reference to its presence on the Sierra Nevada, separated from its nearest genetic relative in North Africa.
Spain (Sierra Nevada).
Holotype. Spain: Granada, Sierra Nevada, Mirador Monte Ahí de Cara, 37.1239°N, -3.4322°W, 2100 m, 6.vi.2021, 1♀, leg. T.J. Wood, on Vella spinosa (Brassicaceae), OÖLM [BOLD accession number WPATW972-22].
Paratypes. Spain: Granada, Sierra Nevada, Mirador Monte Ahí de Cara, 2100 m, 6.vi.2021, 2♀, leg. T.J. Wood, on Vella spinosa (Brassicaceae), OÖLM/TJWC; Granada, Sierra Nevada, Puerto de la Ragua, Barranco Maja Caco, 2000 m, 10.vi.2021, 1♀, leg. T.J. Wood, TJWC; Granada, Sierra Nevada 1900 m, ri. Veleta, 1.vi.1982, 1♀, leg. R. Leys,
Female. Body length: 7 mm (Fig.
Male. Unknown.
Andrena ortizi can quickly be recognised as a Micrandrena due to its small body size, dark integument, and entirely rugose propodeal triangle. Due to the comparatively (for a Micrandrena) long face and clypeus (head overall only 1.1 times wider than broad; apical margin of clypeus clearly ventrally exceeding level of a line drawn between the lower margins of the compound eyes), narrow facial fovea (dorsally occupying ¼ of space between the lateral ocellus and the compound eye, consistently wide along its length, and densely punctate scutum (punctures separated by <1 puncture diameter) they are comparable to A. rugulosa Stöckhert, 1935 (Switzerland to Lebanon and the Caucasus), A. atlantea Wood, 2021 (High and Middle Atlas Mountains in Morocco), and an undescribed Micrandrena species from the Middle Atlas (see above).
Andrena ortizi can be separated from A. rugulosa by the scutum which is slightly less densely punctate, punctures separated by 1 puncture diameter (punctures separated by 0.5 puncture diameters in A. rugulosa), the underlying surface of the scutum being finely shagreened and shiny (scutum is densely shagreened and dull in A. rugulosa), the tergal discs are strongly and clearly punctate (tergal discs obscurely punctate in A. rugulosa), and the marginal areas of T2–4 occupy at least ½ the visible length of the tergum, on T3–4 clearly occupying over ½ this length (marginal areas typically occupying ⅓ length of tergum, at most occupying ½ tergum on T4 in A. rugulosa). Andrena ortizi is more similar to A. atlantea, sharing a similarly less densely punctate and weakly shiny scutum, but the same character of wide tergal margins can be used to separate them, with the marginal areas occupying at most ⅓ of the length of the tergum in A. atlantea. Finally, A. ortizi is most similar to the undescribed Micrandrena from the Middle Atlas, and the width of the tergal margins can again be used to separate them, with the tergal margins occupying at most ⅓ of the disc of T2 and ½ of the discs of T3–4. Additionally, A. ortizi has the disc of T2–3 clearly and densely punctate, whereas in the undescribed Micrandrena the discs of T2–3 are at most obscurely punctate, with punctures disappearing into the background microreticulation. Andrena ortizi also has a strongly isolated distribution, separated from the Swiss Alps (A. rugulosa) by c. 1,300 km and the high altitude parts around Ifrane and Azrou in the Middle Atlas (A. atlantea, the undescribed Micrandrena species) by c. 400 km.
At the Mirador Monte Ahí de Cara (Fig.
Dedicated to the Spanish naturalist and hymenopterist Francisco Javier Ortiz-Sánchez who has worked extensively on the Iberian bee fauna for many years, including that of the Sierra Nevada.
Spain (Sierra Nevada).
Holotype. Spain: Málaga, PN Sierra de las Nieves, mountain peak S of Pinsapo Escalereta, 36.6621°N, -5.0362°W, 1600 m, 30.v.2021, 1♀, leg. T.J. Wood, OÖLM [BOLD accession number WPATW239-21].
Paratypes. Spain: Málaga, PN Sierra de las Nieves, mountain peak S of Pinsapo Escalereta, 1600 m, 30.v.2021, 1♀, leg. G. Ghisbain, TJWC; Málaga – Elvira, 11.ii.1981, 4♂, leg. H. Teunissen,
Female. Body length: 15–16 mm (Fig.
Male. Body length: 13–14 mm (Fig.
Andrena ghisbaini can be recognised within Truncandrena due to its characteristically smooth and finely granulate propodeal triangle which contrasts with the similarly granulate dorsolateral parts of the propodeum which bear fine and scattered raised hair-bearing punctures, the rounded pronotum, the linear malar space, the large body size (>13 mm), yellow male clypeus, and typical genital capsule with the inner margins of the flattened apical parts of the gonostyli strongly raised. It can be placed closest to A. villipes Pérez, 1895 (Fig.
The immediate difference between the two taxa is size, with A. villipes averaging 12–13 mm in length in females and 11–12 mm in males, compared to 15–16 mm and 13–14 mm respectively in A. ghisbaini. Structurally, A. ghisbaini females can be separated by the bicoloured scopa, black dorsally and orange ventrally (Fig.
In the male sex, A. ghisbaini can be separated by the same clypeal punctation character (stronger in A. ghisbaini with clearer impunctate midline), but this is slightly more subtle than in the female sex. Direct comparison of the genital capsule shows that flattened apical part of the gonostyli are more strongly elongate and longer than broad, thus appearing triangular (Fig.
The two females from the Sierra de las Nieves (Fig.
Host plant use and dietary classification for selected Iberian Andrena species. n, total number of pollen loads; N, number of pollen loads from different localities. Plant taxa: ADO, Adoxaceae; AMA, Amaryllidaceae; API, Apiaceae; ASP, Asparagaceae; AST, Asteraceae; BOR, Boraginaceae; BRA, Brassicaceae; CAM, Campanulaceae; CAP, Caprifoliaceae; CAR, Caryophyllaceae; CIS, Cistaceae; CRA, Crassulaceae; EUP, Euphorbiaceae; FAB, Fabaceae; FAG, Fagaceae; FRA, Frankeniaceae; GER, Geraniaceae; HYP, Hypericaceae; PAP, Papaveraceae; PLA, Plantaginaceae; PLU, Plumbaginaceae; RES, Resedaceae; RHA, Rhamnaceae; ROS, Rosaceae; SAL, Salicaceae; SAP, Sapindaceae; SCR, Scrophulariaceae. Countries: BE, Belgium; BG, Bulgaria; DZ, Algeria; ESP, Spain; FRA, France; IL, Israel; IR, Iran; MA, Morocco; PT, Portugal; SY, Syria; TJ, Tajikistan; TN, Tunisia.
Species | n | N | Origin (and number) of pollen loads | Result of microscopic analysis of pollen grains (% of pollen grains) | Percentage of pure loads of preferred host | Percentage of loads with preferred host | Host range |
---|---|---|---|---|---|---|---|
Aciandrena Warncke | |||||||
A. fulica Warncke | 12 | 7 | ESP (10), PT (2) | BRA 99.6, CIS 0.4 | 91.7 | 100.0 | Broadly oligolectic (Brassicaceae) |
A. vacella Warncke | 2 | 2 | ESP (2) | BRA 100.0 | 100.0 | 100.0 | Broadly oligolectic (Brassicaceae) |
aegyptiaca-group | |||||||
A. alluaudi Benoist | 4 | 3 | MA (2), PT (2) | AST 100.0 | 100.0 | 100.0 | Broadly oligolectic (Asteraceae; Cichorioideae) |
Aenandrena Warncke | |||||||
A. aeneiventris Morawitz | 15 | 7 | ESP (15) | API 100.0 | 100.0 | 100.0 | Possibly broadly oligolectic (Apiaceae) |
A. hedikae Jäger | 22 | 10 | ESP (11), MA (9), PT (2) | API 100.0 | 100.0 | 100.0 | Possibly broadly oligolectic (Apiaceae) |
A. hystrix Schmiedeknecht | 9 | 5 | ESP (8), PT (1) | BRA 100.0 | 100.0 | 100.0 | Broadly oligolectic (Brassicaceae) |
Avandrena Warncke | |||||||
A. avara Warncke | 2 | 2 | ESP (2) | GER 100.0 | 100.0 | 100.0 | Broadly oligolectic (Geraniaceae) |
A. melacana Warncke | 6 | 2 | ESP (6) | GER 100.0 | 100.0 | 100.0 | Broadly oligolectic (Geraniaceae) |
A. panurgina De Steffani | 11 | 5 | ESP (4), FRA (4), PT (3) | GER 93.6, AST 5.0, BRA 1.4 | 81.8 | 100.0 | Broadly oligolectic (Geraniaceae) |
Blandandrena subgen. nov. | |||||||
A. blanda Pérez | 27 | 9 | ESP (8), MA (19) | RES 100.0 | 100.0 | 100.0 | Narrowly oligolectic (Reseda, Resedaceae) |
Brachyandrena Pittioni | |||||||
A. colletiformis Morawitz | 6 | 4 | ESP (5), PT (1) | API 100.0 | 100.0 | 100.0 | Possibly broadly oligolectic (Apiaceae) |
A. miegiella Dours | 5 | 4 | ESP (2), MA (2), TN (1) | API 99.8, AST 0.2 | 80.0 | 100.0 | Possibly broadly oligolectic (Apiaceae) |
Chlorandrena Pérez | |||||||
A. abrupta Warncke | 1 | 1 | PT (1) | AST 100.0 | 100.0 | 100.0 | Probably broadly oligolectic (Asteraceae; Asteroideae) |
A. cinerea Brullé | 22 | 15 | ESP (6), FRA (3), PT (12), TN (1) | AST 100.0 | 100.0 | 100.0 | Broadly oligolectic (Asteraceae; Cichorioideae) |
A. curtivalvis Morice | 1 | 1 | ESP (1) | AST 100.0 | 100.0 | 100.0 | Broadly oligolectic (Asteraceae; Cichorioideae) |
A. elata Warncke | 13 | 5 | ESP (13) | AST 100.0 | 100.0 | 100.0 | Broadly oligolectic (Asteraceae; Asteroideae) |
A. leucolippa Pérez | 22 | 10 | ESP (12), FRA (10) | AST 100.0 | 100.0 | 100.0 | Broadly oligolectic (Asteraceae; Asteroideae) |
A. rhenana Stöckhert | 8 | 4 | ESP (2), PT (6) | AST 100.0 | 100.0 | 100.0 | Broadly oligolectic (Asteraceae; Cichorioideae) |
A. senecionis Pérez | 21 | 15 | ESP (11), FRA (3), MA (3), PT (4) | AST 100.0 | 100.0 | 100.0 | Broadly oligolectic (Asteraceae; Cichorioideae) |
Chrysandrena Hedicke | |||||||
A. fertoni Pérez | 4 | 3 | ESP (4) | AST 100.0 | 100.0 | 100.0 | Broadly oligolectic (Asteraceae; Cichorioideae) |
Cordandrena Warncke | |||||||
A. vaulogeri Pérez | 10 | 5 | ESP (3), MA (7) | BRA 61.8, ROS 22.6, AST 9.6, FAB 6.0 | 30.0 | 80.0 | Polylectic s. str. |
Cryptandrena Pittioni | |||||||
A. ventricosa Dours | 39 | 10 | ESP (29), FRA (10) | FAB 92.8, API 6.1, others 1.1 | 76.9 | 97.4 | Polylectic with a strong preference (Fabaceae) |
Didonia Gribodo | |||||||
A. mucida Kriechbaumer (1st generation) | 3 | 3 | ESP (1), PT (2) | ASP 100.0 | 100.0 | 100.0 | Possibly narrowly oligolectic (Muscari; Asparagaceae) |
A. mucida Kriechbaumer (2nd generation) | 12 | 8 | BG (2), ESP (9), MA (1) | CAP 100.0 | 100.0 | 100.0 | Broadly oligolectic (Caprifoliaceae) |
Euandrena Pérez | |||||||
A. lavandulae Pérez | 5 | 5 | ESP (3), FRA (1), PT (1) | FAB 27.6, CIS 24.3, SCR 20.7, PLA 10.3, AST 8.3, CAM 4.7, GER 2.9, CAR 1.2 | 20.0 | 20.0 | Polylectic s. str. |
Graecandrena Warncke | |||||||
A. nebularia Warncke | 5 | 3 | ESP (1), MA (4) | BRA 100.0 | 100.0 | 100.0 | Probably broadly oligolectic (Brassicaceae) |
A. verticalis Pérez | 30 | 21 | ESP (20), MA (8), PT (2) | BRA 56.3, API 43.8 | 53.3 | 53.3 | Mesolectic (Apiaceae & Brassicaceae) |
incisa-group | |||||||
A. lateralis Morawitz | 7 | 3 | ESP (3), IR (1), TJ (3) | API 100.0 | 100.0 | 100.0 | Broadly oligolectic (Apiaceae) |
Leucandrena Hedicke | |||||||
A. leptopyga Pérez | 19 | 12 | DZ (1), ESP (1), MA (12), PT (5) | RES 90.8, BRA 6.6, BOR 1.8, SCR 0.7 | 78.9 | 94.7 | Polylectic with a strong preference (Reseda, Resedaceae) |
A. tunetana Schmiedeknecht | 4 | 4 | DZ (1), ESP (2), MA (1) | BRA 100.0 | 100.0 | 100.0 | Probably broadly oligolectic (Brassicaceae) |
Melanapis Cameron | |||||||
A. fuscosa Erichson | 18 | 12 | ESP (15), FRA (1), IL (1), PT (1) | BRA 52.4, API 15.5, PAP 12.7, AST 9.5, ROS 6.1, EUP 3.1, others 0.7 | 50.0 | 72.2 | Polylectic s. str. |
Melandrena Pérez | |||||||
A. albopunctata (Rossi) | 15 | 7 | ESP (14), MA (1) | AST 47.4, API 25.3, CAP 11.2, BRA 6.4, FAB 3.9, PAP 3.1, others 2.7 | 20.0 | 86.7 | Polylectic s. str. |
A. assimilis Radoszkowski | 15 | 6 | ESP (4), FRA (11) | AST 34.8, ROS 21.1, API 21.0, PLU 7.9, AMA 6.6, SAL 2.7, others 6.1 | 6.7 | 73.3 | Polylectic s. str. |
A. bicolorata (Rossi) | 8 | 5 | ESP (2), PT (6) | BRA 100.0 | 100.0 | 100.0 | Broadly oligolectic (Brassicaceae) |
A. florentina Magretti | 9 | 5 | MA (1), PT (8) | BRA 100.0 | 100.0 | 100.0 | Broadly oligolectic (Brassicaceae) |
A. morio Brullé (including A. hispania Warncke) | 9 | 8 | ESP (3), PT (6) | CIS 69.6, API 17.2, AST 10.0, others 3.2 | 22.2 | 77.8 | Polylectic s. str. |
Micrandrena Ashmead | |||||||
A. ampla Warncke | 21 | 10 | ESP (10), FRA (10), PT (1) | API 100.0 | 100.0 | 100.0 | Broadly oligolectic (Apiaceae) |
A. bayona Warncke | 2 | 2 | ESP (2) | API 50.0, BRA 50.0 | 50.0 | 50.0 | Probably polylectic |
A. djelfensis Pérez | 20 | 13 | ESP (3), MA (7), PT (10) | CIS 99.9, FAB 0.1 | 95.0 | 100.0 | Broadly oligolectic (Cistaceae) |
A. fabrella Pérez | 22 | 13 | ESP (8), FRA (1), MA (5), PT (8) | CIST 99.9, AST 0.1 | 90.0 | 100.0 | Broadly oligolectic (Cistaceae) |
A. icterina Warncke | 9 | 6 | ESP (9) | BRA 69.1, CIS 10.6, SAL 10.3, EUP 5.8, others 4.2 | 22.2 | 88.9 | Polylectic s. str. |
A. longibarbis Pérez | 13 | 8 | ESP (2), MA (8), PT (3) | BRA 99.6, AST 0.4 | 92.3 | 100.0 | Broadly oligolectic (Brassicaceae) |
A. nana (Kirby) | 51 | 28 | ESP (33), FRA (1), MA (8), PT (9) | API 71.2, BRA 28.6, EUP 0.2 | 68.6 | 74.5 | Polylectic with a strong preference (Apiaceae) |
A. nitidula Pérez | 39 | 18 | ESP (16), MA (21), PT (2) | BRA 100.0 | 100.0 | 100.0 | Broadly oligolectic (Brassicaceae) |
A. omnilaevis Wood | 6 | 5 | ESP (2), PT (4) | CRA 100.0 | 100.0 | 100.0 | Probably narrowly oligolectic (Sedum, Crassulaceae) |
A. orana Warncke | 17 | 3 | DZ (9), MA (5), PT (3) | BRA 100.0 | 100.0 | 100.0 | Broadly oligolectic (Brassicaceae) |
A. pauxilla Stöckhert | 11 | 5 | ESP (11) | CRA 82.1, BRA 17.9 | 81.8 | 81.8 | Possibly polylectic with a strong preference (Sedum, Crassulaceae) |
A. spreta Pérez | 19 | 11 | ESP (15), MA (3), PT (1) | BRA 93.1, AST 3.0, EUP 2.8, FAB 1.2 | 78.9 | 100.0 | Polylectic with a strong preference (Brassicaceae) |
A. tenuistriata Pérez | 39 | 27 | ESP (17), FRA (3), MA (6), PT (13) | BRA 99.8, others 0.2 | 94.9 | 100.0 | Broadly oligolectic (Brassicaceae) |
Nobandrena Warncke | |||||||
A. funerea Warncke | 12 | 6 | ESP (12) | BRA 100.0 | 100.0 | 100.0 | Broadly oligolectic (Brassicaceae) |
Notandrena Pérez | |||||||
A. aerinifrons Dours | 25 | 8 | DZ (3), ESP (3), MA (11), PT (8) | BRA 100.0 | 100.0 | 100.0 | Broadly oligolectic (Brassicaceae) |
A. bellidis Pérez | 3 | 3 | ESP (2), PT (1) | AST 38.9, RES 36.4, RAN 18.2, BOR 6.5 | 0.0 | 66.7 | Polylectic s. str. |
A. juliana Wood | 35 | 2 | ESP (35) | API 82.6, FRA 13.1, CIS 2.8, others 1.4 | 74.3 | 82.9 | Polylectic with a strong preference (Apiaceae) |
A. leucophaea Lepeletier | 2 | 2 | ESP (2) | AST 100.0 | 100.0 | 100.0 | Possibly oligolectic (Asteraceae; Asteroideae) |
A. nigroviridula Dours | 9 | 8 | ESP (4), MA (4), PT (1) | BRA 100.0 | 100.0 | 100.0 | Broadly oligolectic (Brassicaceae) |
A. varuga Warncke | 3 | 2 | ESP (3) | BRA 100.0 | 100.0 | 100.0 | Probably broadly oligolectic (Brassicaceae) |
numida-group | |||||||
A. hypopolia Schmiedeknecht | 8 | 7 | ESP (4), FRA (1), PT (3) | BRA 64.8, API 34.6, AST 0.6 | 50.0 | 62.5 | Mesolectic (Apiaceae & Brassicaceae) |
A. ranunculorum Morawitz | 17 | 17 | FRA (17) | BRA 75.8, API 10.2, ROS 5.7, FAG 5.6, AST 2.1, ADO 0.5 | 64.7 | 100.0 | Polylectic with a strong preference (Brassicaceae) |
Orandrena Warncke | |||||||
A. monilia Warncke | 2 | 2 | ESP (1), MA (1) | BRA 100.0 | 100.0 | 100.0 | Probably broadly oligolectic (Brassicaceae) |
Ovandrena subgen. nov. | |||||||
A. farinosa Pérez | 9 | 5 | ESP (9) | FAB 100.0 | 100.0 | 100.0 | Broadly oligolectic (Fabaceae) |
A. oviventris Pérez | 28 | 11 | ESP (9), FRA (4), MA (12), PT (3) | RES 98.3, others 1.7 | 82.1 | 100.0 | Narrowly oligolectic (Reseda, Resedaceae) |
Plastandrena Hedicke | |||||||
A. asperrima Pérez | 56 | 29 | ESP (8), FRA (5), MA (43) | BRA 77.6, RES 18.6, ROS 2.5, AST 1.3 | 69.6 | 87.5 | Polylectic with a strong preference (Brassicaceae) |
A. pilipes Fabricius s. str. | 28 | 21 | ESP (15), FRA (8), PT (5) | BRA 54.4, ROS 18.9, AST 13.8, CIS 6.6, API 5.7, others 0.7 | 35.7 | 57.1 | Polylectic s. str. |
relata-group | |||||||
A. corax Warncke | 10 | 4 | ESP (8), PT (2) | RES 99.7, AST 0.3 | 90.0 | 100.0 | Narrowly oligolectic (Reseda, Resedaceae) |
A. laurivora Warncke | 3 | 1 | MA (1) | RES 100.0 | 100.0 | 100.0 | Probably narrowly oligolectic (Reseda, Resedaceae) |
A. relata Warncke | 2 | 2 | ESP (2) | RES 100.0 | 100.0 | 100.0 | Probably narrowly oligolectic (Reseda, Resedaceae) |
Rufandrena Warncke | |||||||
A. orbitalis Morawitz | 9 | 6 | ESP (2), FRA (4), PT (3) | PLA 100.0 | 100.0 | 100.0 | Narrowly oligolectic (Plantago, Plantaginaceae) |
A. rufiventris Lepeletier | 3 | 1 | MA (3) | PLA 100.0 | 100.0 | 100.0 | Narrowly oligolectic (Plantago, Plantaginaceae) |
Simandrena Pérez | |||||||
A. antigana Pérez | 25 | 14 | ESP (6), MA (7), PT (12) | BRA 99.8, others 0.2 | 96.0 | 100.0 | Broadly oligolectic (Brassicaceae) |
A. cilissaeformis Pérez | 5 | 5 | ESP (2), MA (3) | BRA 83.6, EUP 8.8, RHA 7.6 | 60.0 | 80.0 | Probably polylectic with a strong preference (Brassicaceae) |
A. propinqua Schenck | 43 | 31 | BE (4), ESP (21), FRA (5), MA (3), PT (10) | BRA 46.1, ROS 25.8, FAB 10.2, CIS 4.4, CRA 3.2, BOR 2.5, others 7.7 | 30.2 | 58.1 | Polylectic s. str. |
A. rhypara Pérez | 4 | 3 | MA (4) | RES 100.0 | 100.0 | 100.0 | Possibly narrowly oligolectic (Reseda; Resedaceae) |
A. vetula Lepeletier | 31 | 16 | ESP (20), FRA (2), MA (7), SY (1), TN (1) | BRA 99.8, others 0.2 | 93.5 | 100.0 | Broadly oligolectic (Brassicaceae) |
Truncandrena Warncke | |||||||
A. doursana Dufour | 8 | 3 | MA (7), PT (1) | BRA 100.0 | 100.0 | 100.0 | Broadly oligolectic (Brassicaceae) |
A. ferrugineicrus Dours | 28 | 18 | DZ (1), ESP (16), MA (2), PT (9) | BRA 100.0 | 100.0 | 100.0 | Broadly oligolectic (Brassicaceae) |
A. nigropilosa Warncke | 23 | 8 | ESP (16), FRA (5), MA (2) | BRA 100.0 | 100.0 | 100.0 | Broadly oligolectic (Brassicaceae) |
A. villipes Pérez | 6 | 2 | FRA (1), PT (5) | CIS 100.0 | 100.0 | 100.0 | Probably broadly oligolectic (Cistaceae) |
Dedicated to my friend and colleague Guillaume Ghisbain (Mons, Belgium) who accompanied me during fieldwork in Málaga province, and who is an accomplished hymenopterist in his own right.
Spain (Málaga province).
Description. Male. Body length 6.5–7 mm (Fig.
Diagnosis. Andrena alma can be recognised due to its combination of small body size, dark integument, pronotum with humeral angle, evenly shagreened and weakly shiny terga, gena exceeding the width of the compound eye (Figs
Distribution. Central and southern Portugal and Spain.
Material examined. Portugal: Algarve, Monte Gordo, Retur, Praia do Cabeço, 29.iii.2022, 1♂, leg. T.J. Wood, TJWC; Algarve, Tavira, Cacela Velha, 28.iii.2022, 1♂, leg. T.J. Wood, TJWC; Spain: Almodóvar del Campo (Ciudad Real), 700 m, 24.iii.2005, 1♂, leg. F.J. Ortiz-Sánchez, FJOS; Santa Ana la Real, Sierra Aracena (Huelva), 630 m, 13.iv.2006, 2♂, leg. F.J. Ortiz-Sánchez, FJOS; El Hongo (P.N. Doñana), 30.iii.2018, 1♂, leg. F. Molina,
Description. Male. Body length 8–10 mm (Fig.
Diagnosis. The male of A. ramosa is morphologically most similar to A. (Euandrena) solenopalpa due to the long head (only marginally wider than long) and clypeus that is shiny at least in its apical half. The two species are easily separated by the mouthparts, as in A. ramosa the mouthparts that protrude in front of the head are at most as long as the head (viewed frontally or laterally, Fig.
Distribution. South-western Spain (Cádiz, Sevilla).
Remarks. The phylogenetic placement of A. ramosa remains somewhat obscure even following the discovery of the male sex and generation of a barcode sequence. A 658-bp fragment was generated from the female type specimen [BOLD accession number: IBIHM524-21], but this did not fall unambiguously close to any species or subgenus. The most similar sequences belonged to the subgenus Euandrena, specifically to A. symphyti (90.26%), A. montana Warncke, 1973 (90.31%), A. fulvida Schenck, 1853 (89.98%), and A. rufula Schmiedeknecht, 1883 (89.84%). Morphologically, A. ramosa does not fall nicely into Euandrena, as the female sex has foveae which are narrow but which do not narrow ventrally. However, Euandrena are part of the most highly derived clade of Andrena (
Examination of additional material from the province of Cádiz has shown that A. ramosa is commonly encountered in the Parque Natural Los Alcornocales area. Here it can be encountered between January and March, and is most frequently observed on Erica (Ericaceae; Pérez Gómez in litt.). However, the pollen host is still obscure, since none of these bees have been observed collecting pollen. Moreover, Ericaceae pollen is small, with the grains typically having a diameter of 25 μm. The widely spaced and strongly branched and plumose pollen collecting hairs of A. ramosa (described and illustrated by
Material examined. Spain: Carretera Marrufo, Herriza (Cádiz; 3 km E Puerto de Gáliz), 11.xi.2020, 1♀, leg. Á. Pérez Gómez, APGC; Sevilla, Los Pinares de Aznalcázar [37.2782°N, -6.2356°E], 10.iii.2020, 1♀, leg. F. Molina, OÖLM (holotype); Cádiz, Sierra de Montecoche, 31.i.2022, 4♂, 1♀, leg. Á. Pérez Gómez, APGC/TJWC; 18.i.2021, 1♂, leg. Á. Pérez Gómez, APGC; Cádiz, Pico del Montero, 2.ii.2022, 3♂, 1♀, leg. Á. Pérez Gómez, APGC/TJWC; Cádiz, Sierra de Fates, 21.iii.2022, 1♀, leg. Á. Pérez Gómez, APGC; Cádiz, Pico del Montero, Alcalá de los Gazules, 26.iii.2022, 1♂, leg. Á. Pérez Gómez, APGC.
Andrena (Aenandrena) hystrix Schmiedeknecht, 1883: 618, ♀ [France, lectotype by present designation: RMNH].
Remarks.
Material examined. France: Marseille [43.3612°N, 5.3942°E], 1♀,
Andrena (Notandrena) ranunculi Schmiedeknecht, 1883: 617, ♀♂ [France, lectotype by present designation: RMNH].
Remarks.
Material examined. France: Bordeaux [44.8352°N, -0.5888°E], 1♀,
Andrena (Euandrena) symphyti Schmiedeknecht, 1883: 583, ♀♂ [France, lectotype by present designation: RMNH].
Remarks. As for the previous two species, material labelled by Pérez was found in the
Material examined. France: Bordeaux [44.8352°N, -0.5888°E], 1♀,
Andrena (Chlorandrena) distincta Lucas, 1849 nec. Smith, 1847 [Algeria: MNHN, not examined].
Andrena (Chlorandrena) boyerella Dours, 1872: 429, ♀♂ [Morocco: OÖLM].
Neotype. Morocco: Fès-Meknès, Azrou, 4 km SWW of Bakrit, Cascades Bakrit, 33.0466°N, -5.2681°E, 1650 m, 17.v.2022, 1♂, leg. T.J. Wood, OÖLM [BOLD accession number WPATW495-22] (Fig.
Remarks. As discussed above,
Distribution. Morocco, Algeria, Tunisia, Italy (Sicily).
Andrena (Notandrena) erythrocnemis auctorum. nec. Morawitz, 1871.
Andrena (Notandrena) griseobalteata Dours, 1872: 427, ♀ [France: RMNH].
Neotype. France: Pyrénées-Atlantiques, Bérenx [43.4994°N, -0.8575°W], 6.vi.1987, 1♀, leg. E.A.M. Speijer,
Remarks. The correct name to apply to this distinctive taxon has been confused for many years. Through the combination of its large size (for a Notandrena) and densely punctate scutum it is comparable only to A. ungeri Mavromoustakis, 1952. The name A. erythrocnemis Morawitz, 1871 was used by many authors to refer to this taxon (e.g.
Although
Finally, though listed from Spain by
Material examined. Spain: Sierra de Gredos, 12 km SSW Hoyos del Espino, 1950–2100 m, 4.vii.1972, 1♀, leg. J.A.W. Lucas,
Distribution. Spain, France, Italy, Croatia, Hungary, Albania, Romania, North Macedonia, Bulgaria, Greece, Turkey (western and northern Turkey;
Andrena (Taeniandrena) poupillieri Dours, 1872: 430, ♀ [Algeria: OÖLM].
Andrena (Taeniandrena) poupillieri incana Warncke, 1975a: 310, ♀♂ [Spain, Mallorca: OÖLM, examined].
Neotype. Algeria: Tizi-Ouzou, Tigzirt, 36.8877°N, 4.1140°E, 6 m, 31.iii.2017, 1♀, leg. H. Ikhlef, OÖLM [BOLD accession number HYMAA322-22] (Fig.