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.
Remarks. This is the taxon referred to as ‘A. poupillieri 2’ by
Distribution. Morocco, Algeria, Tunisia, Spain (mainland and Balearic Islands). Records (
Andrena succinea Dours, 1872: 424, ♀ [Morocco: OÖLM].
Neotype. Morocco: Oriental, Guercif, P5427, 2 km SW of Bou Rached, 33.8844°N, -3.6154°W, 950 m, 13.v.2022, 1♀, leg. T.J. Wood, OÖLM [BOLD accession number WPATW389-22] (Fig.
Remarks. As discussed above, it is preferable to designate a neotype for A. succinea in order to maintain nomenclatural stability. The barcoded specimen pictured in Fig.
Distribution. Morocco, Algeria, Tunisia, Libya, Egypt, Israel and the West Bank, Jordan, Syria, Saudi Arabia, Iran (
Andrena numida Lepeletier, 1841: 252, ♀ [Morocco: OÖLM].
Neotype. Morocco: Fès-Meknès, Azrou, P7311, 10 km S of Ain Leuh, 1750 m, 33.2220°N, -5.3411°W, 18.v.2022, 1♀, leg. T.J. Wood, OÖLM [BOLD accession number WPATW484-22] (Fig.
Remarks. As discussed above, it is beneficial to designate a neotype for A. numida since the original type series cannot be located in the
Distribution. Morocco, Algeria, Tunisia, Libya, Italy (Sicily, Calabria, Campania).
Following the changes detailed in the previous sections, A. boyerella, A. creberrima, A. curtula, A. hispania, A. mariana s. str., A. potentillae, A. pusilla, and A. truncatilabris are removed from the Iberian fauna following their listing by
Andrena (Cnemidandrena) simillima was listed by
Finally, A. (Melandrena) chrysopyga is listed as present in Iberia. I have seen no Iberian material of this taxon which is generally very rare in collections. It is often confused with forms of A. gravida with light hairs in the terminal fringe. In the distribution maps of Warncke (
In addition to the species elevated above or newly described below, the following 15 Andrena species were explicitly added to the Iberian fauna by the following works: A. (incertae sedis) laurivora, A. (Notandrena) juliana Wood, 2021, A. (Euandrena) fortipunctata Wood, 2021, A. (Taeniandrena) benoisti Wood & Praz, 2021, and A. (Taeniandrena) levante Wood & Praz, 2021 (
Therefore, relative to the baseline of
Results are presented here for Iberian species for which no or very little previous dietary data have been published. Consequently, these results are not comprehensive, but it is not considered necessary to duplicate here previous analyses that have been conducted in Central Europe (e.g.
For many Iberian endemic or West Mediterranean species, an oligolectic dietary niche was clearly and unambiguously demonstrated by pollen analysis. In many cases, specialised pollen use was as expected based on known subgeneric traits, such as the exclusive use of Asteraceae by the subgenus Chlorandrena and Brassicaceae by the subgenera Aciandrena and Nobandrena Warncke, 1968.
It is important to note some pollen collection preferences. Within the Notandrena, members of the former Carandrena are typically associated with Brassicaceae such as A. aerinifrons (Fig.
Pollen specialist (oligolectic) Andrena species in Iberia A Andrena (Notandrena) aerinifrons Dours, 1873 (Brassicaceae) B Andrena (incertae sedis) corax Warncke, 1967 (Reseda, Resedaceae) C Andrena (Chlorandrena) elata Warncke, 1975 (Asteroideae, Asteraceae) D Andrena (Ovandrena) farinosa Pérez, 1895 (Lotus dorycnium, Fabaceae) E Andrena (Ovandrena) oviventris Pérez, 1895 (Reseda, Resedaceae) F Andrena (Simandrena) vetula Lepeletier, 1841 (Brassicaceae).
Members of the relata-group appear to be specialised on Reseda (Resedaceae), including A. corax (Fig.
All studied members of the subgenus Chlorandrena are specialists of Asteraceae. However, the only published associations relate to the subfamily Cichorioideae (e.g.
Within the newly erected subgenus Ovandrena, no clear pattern was seen, as A. farinosa is a specialist of small-flowered Fabaceae (Fig.
Within the subgenus Simandrena, central and northern European species are well-known to be polylectic (
Most studied members of the subgenus Truncandrena are specialised on Brassicaceae (
The subgenus Micrandrena contains mixture of oligolectic and polylectic species in central and northern Europe, though polylectic species predominate (
There was one species for which the empirical data slightly conflicts with what I believe to be the true dietary niche. For A. (Avandrena) panurgina, the pollen results showed that 93.6% of collected pollen came from Geraniaceae, with the remaining pollen from Asteraceae and Brassicaceae. Following the criteria of
The data presented here also resolve the pollen collection preferences of the West Mediterranean A. (Rufandrena) orbitalis Morawitz, 1871 and A. (Rufandrena) rufiventris Lepeletier, 1841 which belong to the subgenus Rufandrena Warncke, 1968 that may contain three species, with a further species known from Syria and Hatay province in Turkey (
Use of Plantago (Plantaginaceae) pollen by Rufandrena Warncke, 1968 species. Andrena (Rufandrena) orbitalis Morawitz, 1871 A female, manipulating Plantago anther B female, vigorously collecting Plantago pollen; Andrena (Rufandrena) rufiventris Lepeletier, 1841 C female approaching Plantago flower head D female manipulating Plantago anthers with forelegs and mandibles E female scraping pollen from Plantago anther using mandibles F female drinking nectar at Reseda (Resedaceae) with empty scopae.
As Plantago is wind-pollinated, it does not provide a nectar source. Nectar is therefore collected from other plants such as Crepis (Asteraceae), Malva (Malvaceae, Álvarez Fidalgo in litt.), or Reseda (Fig.
Finally, a note on the pollen collection preferences of A. afzeliella and A. ovatula is beneficial.
A widespread behaviour within Andrena is bivoltinism, i.e. species producing a spring and a summer generation. For several bivoltine species, their dietary niche is not yet clear. There is a lack of clarity over two species within Aenandrena. For the 15 pollen loads of A. aeneiventris (collected between 11th June and 25th July) and the 22 pollen loads of A. hedikae (collected between 21st May and 25th July), each load was comprised entirely of Apiaceae pollen. However, both species are bivoltine, and pollen loads from the spring generation must be analysed.
In contrast, A. verticalis provides an example of a bivoltine species that uses Apiaceae, but not exclusively. Andrena verticalis flies between March and August, and in this dataset collects 56.3% of its pollen from Brassicaceae (mostly in the spring) and 43.8% of its pollen from Apiaceae (mostly in the summer). However, the use of specific botanical families is not restricted to the two generations, and instead is based on local availability. Interestingly, no mixed pollen loads were detected in the 30 pollen loads examined here, each consisting only of a single botanical family. This recurring pattern of Apiaceae and Brassicaceae use within bivoltine species can be seen for A. hypopolia, which also displays mesolecty on these two families. Additional study is required across these aforementioned species to robustly test the limits of mesolecty on Apiaceae and Brassicaceae across two generations (A. verticalis and A. hypopolia), oligolecty on Apiaceae in both generations (possibly A. colletiformis and A. miegiella), and possible oligolecty in each generation but upon a different botanical family (potentially A. aeneiventris and A. hedikae).
Finally, it is necessary to discuss the case of A. mucida which appears to be differentially oligolectic in its spring and summer generation. The terminology surrounding pollen use in bees has been refined and categorised several times in recent years (
For the 75 Iberian species (not including A. rufiventris) for which pollen data are presented, the majority are oligolectic or assessed as likely to be oligolectic. Excluding species for which insufficient data are available to allow confident classification (A. aeneiventris, A. colletiformis, A. hedikae, and A. miegiella), 49 species are classified as oligolectic, 21 as polylectic, and one (A. mucida) as heterolectic. This high proportion of oligolectic species, most of which have their pollen preferences demonstrated here for the first time, illustrates the degree to which the Iberian fauna i) hosts many specialised species and ii) has been chronically understudied with regard to the basic biology of its constituent bee species. Within the oligolectic species studied here, there is a clear preference for Brassicaceae, this family hosting 22 species, followed by Asteraceae (n=10), Resedaceae (n=6), Cistaceae (n=3), Geraniaceae (n=3), Apiaceae (n=2), and Crassulaceae, Fabaceae, and Plantaginaceae (n=1 each). The broader faunal trends shown by Iberian Andrena species will be examined in detail in a subsequent publication.
The males of A. cilissaeformis, A. erodiorum, A. foeniculae, A. juliae, A. macroptera, A. aff mica, A. ortizi, A. tenostra, and A. urdula are currently unknown, so they are not included in the key; a level of caution must therefore be taken when working with morphologically similar species. The Spanish A. allosa Warncke, 1975 is unknown to me, so the relevant couplet is based on material from the Alps (see
Unfortunately, after consideration, I have also decided to exclude the male of A. exigua from the key because its confident determination is not clear to me. Existing literature (
For supplementary illustrations, please consult the works cited in the Methodology, particularly
The female and male keys are separated for convenience. For the female key, the following shortcuts can be used:
A. Posterior face of hind femur with latitudinal row of spines or teeth go to 2
B. Hind tibial spur clearly broadened, either at its base or submedially. Large to very large species, at least 12 mm in length go to 25
C. Hairs of the tibial scopae clearly plumose go to 42
D. Propodeal triangle clearly defined by strongly raised carinae, internal surface rugose-areolate go to 51
E. Scutum and scutellum with squamous hairs go to 55
F. Viewed laterally, propodeal corbicula with internal surface (lateral faces of the propodeum) glabrous AND propodeal corbiculae complete (possessing both a dorsal and anterior fringe (subgenus Simandrena) go to 64
G. Fovea strongly constricted medially, strongly diverging from the inner margin of the compound eye dorsally (former subgenus Hyperandrena) go to 75
H. At least some tergal discs extensively red-marked go to 77
I. Head, mesosoma, or tergal discs with metallic reflections (note, A. nigroaenea (Kirby) can have bronzy reflections on the terga; if the tibial scopa is composed at least partly of orange-red hairs, go to 205) go to 84
J. Small black species, body length under 8 mm, or if up to 10 mm then with lateral faces of the propodeum with clear pattern of raised star-shaped wrinkles (all members of the subgenera Aciandrena, Graecandrena, Micrandrena) go to 92
K. Clypeus flattened over majority of its surface (subgenus Taeniandrena) go to 139
L. Fovea dorsally narrow, occupying at most ⅓ of space between lateral ocellus and compound eye, ventrally narrowing strongly (subgenus Euandrena) go to 152
M. Clypeus punctate, interspaces forming weakly raised longitudinal wrinkles (former subgenus Zonandrena) go to 163
N. Dorsolateral surface of the propodeum reticulate, with large and shallow punctures, clearly contrasting the shagreened and shiny propodeal triangle, this lacking lateral carinae and becoming shinier on the declivity (subgenus Hoplandrena) go to 168
O. Pronotum laterally keeled, angulate, keel runs up dorsally to an angled corner go to 174
P. Large species (over 12 mm in length). Typically with abundant black, brown, and/or white pubescence. Clypeus strongly domed. Ocelloccipital distance long, at least 2 times the diameter of a lateral ocellus (subgenus Melandrena partim) go to 206
Q. Without this combination of characters; remaining species go to 212
1 | Posterior face of hind femur with latitudinal row of spines or teeth (Fig. |
2 |
– | Posterior face of hind femur without latitudinal row of spines or teeth. A latitudinal carina may be present (e.g. Andrena flavipes) | 24 |
2 | Mesonotum dorsally with short squamous hairs (Fig. |
3 |
– | Mesonotum with longer hairs, these never squamous | 9 |
3 | At least the disc of T2 red-marked | 4 |
– | All terga dark | 5 |
4 | Metasoma entirely red-marked; mesonotum with squamous hairs black. Widespread throughout Iberia, associated with Asphodelus (Asphodelaceae) | sardoa Lepeletier |
– | Metasoma with red markings typically restricted to T2–3; mesonotum with squamous hairs brown-grey. Restricted to the Pyrenees, associated with Campanula (Campanulaceae) | rufizona Imhoff |
5 | T1 comparatively sparsely punctate, large punctures separated by >1 puncture diameter, with scattered micropunctures between these large and relatively coarse punctures. Foveae with outer margin clearly defined, foveae depressed, separated from lateral ocellus by a distance subequal to its diameter (Fig. |
leucolippa Pérez |
– | T1 finely and densely punctate, punctures separated at most by 1 puncture diameter, usually by 0.5 puncture diameters, without micropunctures. Foveae with outer margin poorly defined, foveae not clearly depressed, separated from lateral ocellus by a distance greater than its diameter (Fig. |
6 |
6 | Galea shagreened and dull, clypeus also shagreened and dull with clear impunctate longitudinal midline. Hind tibiae orange. Associated with Ornithogalum (Asparagaceae) | baetica Wood |
– | Galea smooth and shiny, clypeus also at least partially shiny, without impunctate longitudinal midline. Hind tibiae dark. Associated with Campanula (Campanulaceae) | 7 |
7 | Tarsal segment 5 of the hind leg elongate and strongly bent. Squamous hairs light brown. Larger, 13–14 mm | curvungula Thomson |
– | Tarsal segment 5 of the hind leg shorter, at most as long as the two preceding segments taken together, only weakly bent. Squamous hairs darker grey brown. Somewhat smaller, not larger than 12 mm | 8 |
8 | Squamous hairs very thick, in fresh specimens the underlying scutal punctures are obscured. Midline of the scutum is only slightly impressed. Process of labrum markedly elongate with a clear apical emargination in the fore margin. Larger, 10–12 mm | pandellei Pérez |
– | Squamous hairs moderately thick, the underlying scutal punctures clearly visible. Midline of the scutum clearly impressed. Process of labrum regularly trapezoidal, not elongate, fore margin almost straight. Smaller, 8–10 mm | paucisquama Noskiewicz |
9 | Tibial scopae composed predominantly of simple hairs, at most with occasional and scattered plumose hairs (Fig. |
10 |
– | Tibial scopae extensively composed of plumose hairs (Fig. |
13 |
10 | Propodeal triangle poorly defined, without lateral carinae, smooth and with granular shagreen, more or less shiny over the majority of its area. Larger, 9–10 mm | monilia Warncke |
– | Propodeal triangle shagreened and dull or with clearly raised carinae laterally and medially. Smaller, length never exceeding 8 mm | 11 |
11 | Terga strongly and densely punctate, punctures separated by 0.5 puncture diameters, underlying surface smooth and shiny. Propodeal triangle clearly delineated laterally by strongly raised carinae | ventricosa Dours |
– | Terga shagreened and matt, with obscure and shallow punctures. Propodeal triangle lacking lateral raised carinae, poorly defined | 12 |
12 | Process of the labrum narrow, clearly produced into a narrow, apically pointed triangle. Head and mesosoma with black pubescence | panurgina De Steffani |
– | Process of the labrum broad, clearly at least three times broader than long. Head and mesosoma with brownish to greyish-white pubescence | avara Warncke aggregate (potentially including multiple valid species) |
13 | Scutum and scutellum with clear longitudinal striations | rhyssonota Pérez |
– | Scutum and scutellum without longitudinal striations | 14 |
14 | Fovea with inner margin clearly dorsally diverging from the inner margin of the compound eye, curved towards the lateral ocellus (livens-group) | 15 |
– | Fovea with inner margin not dorsally diverging from the inner margin of the compound eye, not noticeably curved towards the lateral ocellus | 17 |
15 | Terga with weak but clear metallic green-blue reflections. Antennae usually ventrally dark, at most obscurely lightened dark brown. Process of the labrum trapezoidal, small, only lightly broader than long | nigroolivacea Dours |
– | Terga dark, without any metallic reflections. Antennae usually extensively lightened orange ventrally. Process of the labrum broader, at least two times wider than long | 16 |
16 | Discs of T2–3 at their bases consistently and densely punctate, punctures typically separated by 1 puncture diameter, at most by 2 puncture diameters. Distributed throughout Iberia | livens Pérez |
– | Discs of T2–3 at their bases more sparsely punctate, punctures separated by more than 3 puncture diameters. Rare, known only from a single specimen captured near Madrid | agnata Warncke |
17 | Foveae strongly constricted ventrally, here narrower than the width of a flagellum (taraxaci-group) | 18 |
– | Foveae not strongly constricted ventrally, more or less as wide as dorsally, not narrower than the width of a flagellum | 20 |
18 | Scutum medially with greatly reduced shagreenation, here more or less smooth and shiny. Facial pubescence relatively dark, with black to dark brown hairs along the inner margins of the compound eyes. Rare, southern and south-eastern Spain only | curtivalvis Morice |
– | Scutum shagreened, uniformly dull across its entire surface. Facial pubescence bright, with at most scattered dark hairs. Throughout Iberia | 19 |
19 | Depressions of terga, especially T3–4, extensively lightened orange-yellow, semi-transparent. Discs of T2–3 densely punctate, punctures with strongly raised rims | senecionis Pérez |
– | Depressions of terga at most with apical rim narrowly lightened orange-yellow. Discs of terga weakly and relatively obscurely punctate, puncture rims weakly raised | rhenana Stöckhert |
20 | Tergal margins entirely lightened whitish-yellow hyaline. Face (particularly vertex), scutum, and scutellum with subtle metallic blue-green reflections | 21 |
– | Tergal margins never entirely lightened, at most narrowly lightened yellow hyaline. Face (particularly vertex), scutum, and scutellum dark, without metallic reflections | 22 |
21 | Found in dry to steppic areas in central Spain. Scutum with clear punctures that are visible against the strong background shagreenation. Scutellum with shiny interspaces between punctures |
elata Warncke |
– | Found in areas close to or on the coast in southern Portugal and Spain. Scutum with obscure punctures that disappear into the strong background shagreenation. Scutellum with interspaces shagreened |
abrupta Warncke |
22 | T2–4 depressed at their base, marginal areas of T2–4 with apical white hair bands that obscure the underlying surface in fresh specimens. Posterior face of the hind femur with long spines | orbitalis Morawitz |
– | T2–4 not depressed basally, apically with marginal areas with at most scattered orange-yellow hairs, these not forming distinct bands which obscure the underlying surface. Posterior face of the hind femur with short teeth | 23 |
23 | T2 between the punctures weakly shagreened, comparatively shiny. Larger species, over 10 mm. Found in more temperate parts of Iberia, generally absent in hot Mediterranean habitats |
humilis Imhoff |
– | T2 between the punctures strongly shagreened and dull. Smaller species, under 10 mm. Found in hotter parts of Iberia, the dominant taxon in Mediterranean habitats |
cinerea Brullé |
24 (1) | Hind tibial spur clearly broadened, either at its base (Fig. |
25 |
– | Hind tibial spur not broadened at its base or submedially, more or less parallel-sided. Size variable | 41 |
25 | Propodeum dorsolaterally with dense punctures, punctures separated by <1 puncture diameter (Fig. |
26 |
– | Propodeum dorsolaterally with raised reticulation or rugosity (Fig. |
29 |
26 | Vertex wide, ocelloccipital distance at least as wide as three times the diameter of a lateral ocellus | variabilis Smith |
– | Vertex narrower, ocelloccipital distance never as wide as three times the diameter of a lateral ocellus | 27 |
27 | Disc of T1 densely punctate, punctures typically separated by 1 puncture diameter. Scutellum dull between punctures. Tergal hair bands interrupted medially | labialis (Kirby) |
– | Disc of T1 more sparsely punctate, punctures separated by greater than 1 puncture diameter, typically by 2 puncture diameters. Scutellum shiny between punctures. Tergal hair bands complete in fresh specimens | 28 |
28 | Flying in the spring (April–May). Terminal fringe dark blackish-brown. Ocelloccipital distance narrower, 1 times the diameter of a lateral ocellus |
flavilabris Schenck |
– | Flying in the summer (July–August). Terminal fringe light. Ocelloccipital distance wider, 1–1.5 times the diameter of a lateral ocellus |
decipiens Schenck |
29 | Hind tibial spur broadened submedially. Propodeal triangle simply defined laterally by weak and obscurely raised carinae, internal surface without a dense network of strongly raised honeycomb-like rugosity. Terga often red-marked. Exclusively summer flying species, from May onwards | 30 |
– | Hind tibial spur broadened at its base. Propodeal triangle clearly defined laterally by strongly raised carinae (Fig. |
31 |
30 | Tibial scopa ventrally composed of long plumose hairs. Posterior face of the hind femora with clear latitudinal carina. Terga usually red marked, though an entirely melanic form can be found in south-eastern Spain (ssp. nigricauda Wood). Associated with scabious (former Dipsacaceae = Caprifoliaceae). Restricted to montane grasslands in northern and central Spain with isolated populations in the Sierra de Cazorla and Sierra Nevada | hattorfiana (Fabricius) |
– | Tibial scopa simple or with at most occasional obscurely plumose hairs. Posterior face of the hind femora without a latitudinal carina. Terga never red-marked. Associated with yellow Cichorioideae (Asteraceae). Restricted to the Pyrenees and Cantabrian Mountains | polita Smith |
31 | Terminal fringe dark medially and white laterally (Fig. |
32 |
– | Terminal fringe uniformly dark. Lateral faces of the propodeum without ridges, either unsculptured or at most with individually raised points which do not joint together to form a network | 34 |
32 | Disc of T1 strongly and densely punctate, punctures separated by <1 puncture diameter | asperrima Pérez |
– | Disc of T1 with sparse punctate, punctures on average separated by 2 puncture diameters | 33 |
33 | Metasoma with metallic blue reflections. T4 laterally with thick white hair patches. Pygidial plate narrow, laterally with a shiny depressed marginal area | agilissima Scopoli |
– | Metasoma black, without metallic blue reflections. T4 laterally with only loose white hair, not forming dense hair patches. Pygidial plate broad, flat, without a depressed and shiny marginal area | afrensis Warncke |
34 | Terga with metallic blue-green reflections. Mesepisternum and dorsolateral faces of the propodeum more or less smooth, granularly shagreened, weakly and obscurely punctate (subgenus Suandrena) | 35 |
– | Terga black or red-marked, but never with metallic reflections. Mesepisternum and dorsolateral faces of the propodeum with dense network of raised rugosity | 37 |
35 | Propodeal triangle weakly defined, internal rugosity fine and obscure. Known only from Cádiz province, flying in December | gades Wood & Ortiz-Sánchez |
– | Propodeal triangle strongly defined, with pronounced internal rugosity. More widespread across Iberia | 36 |
36 | Process of the labrum triangular with clear apical point. Throughout Iberia |
suerinensis Friese |
– | Process of the labrum broadly triangular with apical margin truncate. Confirmed males known only from central, eastern, and south-eastern Spain |
cyanomicans Pérez |
37 | Pronotum laterally with strong humeral angle. Terga with extremely fine punctation, punctures minute, separated by <0.5 puncture diameters | fuscosa Erichson |
– | Pronotum laterally rounded, without humeral angle. Terga with much coarser punctation, punctures separated by >1 puncture diameter | 38 |
38 | Pubescence of body predominantly black, tibial scopa predominantly composed of white hairs | 39 |
– | Pubescence of body variable, but usually with extensive brown hairs on mesosoma; tibial scopa usually orange, never predominantly composed of white hairs | 40 |
39 | Bivoltine (typically March–April and July–August). Common and widely distributed throughout Iberia |
pilipes Fabricius |
– | Univoltine (typically May–June). Rare, restricted to mountainous parts of Iberia; known from the Pyrenees, the Sistema Central, Serra do Gerês, Sistema Ibérico, and Sierra de Cazorla |
nigrospina Thomson |
40 | Depression of T1 sparsely punctate, punctures separated by more than 2 puncture diameters. Nominally univoltine, flying only in the spring |
tibialis (Kirby) |
– | Depression of T1 densely punctate, punctures separated by 0.5 puncture diameters. Bivoltine, flying in the spring and the summer |
bimaculata (Kirby) |
41 (24) | Hairs of the tibial scopae clearly plumose, with a majority of strongly branched hairs | 42 |
– | Hairs of tibial scopae entirely simple, or with at most a mixture of simple and scattered and weakly branched hairs | 50 |
42 | Terga metallic blue, densely punctate, punctures separated by 1 puncture diameter | bellidis Pérez |
– | Terga variably coloured, but never metallic blue. Densely punctate or not | 43 |
43 | Terga with discs extensively red-marked (some dark individual of A. marginata Fabricius with red markings restricted to the base of the tergal margins) | 44 |
– | Terga dark, at most with depressions lightened, never with red markings at the base of the tergal depressions | 45 |
44 | Clypeus with fore margin slightly upturned. Foveae short, ventrally reaching only to the level of the antennal insertions. Tibial scopae uniformly yellowish | pellucens Pérez |
– | Clypeus with fore margin straight, not upturned. Foveae long, ventrally extending beyond the level of the antennal insertions. Tibial scopae bicoloured, black-brown dorsally, brownish-white ventrally. Associated with scabious (former Dipsacaceae = Caprifoliaceae) | marginata Fabricius |
45 | Tibial scopae with short hairs, at most half the length of the diameter of the hind tibia at its maximum apical width | ranunculi Schmiedeknecht |
– | Tibial scopae with long hairs, clearly exceeding half the length of the diameter of the hind tibia at its maximal apical width | 46 |
46 | Fovea narrow, dorsally occupying ⅓ of the space between the compound eye and the lateral ocellus, clearly narrowing ventrally to half its dorsal width. Mid and hind basitarsi and hind tibiae lightened orange-yellow (subgenus Chrysandrena Hedicke) | 47 |
– | Fovea broader, dorsally occupying at least ½ the space between the compound eye and the lateral ocellus, not narrowed ventrally. Basitarsi and hind tibiae dark | 49 |
47 | Clypeus and scutellum strongly shagreened and matt | hesperia Smith |
– | Clypeus and scutellum at least partially shiny | 48 |
48 | Disc of scutum shiny. Scutellum with uniformly light hairs. A2 as long as A3+4. Restricted to temperate areas in northern Portugal and Spain | fulvago (Christ) |
– | Disc of scutum shagreened and dull. Scutellum with intermixed light and dark brown to black hairs. A2 as long as A3+4+5. Found in Mediterranean areas from southern Portugal to southern and eastern Spain | fertoni Pérez |
49 | Process of the labrum large, as long as broad, ventral surface covered with latitudinal wrinkles (Fig. |
mucida Kriechbaumer (partim, 2nd generation) |
– | Process of the labrum twice as broad as long, without wrinkles. Foveae dorsally strongly impressed and therefore well-defined. Terminal fringe composed of densely plumose orange hairs. Flying in the spring (April-May), Associated with yellow Cichorioideae (Asteraceae) | alluaudi Benoist |
50 (41) | Propodeal triangle clearly defined by strongly raised carinae, internal surface rugose-areolate (Fig. |
51 |
– | Propodeal triangle not strongly defined by lateral carinae with its internal surface rugose-areolate (Fig. |
54 |
51 | Forewing with two submarginal cells. Clypeus with longitudinal striations | lagopus Latreille |
– | Forewing with three submarginal cells. Clypeus without longitudinal striations | 52 |
52 | Hind tibiae usually orange. Foveae clearly medially constricted. Terga without hair bands, T2 laterally without a pair of small foveae. Terminal fringe orange | haemorrhoa (Fabricius) |
– | Hind tibiae always dark. Foveae lacking medial constriction. Terga in fresh specimens with apical white hair bands, T2 laterally with a pair of small foveae. Terminal fringe whitish-yellow | 53 |
53 | Process of the labrum clearly deeply emarginate | colletiformis Morawitz |
– | Process of the labrum triangular, anterior margin truncate, never emarginate | miegiella Dours |
54 (50) | Scutum and scutellum with short squamous hairs (Figs |
55 |
– | Scutum and scutellum without squamous hairs. If specimen shows longer semi-squamous hairs (e.g. A. farinosa Pérez, Fig. |
63 |
55 | Ocelloccipital distance at least three times the diameter of a lateral ocellus (Figs |
56 |
– | Ocelloccipital distance less than two times the diameter of a lateral ocellus. Smaller bees, exceptionally reaching 11 mm in length | 60 |
56 | Dorsolateral faces of the propodeum clearly, densely, and deeply punctate, punctures separated by <0.5 puncture diameters, with shiny interspaces | 57 |
– | Dorsolateral faces of the propodeum either impunctate, with dense network of raised rugosity, or with shallow and sparser punctures, punctures separated by 0.5–2 puncture diameters, interspaces dull | 59 |
57 | 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. Restricted to eastern and south-eastern Spain | nilotica Warncke |
– | Pubescence of scutum short and strongly squamous, anterior dorsolateral corners of scutum with pubescence short, clearly shorter than width of antennae. Terga variable, red to black or any intermediate combination. Terga with or without clear apical hair bands. Punctation of T1 denser, punctures separated at most by 1 puncture diameter | 58 |
58 | 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 (Fig. |
pruinosa Erichson |
59 | Scutal hairs orange-brown (Fig. |
limbata dusmeti Warncke |
– | Scutal hairs black and whitish-brown. Hind tibiae dark. Terga with clear and thick white apical hair bands. Terminal fringe dark brown | lateralis Morawitz |
60 | T1 strongly and densely punctate, interspaces shiny (Fig. |
oviventris Pérez |
– | T1 weakly and obscurely punctate, underlying surface shagreened. Mesepisternum ventrally with an indentation anterior to the attachment point of the mid leg | 61 |
61 | Clypeus without latitudinal ridges. T3–4 laterally with clear squamous hairs between the disc and marginal areas. Terga usually extensively red-marked. Basitarsi and hind tibiae orange | hystrix Schmiedeknecht |
– | Clypeus with latitudinal ridges. T3–4 laterally without squamous hairs. Terga never red marked. Legs dark | 62 |
62 | T2–3 laterally strongly shagreened, with obscure and scattered punctures | aeneiventris Morawitz |
– | T2–3 laterally finely shagreened, clearly and densely punctate, punctures separated by 0.5 puncture diameters | hedikae Jäger |
63 (54) | Viewed laterally, propodeal corbicula with internal surface (lateral faces of the propodeum) glabrous AND propodeal corbiculae complete (possessing both a dorsal and anterior fringe) (Fig. |
64 |
– | Propodeal corbiculae either incomplete (possessing only a dorsal fringe) AND/OR internal surface with hairs | 74 |
64 | Tibial scopae bicoloured, dorsally dark and ventrally pale | 65 |
– | Tibial scopae unicolourous, dorsally and ventrally light | 68 |
65 | Fovea extremely broad, occupying entirety of space between the compound eye and the lateral ocellus (Fig. |
rhypara Pérez |
– | Fovea narrower, not occupying entirety of space between the compound eye and the lateral ocellus. T1 strongly shagreened, impunctate or with scattered punctures | 66 |
66 | T2 impunctate. A2 longer than A3+4, almost as long as A3+4+5 | vetula Lepeletier |
– | T2 with fine and dense punctures, punctures separated by 0.5 puncture diameters. A2 as long as A3+4 | 67 |
67 | Terga with thick white apical hair bands in fresh specimens (Fig. |
cilissaeformis Pérez |
– | Terga with at most obscure narrow brownish hair bands. Foveae with outer margin constricted, clearly deviating from inner margins of the compound eyes submedially (Fig. |
antigana Pérez |
68 | Tibial scopae with short hairs, dorsally these hairs not greatly exceeding the width of a lateral ocellus. Metasoma with punctures on tergal discs dense medially, becoming sparse laterally | 69 |
– | Tibial scopae with long hairs, very clearly greatly exceeding the width of a lateral ocellus. Metasoma with consistently dense punctures, not becoming sparser laterally | 70 |
69 | Scutum medially strongly shagreened and dull (Fig. |
dorsata (Kirby) |
– | Scutum medially polished and shiny (Fig. |
propinqua Schenck |
70 | Disc of T1 strongly and densely punctate, punctures separated by <1 puncture diameter | 71 |
– | Disc of T1 finely and sparsely punctate, punctures separated by >1 puncture diameter | 72 |
71 | Clypeus with pattern of raised latitudinal ridges | combinata (Christ) |
– | Clypeus smooth in the middle, at most transversely wrinkled at the base, ventro-laterally with slight longitudinal wrinkles | lepida Schenck |
72 | Head and mesosoma white-haired. Clypeus smooth and shiny between punctures. Restricted to areas close to the Pyrenees | thomsonii Ducke |
– | Head and mesosoma with rich chestnut-brown hair. Clypeus shagreened and dull between punctures | 73 |
73 | T2–4 with discs finely and densely punctate. Terminal fringe dark brown. Hind tibiae sometimes lightened orange. More widespread across Iberia |
congruens Schmiedeknecht |
– | T2–4 with discs sparsely and obscurely punctate. Terminal fringe reddish-brown. Restricted to the Pyrenees and Cantabrian Mountains |
confinis Stöckhert |
74 (63) | Fovea strongly constricted medially, strongly diverging from the inner margin of the compound eye dorsally (Fig. |
75 |
– | Fovea not constricted medially, not strongly diverging from the inner margin of the compound eye | 76 |
75 | Tibial scopae unicolourous black. Mesosoma entirely covered with light grey hairs | bicolorata (Rossi) |
– | Tibial scopae dorsally black, ventrally orange-red. Mesosoma entirely covered with dark to light brown hairs | florentina Magretti |
76 (74) | At least some tergal discs extensively red-marked (Fig. |
77 |
– | Tergal discs never red-marked, at most with tergal margins lightened | 83 |
77 | Small bees, never exceeding 9 mm in length. Fovea narrow, at most occupying ⅓ of space between the compound eye and the lateral ocellus | 78 |
– | Large bees, greater than 11 mm in length. Fovea broader, occupying at least ½ the space between the compound eye and the lateral ocellus | 80 |
78 | Propodeal triangle broad, laterally delineated with raised straight carinae, internal surface with fine network of raised rugae. Head and scutum without metallic reflections | labiata Fabricius |
– | Propodeum narrow, poorly defined, lacking lateral carinae, internal surface at most with short and weak rugae basally, surface with fine granular shagreenation. Head and scutum with strong or weak metallic reflections | 79 |
79 | Clypeus medially smooth and shiny between the punctures. Scutum anteriorly shagreened, medially becoming smooth and shiny, irregularly but clearly punctate, with strong greenish metallic reflections | binominata Smith |
– | Clypeus medially completely shagreened and dull. Scutum shagreened and shallowly punctate, with weak bronzy metallic reflections | leucophaea Lepeletier |
80 | Tibial scopae unicolourous light orange. Propodeal triangle clearly delineated laterally by fine carinae, internal surface with fine network of raised rugae. Clypeus densely and uniformly punctate, without a longitudinal impunctate midline | schencki Morawitz |
– | Tibial scopae bicoloured, dorsally dark, ventrally light (Fig. |
81 |
81 | Terga strongly and densely punctate, punctures separated by 1 puncture diameter. Compound eyes with inner margins diverging ventrally (Fig. |
florea Fabricius |
– | Terga shallowly and obscurely punctate, punctures separated by 2–3 puncture diameters. Compound eyes with internal margins more or less parallel. Excluding impunctate midline, clypeus comparatively densely punctate, punctures separated by 0.5–1 puncture diameters | 82 |
82 | T1–2 with long hair, equalling the length of the hair on the mesosoma |
trimmerana (Kirby) (partim, light form) |
– | T1–2 with shorter hair, never equalling the length of the mesosomal hair |
rosae Panzer (partim, light form) |
83 (76) | Head, mesosoma, or tergal discs with metallic reflections (note, A. nigroaenea (Kirby) can have bronzy reflections on the terga; if the tibial scopae is composed of orange-red hairs or the tibial scopae is mainly dark with only the ventral hairs orange, go to 206) | 84 |
– | Body without metallic reflections | 91 |
84 | Mesosoma with strongly contrasting pattern of black and white hairs; mesosoma anteriorly and posteriorly with white hairs, medially with a band of black hairs | 85 |
– | Mesosoma without strongly contrasting pattern of black and white hairs | 86 |
85 | Wings infuscate over their apical 2/3rds. Sterna laterally and apically with black hairs. A3 comparatively shorter, shorter than A4+5. Bivoltine (April–May; July–August), restricted to mountains in northern Spain and the Pyrenees |
barbareae Panzer |
– | Wings hyaline over the majority of their area, only slightly darkened apically. Sterna laterally and apically with white hairs. A3 comparatively longer, equalling A4+5. Univoltine (April–May), more widespread across northern Portugal and Spain |
cineraria (Linnaeus) |
86 | Scutum uniformly densely shagreened and completely dull over its entire surface, extremely densely punctate, punctures shallow, flat, and confluent. Mesosoma and discs of T1–2 dorsally with long chestnut-brown hair | doursana Dufour |
– | Scutum either at least partly shiny or less densely punctate, punctures clearly separated by at least 1 puncture diameter. Mesosoma and discs of T1–2 never with long chestnut-brown hair, either glabrous, with shorter hairs, or with hairs of a different colour | 87 |
87 | Terga clearly punctate, punctures separated by up to 2 puncture diameters | 88 |
– | Terga obscurely and sparsely punctate, punctures separated by 4–5 puncture diameters | 89 |
88 | Fovea broad, occupying 2/3rds of the space between the compound eye and a lateral ocellus. Tergal punctation comparatively sparse, punctures separated by 1–2 puncture diameters. Scutum medially becoming smooth and shiny between the punctures. Restricted to south-western Spain (Huelva, Sevilla) | laurivora Warncke |
– | Fovea narrow, occupying less than ½ the space between the compound eye and a lateral ocellus. Tergal punctation dense, punctures separated by up to 1 puncture diameter. Scutum uniformly shagreened and dull. Restricted to the Pyrenees | viridescens Viereck |
89 | Larger species, 11–12 mm. Clypeus shagreened and dull over the majority of its surface. Fovea narrow, but uniformly wide along their length, not narrowing ventrally | aerinifrons Dours |
– | Smaller species, under 9 mm. Clypeus shiny over the majority of its area. Foveae narrow, but also narrowing ventrally to approximately half of their dorsal width | 90 |
90 | Scutum with clear metallic green reflections. Hind tibial spurs straight (Fig. |
nigroviridula Dours |
– | Scutum dark, with at most weak and obscure metallic reflections. Hind tibial spurs strongly bent at their apexes (Fig. |
varuga Warncke |
91 (83) | Small black species without a keel laterally on the pronotum, body length under 8 mm, or if up to 10 mm then with lateral faces of the propodeum with clear pattern of raised star-shaped wrinkles (all members of the subgenera Aciandrena, Graecandrena, Micrandrena) |
92 |
– | Species larger than 9 mm in length or with pronounced keel on the pronotum laterally | 138 |
92 | Propodeal triangle smooth, not defined laterally by raised carinae, with internal surface lacking network of raised rugosity, at most with very short rugae at the base of the propodeal triangle (Fig. |
93 |
– | Propodeal triangle either strongly defined by raised carinae, or with internal surface with network of raised rugosity covering at least the basal half (Fig. |
106 |
93 | Clypeus with clear raised longitudinal striations (Fig. |
94 |
– | Clypeus without any striations | 96 |
94 | Clypeus flattened, medially slightly depressed, weakly concave. Process of the labrum short, twice as broad as long, forming a triangular point. A3 almost as long as A4+5+6. Central and eastern Spain only | fria Warncke |
– | Clypeus evenly arched. Process of the labrum narrower, as long as broad. A3 only slightly exceeding A4+5. More widespread | 95 |
95 | Tibial scopa composed of unicolourous light hairs. T3 impunctate, with weak apical yellowish hair band, marginal area at most weakly depressed. Widespread across Iberia | longibarbis Pérez |
– | Tibial scopa pale ventrally, dark dorsally. T3 with obscure punctures, with strong apical white hair band overlying the clearly depressed marginal area. Restricted to sandy and usually coastal habitats in southern Iberia | orana Warncke |
96 | Foveae not ventrally narrowed, in their lower half at least half as wide as the distance from the inner margin of the compound eye (Fig. |
97 |
– | Foveae ventrally narrowed, in their lower half as wide as the distance from the inner margin of the compound eye (Fig. |
98 |
97 | T2–4 with narrow, widely interrupted hair bands. Wing venation brownish. Clypeus shagreened to smooth and shiny, densely punctate, punctures separated by 1 puncture diameter. Process of the labrum narrow and triangular with a pointed tip (Fig. |
pandosa trigona Warncke |
– | T2–4 with complex wide and dense white hair bands. Wing venation light yellow. Clypeus uniformly shagreened, irregularly punctate, punctures separated by 1–3 puncture diameters. Process of the labrum with the apical margin truncate. Smaller, 6 mm. Rare, known only from central Spain | montarca Warncke |
98 | Tergal discs clearly and regularly punctate, punctures separated by 1 puncture diameter, with punctures extending onto marginal areas | fulica Warncke |
– | Terga either impunctate or obscurely punctate, but punctures never extending onto marginal areas | 99 |
99 | Supraclypeal area covered with longitudinal striations (Fig. |
100 |
– | Supraclypeal area without any striations (Fig. |
105 |
100 | Longitudinal striations on the paraocular areas strong and pronounced, continuing ventrally to the lateral margins of the clypeus without becoming weaker. Clypeus flattened, with distinct longitudinal impression or furrow medially. Restricted to the extreme north-east of Spain | impunctata Pérez |
– | Longitudinal striations on the paraocular areas ventrally extending to the lateral margins of the clypeus but here clearly weaker than their strength adjacent to the foveae. Clypeus either domed or if flattened then without longitudinal impression | 101 |
101 | Clypeus strongly flattened, coarsely shagreened and dull over almost its entirely surface, apical margin narrowly and obscurely shiny; clypeus with shallow and obscure punctures that disappear into the underlying shagreen, punctures separated by 1–2 puncture diameters (Fig. |
verticalis Pérez |
– | Clypeus either domed, or if flattened then with a broad shiny apical margin (at least as broad at the width of a flagellum) and punctures that are clearly visible against the underlying shagreenation | 102 |
102 | A3 exceeding the length of A4+5. Striations of supraclypeal area weakly continue onto clypeus basally and laterally, here shagreened with weakly raised striations. Clypeus shagreened basally and laterally, becoming smooth and shiny medially and apically, with narrow medial shagreened projection, shagreenation thus forming a weak trident shape. Known only from a few specimens from the extreme south of Spain, probably representing an undescribed species (Málaga, Sevilla) | aff mica Warncke |
– | A3 at most equalling A4+5. Striations of supraclypeal area not continuing onto clypeus, entire clypeus free of even a hint of striations | 103 |
103 | Clypeus with fine punctures, punctures separated by 1–3 puncture diameters. Underlying shagreenation weak basally. Process of the labrum slightly broader than long. Presence and distribution in Iberia unclear | abjecta Pérez |
– | Clypeus with strong and coarse punctures, punctures separated by 0.5–2 puncture diameters. Underlying shagreenation strong and coarse basally. Process of the labrum narrow, slightly longer than broad | 104 |
104 | Scutellum polished and shiny between punctures. Clypeus domed and somewhat flattened medially. Restricted to south-eastern Spain (Alicante, Almería, Granada, Murcia) | tenostra Warncke |
– | Scutellum dull and shagreened. Clypeus largely flattened across its entire surface. More widespread across Iberia | alma Warncke |
105 | Larger, 7–8 mm. T3–4 with discs obscurely punctate. Hind basitarsi lightened orange. Nervulus of the forewing interstitial to weakly antefurcal. Shagreenation of the clypeus becoming weaker at its apical margin | nebularia Warncke |
– | Smaller, 4–5 mm. T3–4 with discs impunctate. Hind basitarsi dark. Nervulus of the forewing strongly antefurcal. Clypeus uniformly shagreened | vacella Warncke |
106 | Lateral shoulders of T1 with a pair of strongly produced sharp ridges (Fig. |
taxana Warncke |
– | Lateral shoulders of T1 never with clearly produced ridges, or if with ridges (A. strohmella Stöckhert) then terga never shiny and deeply punctate. Tergal punctation otherwise. Mesepisternum evenly rounded | 107 |
107 | Hind tibiae and basitarsi orange, tibial scopa composed of extremely short hairs. Restricted to the Pyrenees and the Cantabrian Mountains, flying July–August, associated with Potentilla (Rosaceae) | tarsata Nylander |
– | Hind legs dark, tibial scopa with hairs normal, not extremely short | 108 |
108 | Foveae long, dorsally extent reaching a line parallel to the hind margin of the lateral ocelli, foveae deeply impressed. Propodeal triangle not laterally delineated by carinae, internal surface weakly elevated, with irregular raised rugae that do not cover the entire area. Terga laterally with loose, white interrupted hair bands. Restricted to cooler areas in and around the Pyrenees and Cantabrian Mountains, flying July–August | coitana (Kirby) |
– | Foveae shorter, not reaching level of the lateral ocelli dorsally, only weakly impressed. Propodeal triangle clearly delineated laterally by carinae, internal surface evenly and regularly covered by fine network of raised rugae. Terga with lateral hair bands or not | 109 |
109 | Foveae either uniformly narrow or strongly narrowed ventrally, in their ventral half at most as wide as the distance to the inner margin of the compound eye | 110 |
– | Foveae not or only weakly narrowed ventrally, in their ventral half clearly wider than the distance to the inner margin of the compound eye | 115 |
110 | Clypeus with longitudinal striations (c.f. Fig. |
111 |
– | Clypeus without striations | 112 |
111 | Disc of T1 shagreened and dull. T2–4 with hair bands long, longer than the diameter of a flagellum. Scutum uniformly shagreened and dull. Restricted to the extreme north-east of Spain | distinguenda Schenck |
– | Disc of T1 polished and shiny, contrasting the shagreened marginal area. T2–4 with short hair bands, shorter than the diameter of a flagellum. Scutum laterally shagreened, medially smooth and shiny. Widespread throughout Iberia | nitidula Pérez |
112 | T1 finely shagreened to polished and shiny. Mesonotum densely and strongly punctate | 113 |
– | T1 strongly shagreened and dull. Mesonotum finely and sparsely punctate | 114 |
113 | Foveae uniformly narrow, dorsally narrower than the diameter of a flagellum. Disc of T1 with scattered punctures, separated by at least 2 puncture diameters | fabrella Pérez |
– | Foveae dorsally broader than ventrally, dorsally as wide as the diameter of a flagellum. Disc of T1 more densely punctate, punctures separated by 0.5–1 puncture diameters | nana (Kirby) |
114 | Foveae uniformly narrow, dorsally narrower than the diameter of a flagellum. Terga impunctate. Clypeus evenly domed, densely shagreened and dull. Associated with Cistaceae | djelfensis Pérez |
– | Foveae dorsally broader than ventrally, dorsally as wide as the diameter of a flagellum. Terga obscurely punctate. Clypeus weakly three-faced, anterior margin shiny. Associated with Brassicaceae | tenuistriata Pérez |
115 | Large species, 10 mm in length. Clypeus densely covered with latitudinal wrinkles. Lateral faces of the propodeum covered with pattern of raised star-shaped wrinkles | ampla Warncke |
– | Most species smaller. Clypeus without latitudinal wrinkles and propodeum without pattern of raised star-shaped wrinkles | 116 |
116 | Larger species, 8–9 mm. Discs of T1–3 extremely densely punctate, punctures almost confluent, separated by <0.5 puncture diameters. Clypeus densely punctate, punctures separated by 1 puncture diameter, interspaces with weakly raised ridges that form subtle pattern of longitudinal striations. Rare, known from eastern Spain (Jáen, Soria, Teruel) | vaulogeri Pérez |
– | Smaller, usually shorter than 8 mm. Discs of T1–3 less densely punctate, punctures separated by >1 puncture diameter, or if punctures dense, then much smaller than 8 mm in length. Clypeus without pattern of longitudinal ridges or striations | 117 |
117 | T3–4 with the marginal areas strongly depressed relative to the discs (Fig. |
118 |
– | T3–4 with the marginal areas only superficially depressed (Fig. |
131 |
118 | T2–3 laterally without a gradulus | 119 |
– | T2–3 laterally with gradulus present (c.f. Fig. |
122 |
119 | Tergal discs and marginal areas completely smooth and shiny, without any shagreenation (Fig. |
omnilaevis Wood |
– | At least tergal discs shagreened | 120 |
120 | Terga with marginal areas not noticeably differentiated structurally from discs, strongly and densely shagreened and dull (Fig. |
pauxilla Stöckhert |
– | Terga with marginal areas noticeably shinier than discs, with weaker shagreenation. Tergal discs with clear punctures visible against the underlying shagreenation (Fig. |
121 |
121 | Depressed tergal margins polished and shiny, clearly shinier than shagreened tergal discs. In dorsal view, lateral hairs at the base of the marginal area of T2 longer, erect, projecting at a 45° angle, breaking profile of metasoma laterally. Margin of T2 comparatively strongly depressed. Restricted to the Pyrenees and the Cantabrian Mountains | semilaevis Pérez |
– | Depressed tergal margins shagreened, similar to and not strongly contrasting shagreened tergal discs (Fig. |
anthrisci Blüthgen |
122 | Disc of T1 smooth and shiny, clearly punctate, punctures medially separated by 0.5–1 puncture diameter, not extending onto the broad impunctate marginal area. Rare, restricted to the Pyrenees and northern Spain | floricola Eversmann |
– | Disc of T1 never smooth and shiny, always at least partially shagreened, punctate or not | 123 |
123 | Gena entirely smooth and shiny, without any shagreenation. Scutum and scutellum polished and shiny. T2–3 with discs strongly and densely punctate, punctures separated by 1 puncture diameter (Fig. |
bayona Warncke |
– | Gena shagreened and dull, at most with a narrow shiny strip along the outer margin of the compound eye. Scutum and scutellum variable, usually at least partially shagreened. T2–3 usually less densely and strongly punctate | 124 |
124 | Head comparatively long, only slightly shorter than broad (Fig. |
ortizi sp. nov. |
– | Head shorter, clearly broader than long. T1–3 never so densely punctate, punctures when visible separated by at least 1 puncture diameter | 125 |
125 | Propodeal triangle poorly defined (Fig. |
126 |
– | Propodeal triangle well-defined, lateral carinae consistently strong from the base to the apex. Tergal discs weakly to strongly shagreened, never shiny | 127 |
126 | Tergal discs entirely shiny, polished, and smooth (Fig. |
lecana Warncke |
– | Tergal discs at least partially shagreened, never entirely shiny (Fig. |
niveata Friese |
127 | T1 laterally with slightly raised corners, visibly projecting and disrupting rounded profile when viewed dorsally. Clypeus slightly flattened centrally, with clear impunctate mid-line between large and well defined clypeal punctures. Terga 2–3 centrally impunctate, laterally with large punctures with raised margins (crater punctures). Restricted to areas close to the Pyrenees | strohmella Stöckhert |
– | T1 laterally without such raised projections, in dorsal view evenly rounded. Clypeus with or without a clear impunctate mid-line. Terga punctation with or without lateral crater punctures. Distribution otherwise | 128 |
128 | Clypeus weakly domed, sparsely punctate, punctures large, separated by 2–3 puncture diameters. T2–3 laterally with large hair-bearing punctures that contrast the underlying shagreenation and do not disappear into it | icterina Warncke |
– | Clypeus more densely punctate, punctures separated by 1–2 puncture diameters. T2–3 laterally with at most obscure punctures | 129 |
129 | Terga comparatively weakly shagreened and finely punctate, punctures relatively clearly visible against background shagreen, punctures extending onto tergal margins, most clearly visible on T1–3. Anterior ½ to ⅓ of clypeus usually polished and shiny. Probably univoltine (March-May). Central, eastern, and southern Spain, rare |
exigua Erichson |
– | Terga comparatively strongly shagreened, obscurely punctate, punctures disappearing into underlying shagreenation (Fig. |
130 |
130 | Scutum strongly and clearly punctate, punctures separated by 1–2 puncture diameters, underlying surface variable but always at least partially shiny. Bivoltine (March-July). Throughout Iberia, common |
spreta Pérez |
– | Scutum obscurely punctate, punctures disappearing into the abundant underlying shagreen, scutum dull. Univoltine (April-May). Iberian distribution unclear, but probably restricted to southern Spain (Cádiz, Málaga) |
tiaretta Warncke |
131 | Disc of T1 clearly punctate, though sometimes finely and sometimes only punctate basally (Fig. |
132 |
– | Disc of T1 finely to strongly shagreened, at most obscurely punctate with punctures disappearing into the underlying surface sculpture (Fig. |
134 |
132 | Marginal area of T1 wide, strongly thickened, impunctate (Fig. |
falsifica Perkins |
– | Marginal area of T1 normal, not widened or thickened | 133 |
133 | Disc of T1 strongly shagreened, predominantly punctured medially. Antennae ventrally at most lightened dark brown. Throughout Iberia, bivoltine | alfkenella Perkins |
– | Disc of T1 finely shagreened, evenly and finely punctate over its entire area, with shiny interspaces. Antennae ventrally orange. Rare, restricted to northern Spain, univoltine (August–September) | nanula Nylander |
134 | Process of the labrum narrow, as long as broad. Scutum strongly shagreened and matt, densely punctate, punctures difficult to discern against the strong underlying sculpture. Clypeus sparsely punctate, with obscure and weak latitudinal striations | simontornyella Noskiewicz |
– | Process of the labrum wider, broader than long. Scutum less strongly shagreened, with punctures of variable density that are clearly visible against the underlying sculpture. Clypeus variably punctate, but never with latitudinal striations | 135 |
135 | Clypeus strongly flattened and sparsely punctate, punctures separated on average by more than 2 puncture diameters. Process of the labrum medially emarginate. Associated with Ornithogalum (Asparagaceae) | saxonica Stöckhert |
– | Clypeus domed, never strongly flattened, punctures dense or sparse. Process of the labrum truncate or apically rounded | 136 |
136 | Clypeus sparsely and obscurely punctate, punctures separated by 2–3 puncture diameters or more. Scutum very sparsely and finely punctate, punctures small, separated by 2–3 puncture diameters, underlying surface shagreened and dull. Restricted to temperate areas close to the Pyrenees | subopaca Nylander |
– | Clypeus clearly punctate, punctures separated by 1–2 puncture diameters. Scutum more densely punctate, punctures larger, separated by up to 2 puncture diameters, underlying surface dull to shiny. Throughout Iberia | 137 |
137 | Scutellum shagreened, at most weakly shiny. Scutum more densely punctate, punctures separated by 1 puncture diameter, underlying surface shagreened and dull (1st generation) to weakly shiny (2nd generation). Terga with dense and narrow apical hair bands laterally, individual hairs touching each other in fresh specimens (Fig. |
minutula (Kirby) |
– | Scutellum always polished and shiny. Scutum less densely punctate, punctures less regular, separated by 1–2 puncture diameters, underlying surface finely shagreened and weakly shiny (1st generation) to smooth and shiny (2nd generation). Terga with sparse apical hair bands, occasionally with some individual hairs touching each other (Fig. |
minutuloides Perkins |
138(91) | Clypeus flattened over majority of its surface. Species strongly associated with Fabaceae (subgenus Taeniandrena) |
139 |
– | Clypeus not noticeably flattened. Species not usually associated with Fabaceae | 151 |
138 | Face and foveae black haired. Terminal fringe dark brown (Fig. |
lusitania Wood & Ortiz-Sánchez |
– | Combination of characters otherwise; either face with pale hairs, terminal fringe light, or tergal densely punctate | 140 |
140 | Terminal fringe composed of dark brown to brownish grey hairs. Hairs flanking the basitibial plate of the hind tibia also dark (Fig. |
141 |
– | Terminal fringe and hairs flanking basitibial plate of hind tibia light, golden to yellow-orange (Fig. |
143 |
141 | Discs of T2–4 strongly and densely punctate, punctures clearly visible against the underling shagreenation. Widespread throughout Iberia, most common in areas with an Atlantic climate and abundant Genisteae. Bivoltine (typically March–May; June–July) | ovatula (Kirby) |
– | Discs of T2–4 obscurely and shallowly punctate, punctures disappearing into background shagreenation. Iberian distribution more restricted (see below) | 142 |
142 | Scutum with punctures shallow and obscure, separated by >1 puncture diameter. T3 with apical hair band interrupted medially. Currently known only from the steppe of central Spain (Guadalajara, Salamanca, Segovia). Univoltine (May–June) | ovata Schenck |
– | Scutum with punctures dense and clear, separated by <1 puncture diameter. T3 with apical hair band complete in fresh specimens. Currently only confirmed from the coast of southern Spain (Málaga). Bivoltine (probably February–April; May–June) | poupillieri Dours |
143 | Pygidial plate apically with deep emargination medially (Fig. |
lathyri Alfken |
– | Pygidial plate apically rounded, never emarginate. Clypeus without medial impression, punctures usually clear. Terga with or without punctures | 144 |
144 | Clypeus apically shiny. T2–4 apically with wide white hair bands that are complete in fresh specimens, these clearly exceeding the length of the marginal areas (Fig. |
gredana Warncke |
– | Clypeus uniformly dull. T2–4 with hair bands different, either shorter, broadly interrupted medially, or yellowish (Fig. |
145 |
145 | Declivity of T1 medially strongly and densely punctate, punctures separated by <1 puncture diameter | 146 |
– | Declivity of T1 with at most scattered punctures, never with dense patch of punctures medially | 149 |
146 | T3 with interrupted apical hair band in fresh specimens. T2–4 with the apical hyaline part of the marginal areas narrow, not exceeding 3 times the diameter of a puncture from the tergal discs. Scutum uniformly dull. Restricted to northern Portugal and Spain with isolated populations at elevation in the Sistema Central and Sistema Ibérico | wilkella (Kirby) |
– | T3 with apical hair band complete in fresh specimens (Fig. |
147 |
147 | Scutum shagreened laterally, medially with a circular area which is smooth and shiny; punctation here becoming weaker and sparser. Restricted to the Sistema Central to central and northern Portugal and north-western Spain (Zamora). Not known from the Cantabrian or Pyrenees Mountains | benoisti Wood & Praz |
– | Scutum more or less uniformly shagreened and punctate. Found across mountainous areas in eastern Spain (Sierra Nevada, Sierra de Cazorla, Sistema Ibérico) to the Pyrenees | 148 |
148 | Restricted to the alpine zone of the Sierra Nevada (above 2000 m) | contracta Wood |
– | Found elsewhere | intermedia Thomson aggregate (this likely represents a complex of an unknown number of species, potentially all of which are undescribed; the true intermedia may be absent from Spain) |
149 | Terga obscurely punctate, at least on the base of T2 with punctures disappearing into the underlying shagreenation. Univoltine (April–June) | russula Lepeletier sensu lato (including the distinct mitochondrial lineage from southern Portugal) |
– | Terga clearly punctate, at least on the base of T2 with punctures clearly visible against the underlying shagreenation | 150 |
150 | Terga strongly shagreened, dull to weakly shiny. Scutellum shagreened and weakly shiny. Tergal hair bands yellowish. Larger, 11–12 mm. Restricted to southern and south-eastern Spain (Almería, Granada, Málaga, Murcia, Valencia). Univoltine (April–June) | levante Wood & Praz |
– | Terga less strongly shagreened, shiny comparatively more strongly. Scutellum medially almost without sculpture, brightly shiny. Tergal hair bands whitish. Smaller, 8–10 mm. Throughout Iberia. Bivoltine (typically May–June; July–August) | afzeliella (Kirby) |
151 (138) | Fovea dorsally narrow, occupying at most ⅓ of space between lateral ocellus and compound eye, ventrally narrowing strongly (subgenus Euandrena) | 152 |
– | Foveae either dorsally broader, or not strongly narrowing | 162 |
152 | Head elongate, mouthparts extremely long, twice the length of the head (c.f. Fig. |
solenopalpa Benoist |
– | Head and mouthparts shorter, mouthparts never twice the length of the head. Process of the labrum trapezoidal | 153 |
153 | Clypeus densely and coarsely punctate with a raised longitudinal impunctate shiny line. Restricted to areas close to the Pyrenees. Associated with Symphytum (Boraginaceae) | symphyti Schmiedeknecht |
– | Clypeus densely or weakly punctate, never with a raised longitudinal impunctate shiny line | 154 |
154 | Pronotum with weak but distinct lateral keel. Clypeus medially with shallow longitudinal impression | 155 |
– | Pronotum laterally rounded. Clypeus without longitudinal impression, in one species (A. rufula Schmiedeknecht) with hints of an impression caused by impunctate longitudinal midline | 156 |
155 | T2 with marginal area long, occupying almost ½ the segment, strongly depressed and shiny (Fig. |
angustior (Kirby) |
– | T2 with marginal area shorter, not strongly depressed (Fig. |
lavandulae Pérez |
156 | Tergal discs with extremely coarse and dense punctures, punctures separated by 0.5–1 puncture diameter. Tergal margins strongly depressed and essentially impunctate, margins of T2–4 overlain by sparse whitish hair bands that emerge from the apexes of the tergal discs and which do not obscure the underlying surface of the marginal areas (Fig. |
fortipunctata Wood |
– | Tergal discs with normal punctures, not noticeably coarse, punctures separated on average by 1 puncture diameter. Tergal margins only weakly depressed, without long whitish hair bands, sometimes with obscure brownish lateral hair bands on the apexes of the marginal areas themselves | 157 |
157 | Apical margins of T1–4 extensively lightened yellow-hyaline, this lightened area clearly exceeding the diameter of a lateral ocellus (Fig. |
158 |
– | Apical margins of T1–4 not extensively lightened yellow-hyaline, at most narrowly so, this lightened area not exceeding the diameter of a lateral ocellus | 159 |
158 | Tergal margins comparatively more strongly depressed. Tergal discs on average more coarsely and densely punctate, punctures on the disc of T2 separated by 1.5–2 puncture diameters |
granulosa Pérez |
– | Tergal margins comparatively more weakly depressed. Tergal discs on average less strongly and densely punctate, punctures on the disc of T2 separated by 3–4 puncture diameters |
vulpecula Kriechbaumer |
159 | Facial pubescence predominantly light, with dark hairs restricted to the inner margins of the compound eyes. Mesepisternum entirely pale-haired. Restricted to temperate areas close to the Pyrenees | rufula Schmiedeknecht |
– | Facial pubescence predominantly black, with occasional scattered light hairs (Fig. |
160 |
160 | Face long, majority of the clypeus passing below an imaginary line drawn between the ventral margins of the compound eyes. Clypeus comparatively sparsely punctate, punctures separated by an average of 1 puncture diameter, with impunctate longitudinal midline. Restricted to mountainous areas in northern and north-western Spain | allosa Warncke (note, the Spanish taxon may be distinct from populations in Central Europe) |
– | Face short, only a small part of the clypeus falls below an imaginary line drawn between the ventral margins of the compound eyes. Clypeus comparatively densely punctate, punctures separated on average by 0.5–1 puncture diameter | 161 |
161 | Clypeus extremely densely and coarsely punctate, punctures separated by 0.5 puncture diameters (Fig. |
isolata sp. nov. |
– | Clypeus less densely punctate, punctures separated on average by 0.5–1 puncture diameters. Widespread across Iberia | bicolor Fabricius sensu lato (two mitochondrial lineages are present in A. bicolor; to date, only the southern lineage has been found in Iberia) |
162 (151) | Clypeus punctate, interspaces forming weakly raised longitudinal wrinkles (Fig. |
163 |
– | Clypeus without longitudinal wrinkles | 167 |
163 | Face, mesepisternum, scutum anteriorly, and propodeum with abundant white hairs; scutum medially with contrasting short black pubescence. Marginal areas of T2–4 laterally with broad white hair patches that obscure the underlying surface. Restricted to steppic areas in central Spain | soror Dours |
– | Face with pubescence variable, either black or a mixture of black and brown; if entirely white, then mesosoma with extensive brown pubescence dorsally. Terga with complete apical hair bands that are never interrupted medially in fresh specimens or with hair bands reduced and essentially absent | 164 |
164 | Facial pubescence white. Rare, restricted to the Pyrenees | gravida Imhoff |
– | Facial pubescence black or a mixture of black and brown; never pure white | 165 |
165 | Posterior face of hind femur with a latitudinal carina. Facial pubescence never entirely black, usually a mixture of yellow, brown, and black hairs, sometimes whitish. Terga usually with strong apical hair bands (Fig. |
flavipes Panzer |
– | Posterior face of hind femur rounded, without a latitudinal carina. Facial pubescence dark, often entirely black (Fig. |
166 |
166 | Propodeal triangle with finely raised rugae covering entire surface. Pubescence variable, from almost entirely melanic (Fig. |
vulcana Dours |
– | Propodeal triangle with finely raised rugae covering only basal half, apical parts with fine granular shagreenation. Pubescence variable, but never entirely melanic, mesonotum always with at least some brown hairs (Figs |
discors Erichson |
167 (162) | Dorsolateral surface of the propodeum reticulate, with large and shallow punctures (can be small in A. nuptialis Pérez), clearly contrasting the shagreened and shiny propodeal triangle, this lacking lateral carinae and becoming shinier on the declivity (Fig. |
168 |
– | Without this combination of characters, pronotum keeled or not | 173 |
168 | Hind tibiae and/or tarsi golden-orange | 169 |
– | Both hind tibiae and tarsi dark, at most obscurely reddish-brown | 170 |
169 | Hind tibiae golden-orange. Clypeus medially with a clearly raised longitudinal impunctate area, this area smooth and shiny, strongly contrasting the remaining parts of the clypeus which are densely punctate. Northern and central Spain, in areas with deciduous forest. Associated with Quercus (Fagaceae) | ferox Smith |
– | Hind tibiae dark. Clypeus with faint impunctate longitudinal line medially, but this is not raised and not shiny, therefore not strongly contrasting with the remaining parts of the clypeus which are regularly punctate. Rare, restricted to the Pyrenees | bucephala Stephens |
170 | A3 clearly exceeding length of A4+5 | nuptialis Pérez |
– | A3 not clearly exceeding length of A4+5, usually as long as or slightly shorter than A4+5 | 171 |
171 | Metasomal terga with short hairs, most clearly seen in profile on T2–3 with hairs not exceeding width of a flagellum | rosae Panzer (partim, dark form) |
– | Metasomal terga with extensive and abundant long hairs, most clearly seen in profile on T2–3 with hairs clearly exceeding width of a flagellum | 172 |
172 | Tibial scopa in fresh specimens usually dark dorsally and silver ventrally. Usually with light brown facial hair. Usually univoltine, flying April to mid-June, with potential sporadic emergence in August and September (not yet observed in Iberia). Rare, restricted to cooler parts of Iberia |
scotica Perkins |
– | Tibial scopa in fresh specimens dark dorsally and golden ventrally, but this can be ambiguous and fade to silver in older specimens or pinned material. Facial hair can be dark, particularly in the spring generation, the summer generation usually has lighter facial hair. Bivoltine, usually flying March–May and July–August, with phenology depending on local conditions. Common and widespread across Iberia |
trimmerana (Kirby) (partim, dark form) |
173 (167) | Pronotum laterally keeled, angulate, keel runs up dorsally to an angled corner | 174 |
– | Pronotum laterally rounded, without a keel | 205 |
174 | Mesepisternum deeply and distinctly punctate, punctures separated by <1 puncture diameter, with weakly shiny interspaces. Ocelloccipital distance exceeding 3 times the diameter of a lateral ocellus. Terga extremely densely punctate, punctures separated by <0.5 puncture diameters, with shiny interspaces. No recent records, possibly extinct, restricted to central Spain | incisa Eversmann |
– | Without this combination of characters, mesepisternum usually impunctate or only obscurely punctate, ocelloccipital distance shorter, terga dull or less densely punctate | 175 |
175 | Fore margin of clypeus slightly upturned, forming wide] shape, dorsolateral surface of propodeum reticulate, impunctate, not strongly differentiated from the propodeal triangle, terga with dense apical hair bands on T2–4, fovea broad and occupying over ½ of the distance between the compound eye and the lateral ocellus, tibial scopae with hairs long and loose. Restricted to central, southern, and eastern Spain and associated with Reseda (Resedaceae) | 176 |
– | Without this combination of characters | 177 |
176 | Terminal fringe orange. Tibial scopae orange. Tergal discs with extremely fine shagreen, shiny, clearly and deeply punctate | relata Warncke |
– | Terminal fringe dark brown medially and white laterally (Fig. |
blanda Pérez |
177 | Pygidial plate flat to weakly convex, outer margin usually somewhat elevated, without clearly raised area medially (Fig. |
178 |
– | Pygidial plate with distinctly limited raised area medially (Fig. |
186 |
178 | Scutum with very dense and even punctation across the entire surface of the disc, punctures separated by much less than the diameter of a puncture, in some cases nearly touching (Fig. |
griseobalteata Dours |
– | Scutum with moderate and uneven punctation, punctures separated by 1–3 puncture diameters in some cases. If occasional punctures are close to touching, then this is not consistently replicated across the whole scutum, with other punctures being separated by a clear distance. Underlying integument variable, from shagreened to shiny | 179 |
179 | Hind tibia and all tarsi completely orange. Restricted to areas close to the Pyrenees | chrysosceles (Kirby) |
– | Hind tibia black, tarsi may be black or orange | 180 |
180 | Larger species, body length 11–12 mm. T2–4 with thick white apical hair bands that obscure the underlying surface, only weakly interrupted medially on T2 | langadensis albipila Warncke |
– | Smaller species, body length 7–10 mm. T2–4 usually with weaker hair bands | 181 |
181 | Tibial scopa, when viewed in reverse profile (i.e. looking at the posterior face of the hind tibia) dorsally short and thick, hair length only 1–1.5 times the diameter of a lateral ocellus, clearly shorter than the ventral scopal hairs. Restricted to areas close to the Pyrenees | pallitarsis Pérez |
– | Tibial scopa equally long dorsally and ventrally | 182 |
182 | Terga very sparsely punctate, punctures of discs of T2–4 shallow, obscure, and separated by 3–4 puncture diameters. Clypeus comparatively flattened. Very rare, known only from southern Spain (Cádiz) | microthorax Pérez |
– | Terga densely punctate, punctures on discs of T2–4 strong and deep, separated at most by 2 puncture diameters, usually by <2 puncture diameters. Clypeus comparatively domed | 183 |
183 | Vertex clearly elongate, the hind ocelli separated from the margin of the vertex by a distance greater than the diameter of a lateral ocellus (Fig. |
foeniculae Wood |
– | Vertex not elongate, the hind ocelli separated from the margin of the vertex by a distance clearly less than the diameter of a lateral ocellus (Fig. |
184 |
184 | Central line in the front half of the scutum strongly impressed (Fig. |
nitidiuscula Schenck |
– | Central line in the front half of the scutum only weakly and superficially impressed (Fig. |
185 |
185 | Disc of T1 densely punctate, punctures separated by 1 puncture diameter. Scutellum shagreened and dull. Hind basitarsi orange. Throughout Iberia. Bivoltine (March–August) | fulvicornis Schenck |
– | Disc of T1 sparsely punctate, punctures separated by 2–3 puncture diameters. Scutellum polished and shiny. Restricted to strongly saline habitats in southern Iberia. Voltinism unclear, possibly bivoltine, recorded March–early June | juliana Wood |
186 | Flying exclusively in the summer (July–September). Hind tibiae triangular, clearly much broader apically than basally, with short scopal hairs not greatly exceeding the diameter of a lateral ocellus (Fig. |
187 |
– | Flying predominantly in the spring, some species extending into July. Hind tibiae normal, not greatly broader apically than basally, with long scopal hairs, clearly greatly exceeding the diameter of a lateral ocellus. Terga with or without clear apical hair bands | 189 |
187 | Outer surface of the galea smooth and shiny. Associated with Ericaceae, found in Atlantic habitats across northern, central, and western Iberia | fuscipes (Kirby) |
– | Outer surface of the galea shagreened and dull | 188 |
188 | Mesosoma anteriorly and posteriorly with pale hairs, medially with abundant black hairs. Face with pale hairs. Associated with Asteraceae, restricted to northern Spain | denticulata (Kirby) |
– | Mesosoma with pale to brown hairs, with at most occasional intermixed black hairs. Face black-haired. Polylectic, though often found on Asteraceae. Restricted to areas surrounding the Pyrenees and the Cantabrian Mountains with isolated populations in high mountains in southern Spain (particularly the Sierra Nevada) | nigriceps (Kirby, 1802) |
189 | Process of the labrum either elongate (as long as or slightly longer than broad) or pointed triangular, never medially emarginate (Fig. |
190 |
– | Process of the labrum trapezoidal, always broader than long, usually medially emarginate, at least weakly. Clypeus never with transverse wrinkles (subgenus Andrena s. str.) | 197 |
190 | Fovea very wide, occupying the entirety of the space between the inner margin of compound eye and the lateral ocellus | 191 |
– | Fovea narrower, occupying between half and two-thirds of distance between the inner margin of compound eye and the lateral ocellus | 192 |
191 | Clypeus with network of fine, raised ridges that extend laterally across the disc, underlying surface shiny. Process of labrum short, broader than long, produced to a fine triangular point (Fig. |
tunetana Schmiedeknecht |
– | Clypeus without any lateral ridges, underlying surface shagreened, dull. Process of labrum short and slightly broader than long, but clearly trapezoidal, apical margin slightly thickened and raised (Fig. |
leptopyga Pérez |
192 | Scutum and scutellum shagreened, dull. Process of labrum as long as broad, evenly rounded apically (Fig. |
barbilabris (Kirby) |
– | At least some part of the scutum or scutellum shiny. Process of labrum either pointed triangular or apically truncate, not evenly rounded apically | 193 |
193 | Process of labrum pointed triangular (Fig. |
sericata Imhoff |
– | Process of labrum apically truncate, not pointed (Fig. |
194 |
194 | Terga clearly finely and densely punctured, punctures separated by 1–2 puncture diameters, visible against weakly shagreened integument. Restricted to mountainous areas of central and northern Spain | argentata Smith |
– | Terga with punctures sparse and difficult to see, separated by 3–5 puncture diameters | 195 |
195 | Process of labrum comparatively large, apex more rounded (Fig. |
parviceps Kriechbaumer |
– | Process of labrum comparatively small, more clearly truncate (Fig. |
196 |
196 | Scutum shagreened laterally, but centrally shagreenation is absent, underlying surface therefore smooth and shiny. Lateral faces of the propodeum with clearly raised more or less parallel ridges. Scutum more strongly punctured, individual punctures larger and closer together, particularly anteriorly where they are separated by 1–2 puncture diameters | dinizi Warncke |
– | Scutum shagreened, shagreenation weaker centrally but still clearly visible. Lateral faces of the propodeum with at most very small wrinkles. Scutum less strongly punctured, individual punctures normal and more scattered, separated by 1–3 puncture diameters | ventralis Imhoff |
197 | Terga densely covered with long hairs, in fresh specimens these obscuring the underlying surface (Fig. |
198 |
– | Terga less thickly haired, sometimes with hair tufts on T1–2, but these not obscuring the underlying surface (Fig. |
199 |
198 | Hind tibiae orange. Terga predominantly black-haired, sometimes with light brown hairs on T1. Restricted to northern Spain, associated with Salix (Salicaceae) | clarkella (Kirby) |
– | Hind tibiae dark. Terga 1–5 with extensive reddish-orange pubescence (Fig. |
fulva (Müller) |
199 | Marginal areas of T2–4 very wide, occupying ¾ of each segment | synadelpha Perkins |
– | Marginal areas of T2–4 never occupying more than ½ of each segment, usually covering only ⅓ | 200 |
200 | Face with entirely pale hairs, at most with a few scattered dark hairs along the inner margin of the compound eye | 201 |
– | Face with extensive black hairs, particularly around the antennal insertions and along the inner margin of the compound eye | 203 |
201 | Terga sparsely haired, with at most weak hair tufts on T1–2. Terga finely shagreened and thus weakly shiny. Restricted to northern Spain. Flying later in the year (June–August), associated with shrubs, particularly Rubus (Rosaceae) | fucata Smith |
– | Terga typically with long hairs, in fresh specimens with clear and dense hair tufts on T1–2. Abraded or older specimens may lack such hair tufts, in which case use the following characters: terga strongly shagreened, dull. Flying earlier in the year (March–May), associated with flowering trees | 202 |
202 | Basitarsi of hind tibiae parallel-sided, not converging apically. Restricted to the Pyrenees with an isolated population in the Sierra de Cazorla (Jáen). Associated with various flowering trees and shrubs | helvola (Linnaeus) |
– | Basitarsi of hind tibiae broader basally, narrower apically, therefore converging apically. Restricted to areas around the Pyrenees and the Cantabrian Mountains. Associated with Salix (Salicaceae) | mitis Schmiedeknecht |
203 | Face entirely black haired, without any pale hairs. Terga basally (T1–2) with orange-brown hairs, apically (T3–4) with extensive and strongly contrasting black hairs. Associated with Vaccinium (Ericaceae) | lapponica Zetterstedt |
– | Face with at least some pale hairs, particularly around the antennal insertions. Terga without extensive areas with black hairs, generally with mixture of predominantly yellowish to brownish hairs on T1–4. Species associated with Salix (Salicaceae) | 204 |
204 | Smaller, 10–11 mm. Clypeus predominantly shagreened and dull, with only the narrow longitudinal impunctate midline weakly shiny. Terminal fringe dark brown | praecox (Scopoli) |
– | Larger, 11–14 mm. Clypeus comparatively shinier, only laterally shagreened and dull, apico-medially broadly shiny around the comparatively broader longitudinal impunctate midline. Terminal fringe black | apicata Smith |
205 (173) | Large species (over 12 mm in length). With abundant black, brown, and/or white pubescence. Clypeus strongly domed. Ocelloccipital distance long, at least 2 times the diameter of a lateral ocellus (subgenus Melandrena partim) | 206 |
– | Without this combination of characters; remaining species | 212 |
206 | Tergal discs T1–3(4) with upstanding mixture of short pale whitish to light brown hairs extending over both disc and marginal areas. Tibial scopae orange-red, or bicoloured and dorsally dark, ventrally orange (dark form, active February-April). Terga sometimes with greasy-bronzy metallic reflections | nigroaenea (Kirby) |
– | Tergal discs never with this sort of pubescence, sometimes with white to brownish pubescence in basolateral corners of tergal discs, never extending onto marginal areas. Tibial scopae entirely black or a combination of black and white, never orange-red. Terga dark, without metallic reflections | 207 |
207 | Mesosoma with black and white hairs, never with brown hairs | 208 |
– | Mesosoma with black, brown, and/or pale hairs, never with only black and white hairs | 209 |
208 | T2–4 laterally with thick apical patches of white hairs, these strongly contrasting the black integument | albopunctata (Rossi) |
– | T2–4 laterally without white hair patches, entirely dark | morio Brullé |
209 | Tibial scopae bicoloured, black dorsally and white ventrally. Mesosoma dorsally with bright orange-brown pubescence, laterally with pale pubescence, never with black hairs on the mesepisternum. Univoltine, flying April-June. Restricted to temperate parts of northern Spain | nitida (Müller) |
– | Tibial scopae entirely black. Mesosoma dorsally usually with the pubescence darker, laterally with pubescence never pale, usually with abundant black hairs, at most with pubescence of the mesepisternum brown | 210 |
210 | Disc of T1 shagreened and barely punctate, punctures scattered and obscure against the underlying sculpture | assimilis Radoszkowski |
– | Disc of T1 polished and shiny, strongly punctate, punctures clearly visible against the underlying sculpture | 211 |
211 | Disc of T1 with punctures comparatively sparse, separated by 2 puncture diameters | thoracica (Kirby) |
– | Disc of T1 with punctures dense, punctures separated by up to 1 puncture diameter, often separated by less | limata Smith |
212 (205) | Propodeal corbicula complete, with both anterior and dorsal fringe, fringes composed of long, dense, and extremely plumose yellowish-brown hairs, these plumose hairs present also on the mesepisternum, the flocculus, and the femoral scopae (Fig. |
ramosa Wood |
– | Propodeal corbicula simple, composed of simple or weakly plumose hairs, but if plumose then these not extending onto the mesepisternum, flocculus, and femoral scopae | 213 |
213 | Process of the labrum large, as long as broad, apically rounded, ventral surface covered with latitudinal wrinkles (Fig. |
mucida Kriechbaumer (partim, 1st generation) |
– | Process of the labrum different, either wider than long, trapezoidal, or apically pointed, ventral surface never covered with latitudinal wrinkles. Foveae narrow or broad | 214 |
214 | Head short and broad, at least 1.3 times wider than long. Fovea short and broad, only slightly longer than wide. Small bees, 8–10 mm, with long white pubescence in fresh specimens. Associated with Erodium (Geraniaceae) | 215 |
– | Without this combination of characters, usually larger. Never associated with Erodium | 217 |
215 | Propodeum and mesepisternum entirely microreticulate, without strong or weak raised network of reticulation (Fig. |
melacana Warncke |
– | Propodeum and mesepisternum shiny or with granular microreticulation, overlain by strong or weak network of raised rugosity (Figs |
216 |
216 | Propodeum (including propodeal triangle) and mesepisternum with strongly produced but fine interlinked network of raised rugosity (Fig. |
erodiorum Wood & Ortiz-Sánchez |
– | Propodeum with fine granular shagreen, with weak network of raised rugosity, propodeal triangle slightly depressed, basal 2/3rds with raised longitudinal rugae (Fig. |
juliae sp. nov. |
217 | Terga shagreened and dull, impunctate or with obscure and scattered punctures (Figs |
218 |
– | Terga polished and shiny, at most finely shagreened, clearly and densely punctate | 227 |
218 | Propodeal triangle with internal surface covered with fine network of raised reticulation | 219 |
– | Propodeal triangle with internal surface smooth, entirely granularly shagreened, without raised reticulation (Fig. |
220 |
219 | Mid and hind basitarsi dark. Tergal discs in profile view with sparse and short dark hairs, apically with white hair bands. Throughout Iberia | hypopolia Schmiedeknecht |
– | Mid and hind basitarsi orange (Fig. |
ranunculorum Morawitz |
220 | Propodeum with comparatively few hairs, these largely restricted laterally to the dorsal fringe of the propodeal corbicula, propodeum dorsally with sparse pubescence (Fig. |
funerea Warncke |
– | Propodeum covered with long abundant pubescence, no meaningful difference between hairs of the dorsal fringe of the propodeal corbicula and those on the dorsal surface of the propodeum (Fig. |
221 |
221 | T2–4 with clear apical hair bands | 222 |
– | Terga without apical hair bands, hairs may be present on the tergal discs | 224 |
222 | Fovea relatively narrow, occupying ½ the space between the compound eye and the lateral ocellus. Process of the labrum triangular, slightly truncate. Smaller, not exceeding 10 mm in length. Associated with Brassicaceae | medeninensis donata Warncke |
– | Fovea broad, occupying ¾ of space between the compound eye and the lateral ocellus. Process of the labrum trapezoidal with a strong emargination medially. Larger, exceeding 12 mm in length. Associated with Cistus (Cistaceae) | 223 |
223 | Scopa bicoloured, black dorsally and orange ventrally (Fig. |
ghisbaini sp. nov. |
– | Scopa unicolourous orange (Fig. |
villipes Pérez |
224 | Mid and hind basitarsi entirely lightened orange | 225 |
– | Mid and hind basitarsi dark, at most obscurely dark reddish | 226 |
225 | Mesepisternum laterally with abundant dark hairs. Base of discs of T2–4 strongly depressed (depressed relative to the apical margin of the preceding tergum, often with a physical space between them), this depression therefore laterally exaggerating the gradulus along its inner margin. Tergal margins (and sometimes sides of terga when viewed laterally) sometimes lightened reddish, contrasting the dark discs. Rare, known only from southern Spain (Alicante, Cádiz) | minapalumboi Gribodo |
– | Mesepisternum laterally with entirely pale hairs. Base of T2–4 not noticeably depressed, therefore gradulus not particularly noticeable. Tergal margins always dark. Common throughout Iberia | ferrugineicrus Dours |
226 | Discs of T2–4 with abundant and extensive pale pubescence, in fresh specimens this pubescence forming distinct patches laterally. Marginal areas broad, on T3–4 occupying ²/₅ of the length of the segment. Clypeus shagreened and dull, with a narrow impunctate longitudinal mid line, only slightly shiny at extreme apex. Throughout Iberia | nigropilosa Warncke |
– | Discs of T2–4 with at most scattered and fine hairs, never forming patches. Marginal areas narrow, on T3–4 occupying at most 1/5th of the length of the segment. Clypeus less extensively shagreened and dull, becoming smooth and shiny in its apical half, with broader impunctate longitudinal mid line. Rare, known only from southern Spain (Córdoba) | varia Pérez |
227 | Terga without apical hair bands, or with obscure hair bands that are widely interrupted medially even in fresh specimens (Fig. |
228 |
– | Terga with clear apical hair bands, these complete at least on T3–4 (Fig. |
229 |
228 | Terga with widely interrupted apical hair bands, often abraded and absent. Terga coarsely and densely punctate, tergal margins strongly depressed, with punctures at most half the size of those on the tergal discs (Fig. |
macroptera Warncke |
– | Terga without apical hair bands. Terga extremely finely punctate, no major difference in the size of the punctures on the weakly depressed marginal areas compared to the discs (Fig. |
corax Warncke |
229 | Scutum sparsely punctate, punctures separated by 1–2 puncture diameters, laterally shagreened and dull, shagreenation weakening medially and on the scutellum, here shiny. Process of the labrum short and triangular (Fig. |
urdula Warncke |
– | Scutum and scutellum densely punctate, punctures separated by up to 1 puncture diameter, underlying surface uniformly finely shagreened and weakly shiny. Process of the labrum trapezoidal, margin truncate to weakly emarginate | 230 |
230 | Tergal discs extensively covered with extremely short hairs, forming a velvety pubescence in addition to denser and longer hairs on tergal margins, almost obscuring the underlying surface in fresh specimens (Fig. |
farinosa Pérez |
– | Tergal discs with scattered short orange hairs, not forming a velvety pubescence (Fig. |
murana Warncke |
Andrena (Chlorandrena) humilis Imhoff, 1832, female A posterior face of hind femur; Andrena (Rufandrena) orbitalis Morawitz, 1871, female B posterior face of hind femur H tibial scopa, profile view; Andrena (Lepidandrena) pandellei Pérez, 1895, female C scutum, profile view; Andrena (Chlorandrena) leucolippa Pérez, 1895, female D scutum, profile view E foveae, dorsal view; Andrena (Lepidandrena) curvungula Thomson, 1870, female F foveae, dorsal view; Andrena (Cryptandrena) ventricosa Dours, 1873, female G tibial scopa, profile view.
Andrena (Plastandrena) bimaculata (Kirby, 1802), female A hind tibial spurs, dorsal view D propodeum, dorsal view; Andrena (Holandrena) labialis (Kirby, 1802), female B hind tibial spurs, dorsal view C propodeum, dorsal view; Andrena (Trachandrena) haemorrhoa (Fabricius, 1781), female E propodeum, dorsal view; Andrena (Melandrena) nigroaenea (Kirby, 1802), female F propodeum, dorsal view; Andrena (Andrena) helvola (Linnaeus, 1758), female G propodeum, dorsal view; Andrena (Hoplandrena) scotica Perkins, 1916, female H propodeum, dorsal view.
Andrena (Simandrena) dorsata (Kirby, 1802), female A propodeal corbicula, lateral view D scutum, dorsolateral view; Andrena (Simandrena) rhypara Pérez, 1903, female B foveae, dorsal view; Andrena (Simandrena) antigana Pérez, 1895, female C fovea, dorsofrontal view; Andrena (Simandrena) propinqua Schenck, 1853, female E scutum, dorsolateral view; Andrena (Melandrena) bicolorata (Rossi, 1790), female F fovea, dorsofrontal view; Andrena (Notandrena) nigroviridula Dours, 1873, female G hind tibial spurs, dorsal view; Andrena (Notandrena) varuga Warncke, 1975, female H hind tibial spurs, dorsal view.
Andrena (Micrandrena) longibarbis Pérez, 1895, female A propodeum, dorsal view D clypeus, frontal view; Andrena (Micrandrena) spreta Pérez, 1895, female B propodeum, dorsal view; Andrena (Micrandrena) tenuistriata Pérez, 1895, female C propodeum, dorsal view; Andrena (Micrandrena) pandosa trigona Warncke, 1975, female E face, frontal view; Andrena (Graecandrena) verticalis Pérez, 1895, female F face, frontal view; Andrena (Graecandrena) nebularia Warncke, 1975, female G face, frontal view; Andrena (Parandrenella) taxana Warncke, 1975, female H T1, dorsal view.
Andrena (Micrandrena) spreta Pérez, 1895, female A terga, dorsal view; Andrena (Micrandrena) omnilaevis Wood, 2020, female B terga, dorsal view; Andrena (Micrandrena) anthrisci Blüthgen, 1925, female C terga, dorsal view; Andrena (Micrandrena) pauxilla Stöckhert, 1935, female D terga, dorsal view; Andrena (Micrandrena) bayona Warncke 1975, female E terga, dorsal view; Andrena (Micrandrena) minutula (Kirby, 1802), female F terga, dorsal view (gradulus indicated by white arrow); Andrena (Micrandrena) minutuloides Perkins, 1914, female G terga, dorsal view (gradulus indicated by white arrow); Andrena (Micrandrena) falsifica Perkins, 1915, female H terga, dorsal view.
Andrena (Taeniandrena) lusitania Wood & Ortiz-Sánchez, 2022, female A terga, dorsal view; Andrena (Taeniandrena) ovatula (Kirby, 1802), female B terminal fringe; Andrena (Taeniandrena) afzeliella (Kirby, 1802), female C terminal fringe; Andrena (Taeniandrena) lathyri Alfken, 1900, female D pygidial plate, posterior view; Andrena (Taeniandrena) gredana Warncke, 1975, female E terga, dorsal view; Andrena (Taeniandrena) benoisti Wood & Praz, 2021, female F terga, dorsal view.
Andrena (Euandrena) angustior (Kirby, 1802), female A terga, dorsal view; Andrena (Euandrena) lavandulae Pérez, 1902, female B terga, dorsal view; Andrena (Euandrena) fortipunctata Wood, 2021, female C terga, dorsal view; Andrena (Euandrena) granulosa Pérez, 1902, female D terga, dorsal view; Andrena (Melandrena) flavipes Panzer, 1799, female E head, frontal view F terga, dorsal view; Andrena (Melandrena) vulcana Dours, 1873, female G mesosoma, profile view (dark form) H mesosoma, profile view (light form).
Andrena (Notandrena) griseobalteata Dours, 1872, female A pygidial plate, posterior view; Andrena (Leucandrena) leptopyga Pérez, 1895, female B pygidial plate, posterior view; Andrena (Notandrena) foeniculae Wood, 2020, female C head, dorsal view; Andrena (Notandrena) nitidiuscula Schenck, 1853, female D head, dorsal view E scutum, dorsolateral view; Andrena (Notandrena) fulvicornis Schenck, 1861, female F scutum, dorsolateral view.
Andrena (Cnemidandrena) denticulata (Kirby, 1802), female A hind leg, profile view B terga, dorsal view; Andrena (Leucandrena) barbilabris (Kirby, 1802), female C process of the labrum, frontal view; Andrena (Leucandrena) tunetana Schmiedeknecht, 1900, female D process of the labrum, frontal view; Andrena (Leucandrena) sericata Imhoff, 1868, female E process of the labrum, frontal view; Andrena (Leucandrena) parviceps Kriechbaumer, 1873, female F process of the labrum, frontal view; Andrena (Leucandrena) ventralis Imhoff, 1832, female G process of the labrum, frontal view; Andrena (Leucandrena) leptopyga Pérez, 1895, female H process of the labrum, frontal view.
Andrena (Andrena) fulva (Müller, 1776), female A terga, dorsal view; Andrena (Andrena) helvola (Linnaeus, 1758), female B terga, dorsal view; Andrena (?Euandrena) ramosa Wood, 2022, female C propodeal corbicula, profile view; Andrena (Didonia) mucida Kriechbaumer, 1873, female D process of the labrum, ventral view; Andrena (Avandrena) erodiorum Wood & Ortiz-Sánchez, 2022, female E propodeum, dorsal view; Andrena (Avandrena) melacana Warncke, 1967, female F propodeum, dorsal view; Andrena (Nobandrena) funerea Warncke, 1975, female G propodeum, dorsal view; Andrena (Truncandrena) ferrugineicrus Dours, 1872, female H propodeum, dorsal view.
Andrena (incertae sedis) macroptera Warncke, 1974, female A terga, dorsal view; Andrena (incertae sedis) corax Warncke, 1975, female B terga, dorsal view; Andrena (incertae sedis) urdula Warncke, 1965, female C terga, dorsal view D face, frontal view; Andrena (Ovandrena) farinosa Pérez, 1895, female E terga, dorsal view; Andrena (incertae sedis) murana Warncke, 1975, female F terga, dorsal view.
For the male key, the following shortcuts can be used:
A. Clypeus at least partly yellow-marked go to 2
B. Propodeal triangle clearly defined by strongly raised carinae, internal surface rugose-areolate go to 52
C. Genital capsule distinctive, with clear 90° emargination in the outer parts of the gonostyli. The most common Andrena species in Iberia go to Andrena flavipes Panzer
D. Process of the labrum thickened and expanded, anteriorly projecting beyond the fore margin of the clypeus. Pronotum with strong humeral angle. Active in the summer (mid-June to September) (subgenus Cnemidandrena and remaining Margandrena) go to 68
E. Fore margin of clypeus upturned AND gena conspicuously broadened, wider than the width of the compound eye AND pronotum with a conspicuous humeral angle (remaining Notandrena) go to 72
F. Small species, body length of 5–7 mm (exceptionally 8 mm), terga always dark (remaining Aciandrena, Aenandrena partim, Avandrena, Cordandrena, Cryptandrena partim, Graecandrena, and almost all Micrandrena) go to 77
G. Mesepisternum and/or dorsolateral parts of the propodeum conspicuously punctate AND ocelloccipital distance at least 3 times the diameter of a lateral ocellus (incisa-group, Pruinosandrena) go to 122
H. Clypeus flattened over majority of its surface (subgenus Taeniandrena) go to 127
I. At least some tergal discs red-marked go to 140
J. Tergal discs with metallic blue reflections go to 145
K. Mandibles elongate, sickle-like, strongly crossing apically. Pronotum with strong humeral angle. Gena often expanded, broader than the width of a compound eye go to 148
L. Measured along ventral margin, A3 twice as long as A4 go to 168
M. Measured along ventral margin, A3 shorter than or as long as A4 go to 194
N. Remaining species; measured along ventral margin, A3 slightly longer than A4 go to 215
1 | Clypeus at least partly yellow-marked (Figs |
2 |
– | Clypeus entirely dark, never with yellow markings | 51 |
2 | Clypeus and lower paraocular areas with yellow markings; paraocular markings may be reduced to small spots (Fig. |
3 |
– | Yellow facial markings restricted to the clypeus, paraocular areas entirely dark (Figs |
15 |
3 | Propodeum almost entirely declivous, without clearly differentiated horizontal and vertical parts. Propodeum with dorsolateral parts adjacent to the propodeal triangle densely and deeply punctate, punctures separated by <0.5 puncture diameters (c.f. Fig. |
4 |
– | Propodeum with clearly differentiated horizontal and vertical parts. Propodeum with or without punctures, but never with punctures separated by <0.5 puncture diameters | 6 |
4 | Ocelloccipital distance 2.5–3 times the diameter of a lateral ocellus. Tergal discs occasionally red-marked | variabilis Smith |
– | Ocelloccipital distance <2 times the diameter of a lateral ocellus. Tergal discs always dark | 5 |
5 | S5 apically with extremely dense and pronounced latitudinal tuft of yellowish hairs. Terga with apical hair bands broadly interrupted. Typically flying earlier, May-June | labialis (Kirby) |
– | S5 apically without dense latitudinal hair tuft, at most with scattered hairs. Terga typically with clear and uninterrupted apical hair bands, though beware abraded specimens. Typically flying later, July-August | decipiens Schenck |
6 | Discs of T2–3 entirely and conspicuously red-marked | 7 |
– | Discs of T2–3 dark, at most with the apical margins lightened | 8 |
7 | Large species, 11–13 mm. Mandibles long, sickle-like, crossing in their apical third (Fig. |
schencki Morawitz |
– | Smaller species, never exceeding 9 mm. Mandibles normal, not strongly crossing apically. Head more or less round. Genital capsule with gonostyli apically narrow, strongly converging subapically (Fig. |
labiata Fabricius |
8 | Paraocular areas with markings narrow, running up the inner margin of the compound eye, clearly dorsally exceeding the antennal insertions (Fig. |
orbitalis Morawitz |
– | Paraocular areas with markings more or less rectangular or quadrangular, never narrow and running up the inner margin of the compound eye (Fig. |
9 |
9 | Fore margin of the clypeus clearly and strongly upturned. Pronotum with strong carinate humeral angle. Head broad, clearly broader than long | 10 |
– | Fore margin of the clypeus normal, not upturned. Pronotum without a strong humeral angle. Head variable, but not strongly broadened | 13 |
10 | Scutum strongly and densely punctate, punctures separated by 0.5–1 puncture diameter, underlying surface shagreened. A5 long, clearly exceeding the length of A4; A4 at most 0.6 times as long as A5. Larger, 9–10 mm | griseobalteata Dours |
– | Scutum either strongly shagreened and obscurely punctate, or with shiny interspaces, never strongly punctured and shagreened. A5 short, not noticeably longer than A4, both segments quadrate or subquadrate. Smaller, 8–9 mm | 11 |
11 | Gena posteriorly rounded, without carina. All tarsi and the apex of the hind tibia lightened orange. Scutum strongly shagreened and obscurely punctate. Flying April-June | chrysosceles (Kirby) |
– | Gena posteriorly with clear carina. Scutum with surface shiny between punctures | 12 |
12 | Hind tarsi dark. Genital capsule with penis valves comparatively narrow. Associated with saline soils in southern Iberia. Flying March-June | juliana Wood (partim, with yellow markings on lower paraocular areas) |
– | Hind tarsi lightened orange. Genital capsule with penis valves broad, occupying the majority of the space between the gonostyli. Restricted to the Pyrenees. Flying July-August | pallitarsis Pérez |
13 | Terga clearly and finely shagreened, dull to weakly shiny, with small and weak punctures. Facial markings yellow. Ocelloccipital distance short, equalling the diameter of a lateral ocellus. Restricted to central Spain | funerea Warncke |
– | Terga smooth and shiny between strongly pronounced punctures. Facial markings white. Ocelloccipital distance larger, at least 2 times the diameter of a lateral ocellus | 14 |
14 | Larger, exceeding 10 mm in length. Mandibles basally with white markings. Body with abundant white pubescence, with dense hairs on the clypeus and ventrally on the gena that can obscure the underlying surface in fresh individuals. Widespread across all of Iberia. Flying April–July | leucolippa Pérez |
– | Small species, 7–8 mm. Mandibles dark, without basal white markings (Fig. |
coitana (Kirby) |
15 | Discs of T2–3 clearly and extensively red-marked | 16 |
– | Tergal discs dark, at most with the apical margins lightened | 18 |
16 | Clypeus with yellow marking small, occupying only a small proportion of the clypeus medially, without internal black markings (Fig. |
sardoa Lepeletier |
– | Clypeus almost entirely pale, with two small black markings. Flying later in the year, from late May to August; restricted to temperate habitats across Spain, including mountains in eastern and south-eastern Spain Associated with scabious (former Dipsacaceae = Caprifoliaceae) | 17 |
17 | Pronotum with strong humeral angle. Fore margin of the clypeus laterally produced into two small but distinctly projecting points (Fig. |
marginata Fabricius |
– | Pronotum rounded. Fore margin of the clypeus normal, without lateral projections. Larger, 14–16 mm. Restricted to montane grasslands in northern and central Spain with isolated populations in the Sierra de Cazorla and Sierra Nevada | hattorfiana (Fabricius) (partim, light form) |
18 | Head, mesosoma, and/or metasoma with metallic green or blue reflections | 19 |
– | Body lacking metallic reflections | 24 |
19 | Yellow marking of clypeus very small, reduced to a narrow latitudinal strip at the fore margin of the clypeus (Fig. |
doursana Dufour |
– | Yellow or white marking of the clypeus always larger, covering the majority or the entirety of the clypeus (with exception of pair of small black dots) | 20 |
20 | Discs of T2–4 clearly and densely punctate with strong punctures, punctures separated by ≤1 puncture diameter | 21 |
– | Tergal discs with weaker and more scattered punctures, punctures either fine or separated by >1 puncture diameter | 22 |
21 | Pronotum rounded. Tergal margins not strongly depressed, more or less at the same level as the tergal discs. Gena normal, as wide as the width of the compound eye. Restricted to high elevation sites in the Pyrenees | viridescens Viereck |
– | Pronotum laterally carinate. Tergal margins strongly depressed, clearly lower than the level of the tergal discs. Gena broadened, clearly broader than the width of the compound eye. Found in central and southern Iberia | bellidis Pérez |
22 | Larger, 8–10 mm. Scutum laterally shagreened, medially with a large more or less circular smooth and shiny area, this almost impunctate | aerinifrons Dours |
– | Smaller, 6–7 mm. Scutum uniformly shagreened and punctate, without a smooth and shiny area | 23 |
23 | Inner hind tibial spurs strongly bent at their apexes. Body with subdued metallic reflections, almost without reflections | varuga Warncke |
– | Inner hind tibial spurs straight, parallel-sided. Body with clear metallic reflections | nigroviridula Dours |
24 | Fore margin of the clypeus upturned (Fig. |
25 |
– | Fore margin of the clypeus normal, not upturned. Pronotum with or without a humeral angle | 28 |
25 | Propodeum almost entirely declivous, without clearly differentiated horizontal and vertical parts. Propodeum with dorsolateral parts adjacent to the propodeal triangle densely and deeply punctate, punctures separated by <0.5 puncture diameters | flavilabris Schenck |
– | Propodeum with clearly differentiated horizontal and vertical parts. Propodeum at most obscurely punctate | 26 |
26 | Larger, 10 mm. Scutum and scutellum shagreened, without shiny areas | langadensis albipila Warncke |
– | Smaller, 6–7 mm. Scutum and scutellum at least medially with smooth and shiny areas | 27 |
27 | Gena posteriorly carinate. Surface of gena predominantly smooth and shiny. Associated with saline soils in southern Iberia | juliana Wood (partim, with yellow markings restricted to the clypeus) |
– | Gena posteriorly rounded, without carina. Surface of gena microreticulate and dull. Very rate, recorded only from the Cádiz region, habitat preference unknown | microthorax Pérez |
28 | Mandibles elongate, sickle-like, strongly crossing in their apical third (Fig. |
29 |
– | Without this combination of characters; mandibles either shorter, pronotum laterally rounded, or gena not exceeding width of head | 32 |
29 | Ocelloccipital distance short, <0.5 times the diameter of a lateral ocellus. Genital capsule with gonocoxal teeth apically strongly diverging, apexes pointed (Fig. |
tunetana Schmiedeknecht |
– | Ocelloccipital distance long, 1–2 times the diameter of a lateral ocellus. Genital capsule with gonocoxal teeth not strongly diverging, apexes truncate (Fig. |
30 |
30 | Typically larger, 9–10 mm. Genital capsule more elongate, gonocoxal teeth with apexes comparatively narrow, gonostyli not noticeably constricted medially (Fig. |
sericata Imhoff |
– | Typically smaller, 7–9 mm. Genital capsule more compact, gonocoxal teeth with apexes comparatively broad, gonostyli strongly constricted medially (Fig. |
31 |
31 | Lateral faces of the propodeum with a fine network of raised rugosity on top of the underlying granular microreticulation (Fig. |
dinizi Warncke |
– | Lateral faces of the propodeum with regular granular microreticulation (Fig. |
ventralis Imhoff |
32 | Small species, length not exceeding 8 mm | 33 |
– | Larger species, at least 9 mm in length | 38 |
33 | Propodeal triangle with smooth granular microreticulation, without raised rugosity. Very small species, usually not exceeding 6 mm (yellow-faced Aciandrena) | 34 |
– | Propodeal triangle with at least some raised rugosity, never with uniformly smooth microreticulation. Usually a little larger, 6–8 mm | 35 |
34 | Terga regularly and clearly punctate, punctures extending onto depressed tergal margins. Throughout Iberia | fulica Warncke (partim, light form with yellow clypeus) |
– | Terga obscurely punctate, punctures disappearing into underlying microreticulation, tergal margins more or less impunctate. Restricted to central and eastern Spain | vacella Warncke |
35 | Hind basitarsi entirely lightened orange. Frons with contrasting black and white pubescence, black on the frons and gena posteriorly, intermixing with white around the antennal insertions, becoming white on the clypeus and on the gena ventrally. Restricted to Potentilla-rich (Rosaceae) cool habitats in the Pyrenees. Flying July-August | tarsata Nylander |
– | Hind basitarsi dark. Face with uniformly bright white or yellowish pubescence. Not restricted to the Pyrenees, usually flying earlier in the year | 36 |
36 | Tergal discs covered with upstanding whitish pubescence, this forming dense apical hair bands on tergal margins in fresh individuals. Scutum anteriorly shagreened, becoming smooth and shiny posteriorly. Antennae ventrally strongly and extensively lightened orange. Restricted to south-western Spain (Huelva, Sevilla), flying March-April | laurivora Warncke |
– | Tergal discs never with upstanding pubescence; white apical hair bands can be present on tergal margins. Scutum with uniform sculpture, consistently shiny across its surface. Antennae typically dark ventrally | 37 |
37 | Disc of T1 with scattered punctures, punctures separated by 1–3 puncture diameters, strongly contrasting punctation of the discs of T2–4, here punctures separated by 0.5 puncture diameters. Clypeus yellow-marked across its entire surface with the exception of two small black marks. Genital capsule compact, with pronounced gonocoxal teeth and small and spatulate gonostyli (Fig. |
taxana Warncke |
– | All tergal discs uniformly punctate, punctures separated by 0.5 puncture diameters. Clypeal marking often reduced in size, not covering entire surface. Genital capsule elongate, with weakly pronounced gonocoxal teeth, gonostyli apically produced into triangular wedges (Fig. |
ventricosa Dours (partim, light form) |
38 | Mid and hind basitarsi entirely lightened orange (Fig. |
39 |
– | Mid and hind basitarsi dark | 41 |
39 | Ocelloccipital distance 3 times the diameter of a lateral ocellus. Terga very finely shagreened, more or less smooth and shiny. Tergal discs regularly and deeply punctate, punctures separated by 1–2 puncture diameters, punctures becoming smaller and continuing onto tergal margins. Genital capsule, see Fig. |
limbata dusmeti Warncke |
– | Ocelloccipital distance <2 times the diameter of a lateral ocellus. Terga strongly shagreened, at most weakly shiny, tergal discs with small and obscure punctures that disappear into the underlying structure, tergal margins more or less impunctate | 40 |
40 | T2–5 laterally with extremely strong gradulus emerging from under the preceding terga, tergal margins of T1–4 elevated, not in contact with basal parts of T2–5 (Fig. |
minapalumboi Gribodo |
– | T2–5 laterally with very weak gradulus, margins of T1–4 not elevated, in contact with basal parts of T2–5. Body with paler pubescence, white hairs on face, gena ventrally, and mesepisternum, with light brown hairs dorsally. Common throughout Iberia | ferrugineicrus Dours |
41 | Scutum and scutellum with dense network of raised longitudinal striations, most obvious on the scutellum (Fig. |
rhyssonota Pérez |
– | Scutum and scutellum without longitudinal striations. Tergal margins depressed or not, but not in combination with tergal bases depressed | 42 |
42 | Large species, 14–16 mm. Genital capsule distinctive, elongate with projecting and apically truncate gonocoxal teeth (Fig. |
hattorfiana (Fabricius) (partim, dark form) |
– | Smaller, sometimes reaching 14 mm, but usually 9–12 mm. Genital capsule different; if elongate, then gonocoxal teeth pointed, not truncate. Pubescence predominantly bright, never almost entirely black | 43 |
43 | Face with strongly contrasting black and white pubescence; clypeus medially with white hairs, inner margins of the compound eyes and area around antennal insertions and frons with abundant black hairs | 44 |
– | Face with bright pubescence, at most with occasional scattered dark hairs, never strongly contrasting light pubescence | 46 |
44 | A3 short, only slightly longer than A4. A4 rectangular, only slightly longer than broad, clearly shorter than A5. Antennal segments slightly bulging ventrally. Rare, known only from southern Spain (Córdoba) | varia Pérez |
– | A3 long, clearly longer than A4. A4 rectangular, clearly longer than broad, slightly shorter than A5. Antennal segments parallel-sided, without ventral bulges | 45 |
45 | Genital capsule with apical flattened part elongate, longer than broad, appearing triangular (Fig. |
ghisbaini sp. nov. |
– | Genital capsule with apical flattened part rounded, as long as broad (Fig. |
villipes Pérez |
46 | Propodeal triangle broad, internal surface with very fine granular reticulation, laterally and posteriorly weakly shiny, contrasting the dorsolateral parts of the propodeum (Fig. |
monilia Warncke |
– | Propodeal triangle with clearer structure of raised reticulation or rugosity, without weakly shiny margins. Face in frontal view with compound eyes normal, separated by a distance much greater than the diameter of an individual compound eye (Fig. |
47 |
47 | Terga finely shagreened, more or less smooth and shiny, regularly and deeply punctate, punctures separated by 1–2 puncture diameters, punctures extending onto the tergal margins. A3 comparatively short, only slightly exceeding length of A4. Nervulus antefurcal | 48 |
– | Tergal shagreened to microreticulate, dull to weakly shiny, obscurely or clearly punctate, but marginal areas impunctate. A3 comparatively long, clearly exceeding length of A4. Nervulus interstitial to weakly postfurcal | 49 |
48 | Genital capsule simple, with small gonocoxal teeth, gonostyli spatulate and more or less parallel-sided (Fig. |
relata Warncke |
– | Genital capsule complex, with inner margins progressively diverging, forming 90° inner angle. Gonostyli apically strongly flattened and broadened, more or less triangular shovel-like (Fig. |
murana Warncke |
49 | Disc of T1 with large ‘crater punctures’, punctures with distinctly raised margins. Terga shagreened and weakly shiny. Genital capsule with long, strongly produced gonocoxal teeth (Fig. |
humilis Imhoff |
– | Disc of T1 with small hair-bearing punctures, without raised, crater-like rims. Terga microreticulate and dull. Genital capsule with only weakly produced gonocoxal teeth | 50 |
50 | A3 very long, slightly exceeding the length of A4+5. A4 very short, broader than long. Genital capsule relatively compact, with flattened apical part clearly longer than broad | medeninensis donata Warncke |
– | A3 comparatively shorter, at most equalling the length of A4+5. A4 longer, slightly longer than broad. Genital capsule relatively elongate, with flattened apical part only slightly longer than broad (Fig. |
nigropilosa Warncke |
51 (1) | Propodeal triangle clearly defined by strongly raised carinae, internal surface rugose-areolate (Figs |
52 |
– | Propodeal triangle not strongly defined by lateral carinae with its internal surface rugose-areolate | 66 |
52 | Forewing with two submarginal cells. Clypeus with longitudinal striations | lagopus Latreille |
– | Forewing with three submarginal cells. Clypeus without longitudinal striations | 53 |
53 | Small species, 6–7 mm. Clypeus, scutum, and terga extremely densely and deeply punctate with small punctures, punctures almost confluent. Each side of T2 laterally with a small but deep and clearly defined fovea (Fig. |
54 |
– | Larger species, at least 8 mm in length, usually >10 mm. Body never so densely punctate, punctures always a little separated, never confluent. T2 with fovea obscure | 55 |
54 | T2 laterally with foveae narrow and relatively elongate (Fig. |
colletiformis Morawitz |
– | T2 laterally with foveae shorter and relatively broad (Fig. |
miegiella Dours |
55 | Pronotum laterally with strong humeral angle with well-developed carina. Fore margin of clypeus weakly upturned | fuscosa Erichson |
– | Pronotum laterally rounded. Fore margin of clypeus normal, not upturned | 56 |
56 | Hind basitarsi and majority of hind tibiae lightened orange. Body with integument uniformly dark. Pubescence bright, face and mesepisternum with light brown hairs, scutum and T6 with bright orange hairs. A3 clearly shorter than A4 | haemorrhoa (Fabricius) |
– | Without this combination of characters; either hind legs dark, body with darker pubescence, or A3 clearly longer than A4 | 57 |
57 | Terga with metallic reflections and at most superficial punctures AND genital capsule distinctive, with strongly produced gonocoxal teeth and gonostyli strongly reflexed, with long plumose hairs present on their outer margin (subgenus Suandrena) | 58 |
– | Terga without metallic reflections OR if with metallic reflections, then genital capsule otherwise, either lacking pronounced gonocoxal teeth or without plumose hairs on the outer margin of the gonostyli OR terga clearly and abundantly punctate (subgenus Plastandrena) | 60 |
58 | Genital capsule with penis valves grossly inflated, occupying entirety of space between the gonostyli, laterally produced into bulbous projections (Fig. |
gades Wood & Ortiz-Sánchez |
– | Genital capsule with penis valves narrower, essentially parallel-sided, not occupying entirety of space between the gonostyli, without lateral projections | 59 |
59 | A3 ventrally rounded, lacking any kind of triangular point (Fig. |
cyanomicans Pérez |
– | A3 ventrally produced into triangular point (Fig. |
suerinensis Friese |
60 | Genital capsule with gonocoxae apically rounded (Fig. |
61 |
– | Genital capsule with clearly produced gonocoxal teeth (Fig. |
62 |
61 | Terga with metallic blue reflections. Genital capsule with penis valves medially with and oval hyaline expansion, this breaking the profile of the penis valves themselves (Fig. |
agilissima (Scopoli) |
– | Terga dark, without metallic blue reflections. Genital capsule penis valves uniformly converging apically (Fig. |
afrensis Warncke |
62 | Terga usually extremely densely punctate, punctures almost confluent. Terga with weak metallic blue hints. Genital capsule, see Fig. |
asperrima Pérez |
– | Terga never so densely punctate, punctures never confluent, always separated by at least 0.5 puncture diameters. Terga never with metallic blue reflections. Genital capsule otherwise | 63 |
63 | Mesosoma with brown pubescence dorsally and laterally, at most with occasional black hairs | 64 |
– | Mesosoma with grey and black pubescence, always with at least some black hairs laterally on the mesepisternum | 65 |
64 | Genital capsule with penis valves basally broad (Fig. |
tibialis (Kirby) |
– | Genital capsule with penis valves basally narrow (Fig. |
bimaculata (Kirby) |
65 | Penis valves basally narrow, apical spatulate parts of gonostyli comparatively narrow (Fig. |
pilipes Fabricius |
– | Penis valves basally broad, apical spatulate parts of gonostyli comparatively broad (Fig. |
nigrospina Thomson |
66 (51) | Genital capsule distinctive, with more or less 90° emargination in the outer parts of the gonostyli (Fig. |
flavipes Panzer |
– | Genital capsule otherwise | 67 |
67 | Process of the labrum thickened and expanded, anteriorly projecting beyond the fore margin of the clypeus. Pronotum with strong humeral angle. Active in the summer (mid-June to September) (subgenus Cnemidandrena and remaining Margandrena) | 68 |
– | Without this combination of characters | 71 |
68 | T2–3 extensively red-marked. Clypeus with apical corners produced into distinct anteriorly projecting knobs (c.f. Fig. |
pellucens Pérez |
– | Terga dark, without red markings. Clypeus without apically projecting knobs | 69 |
69 | Outer surface of the galea smooth and shiny. Associated with Ericaceae, found in Atlantic habitats across northern, central, and western Iberia | fuscipes (Kirby) |
– | Outer surface of the galea shagreened and dull | 70 |
70 | Gena posteriorly produced into a strong winged carina. Scutum with abundant black hairs. Associated with Asteraceae, restricted to northern Spain | denticulata (Kirby) |
– | Gena posteriorly rounded. Scutum at most with occasional black hairs, usually entirely brown-haired. Polylectic, though often found on Asteraceae. Restricted to areas surrounding the Pyrenees with isolated populations in high mountains in southern Spain (particularly the Sierra Nevada) | nigriceps (Kirby, 1802) |
71 (67) | Fore margin of clypeus upturned AND gena conspicuously broadened, wider than the width of the compound eye AND pronotum with a conspicuous humeral angle (remaining Notandrena) | 72 |
– | Without this exact combination of characters. (Note, male members of the Ovandrena are very similar, but the gena is as broad as the width of the compound eye, not broader. They can be found at couplet 168; Andrena blanda is also similar, but the gena is only slightly wider than the width of the compound eye. It should be diagnosed by its genital capsule, go to couplet 184) | 76 |
72 | Mandibles long, sickle-shaped, strongly crossing apically. A3 long, exceeding length of A4+5. Clypeus covered with dense plumose hairs than obscure the underlying surface in fresh specimens. Clypeus sometimes with weak metallic reflections | 73 |
– | Mandibles normal, not sickle-shaped, not strongly crossing apically. A3 short, slightly shorter than A4+5. Clypeus without plumose hairs that obscure the underlying surface. Clypeus dark, without metallic reflections | 75 |
73 | Terga uniformly dark, at most with marginal areas lightened dark brown. A3 comparatively short, only moderately exceeding length of A4+5. Discs of T2–4 clearly and regularly punctate, punctures separated by 2 puncture diameters, underlying surface shagreened and weakly shiny | ranunculi Schmiedeknecht |
– | Terga red-marked, with at least marginal areas and lateral parts of T2–3 lightened red. A3 comparatively long, almost equalling or equalling length of A4+5+6. Terga shagreened, obscurely punctate, punctures disappearing into background sculpture | 74 |
74 | Clypeus polished and shiny, with scattered punctures. Mesonotum with shiny areas medially. A3 equalling A4+5+6 | binominata Smith |
– | Clypeus uniformly shagreened and dull. Scutum uniformly shagreened and dull. A3 slightly shorter than A4+5+6 | leucophaea Lepeletier |
75 | Terga with marginal areas of T2–4 puncture-free. A3 shorter than A4+5, never twice as long as A4. Basitarsi usually dark, sometimes apically lightened orange. Restricted to cooler parts of northern and western Iberia; univoltine, flying on July-August | nitidiuscula Schenck |
– | Terga with punctures of discs of T2–4 extending onto marginal areas. A3 almost as long as A4+5, typically twice as long as A4. Basitarsi usually entirely lightened orange. Widespread across Iberia; bivoltine, flying in April-May and July-August | fulvicornis Schenck |
76 (71) | Small species, body length of 5–7 mm, terga always dark, legs always dark (remaining Aciandrena, Aenandrena partim, Avandrena, Cordandrena partim, Cryptandrena partim, Graecandrena, and almost all Micrandrena) | 77 |
– | Larger species, at least 8 mm in length. Terga lightened red in some species, either on the discs or the margins. Hind tarsi or basitarsi lightened orange in some species | 121 |
77 | All tergal discs uniformly punctate, punctures separated by 0.5 puncture diameters, underlying surface weakly shiny. Genital capsule elongate, with weakly pronounced gonocoxal teeth, gonostyli apically produced into triangular wedges (Fig. |
ventricosa Dours (partim, dark form) |
– | Genital capsule otherwise; tergal punctation variable | 78 |
78 | A3 much shorter than A4, at most ½ the length (Fig. |
79 |
– | A3 as long as A4 or longer. Terga without metallic reflections | 81 |
79 | A3 extremely short relative to A4, at most ¹/₅ the length (Fig. |
vaulogeri Pérez (partim, small individuals) |
– | A3 not so short, around ½ the length of A4 (Fig. |
80 |
80 | T3–4 laterally almost impunctate, any obscure punctures disappear into the background sculpture | aeneiventris Morawitz |
– | T3–4 laterally with abundant clear punctures, these distinct against the background structure (Fig. |
hedikae Jäger |
81 | A3 long, equalling or exceeding length of A4+5. Face with abundant dark pubescence, sometimes intermixed with white hairs. Propodeal triangle narrow, with fine granular shagreenation over the majority of its area, sometimes with short, fine, and raised rugae basally (Fig. |
82 |
– | Without this combination of characters | 84 |
82 | Genital capsule relatively compact, gonostyli apically flattened, more or less triangular shovel-like, more or less as broad as long (Fig. |
melacana Warncke |
– | Genital capsule elongate, gonostyli with apical flattened part much longer than broad (Fig. |
83 |
83 | Viewed ventrally, process of S8 narrow, more or less the same width as the basal stem part (Fig. |
panurgina De Steffani |
– | Viewed ventrally, process of S8 large, clearly broader than the basal stem part (Fig. |
avara Warncke aggregate (potentially including multiple valid species) |
84 | Propodeal triangle smooth, not defined laterally by raised carinae, with internal surface lacking network of raised rugosity, at most with very short rugae at the base of the propodeal triangle (Fig. |
85 |
– | Propodeal triangle either strongly defined by raised carinae, or with internal surface with network of raised rugosity covering at least the basal half (specifically, take care with A. tenuistriata, for which the lateral parts of the propodeal triangle present granular shagreen) | 95 |
85 | Clypeus with clear longitudinal striations covering the entire surface. Restricted to sandy and usually coastal habitats in southern Iberia | orana Warncke |
– | Clypeus without striations, or with at most obscure striations at the base of the clypeus, never covering the entire surface | 86 |
86 | Clypeus weakly but distinctly domed, basally densely punctate, punctures separated by 0.5 puncture diameters, punctures becoming sparse apically, here separated by 2–3 puncture diameters; underlying surface smooth and shiny (Fig. |
pandosa trigona Warncke |
– | Without this combination of characters, clypeus usually flattened and/or shagreened and dull, or genital capsule otherwise | 87 |
87 | Terga strongly and clearly punctate, tergal margins strongly depressed, with punctures continuing onto marginal areas; underlying surface finely shagreened and weakly shiny. Very small species, not exceeding 6 mm | fulica Warncke (partim, dark form with black clypeus) |
– | Terga either impunctate (Fig. |
88 |
88 | Clypeus flattened, shagreened in basal half, finely shagreened in apical half, the two half therefore contrasting; surface with obscure and scattered punctures. Genital capsule elongate, with long and sharply pointed penis valves (Fig. |
verticalis Pérez |
– | Clypeus flattened or domed, but without this combination of characters. Genital capsule otherwise | 89 |
89 | Genital capsule unusual, gonocoxae with inner margins forming obtuse angle, without gonocoxal teeth, gonostyli flattened with square truncate apexes (Fig. |
90 |
– | Genital capsule otherwise | 91 |
90 | Genital capsule with gonostyli comparatively narrow, apically narrower than the basal width of the penis valves; penis valves occupying comparatively little of the space between the gonostyli (Fig. |
impunctata Pérez |
– | Genital capsule with gonostyli comparatively broad, apically as broad as the basal width of the penis valves; penis valves occupying majority of space between the gonostyli (Fig. |
nebularia Warncke |
91 | Tergal margins extensively lightened hyaline-yellow, T2–4 with dense and thick complete apical hair bands that occupy and slightly exceed the entire length of the margin, obscuring the underlying surface in fresh individuals. Small species, not exceeding 6 mm. Rare, known only from central Spain | montarca Warncke |
– | Tergal margins with weaker apical hair bands, tergal margins not so extensively lightened yellow. If in doubt, species exceeding 6 mm in length, or with clypeus flattened | 92 |
92 | Genital capsule with penis valves basally strongly swollen, blister-like, gonocoxae with well-developed gonocoxal teeth (Fig. |
93 |
– | Genital capsule with penis valves normal, not noticeably broadened (Fig. |
94 |
93 | A3 shorter than A4+5. Clypeus flattened, with slight metallic green-purple hints medially. Terga essentially impunctate. Widespread across Iberia | longibarbis Pérez |
– | A3 slightly longer than A4+5. Clypeus flattened, almost with slight longitudinal concavity medially, uniformly black. Terga obscurely punctate. Central and eastern Spain only | fria Warncke |
94 | Gena broad, clearly broader than the width of a compound eye (Fig. |
alma Warncke |
– | Gena normal, equalling the width of a compound eye. Scutum extremely obscurely punctate, punctures disappearing into background structure. Pronotum rounded. Presence and distribution in Iberia unclear | abjecta Pérez |
95 | Genital capsule distinctive, with strongly reflexed gonostyli (Fig. |
djelfensis Pérez |
– | Genital capsule otherwise | 96 |
96 | Genital capsule with penis valves clearly and strongly broadened basally, clearly bulbous in appearance (Fig. |
97 |
– | Genital capsule with penis valves not strongly broadened basally | 100 |
97 | Marginal area of T1 wide, strongly thickened, impunctate. Scutum with scattered punctures, punctures separated by 1–3 puncture diameters. Genital capsule with gonocoxal teeth weakly produced (Fig. |
falsifica Perkins |
– | Marginal area of T1 not strongly thickened, flat or slightly depressed. Genital capsule with gonocoxal teeth truncate (Fig. |
98 |
98 | Clypeus flattened, with slight longitudinal impression medially. Inner margin of the gonostyli upturned, diverging dorsally from outer margin, forming a slight but visible ‘kink’ (Fig. |
saxonica Stöckhert |
– | Clypeus less strongly flattened. Inner margin of the gonostyli parallel with the outer margin, without a visible ‘kink’ (Fig. |
99 |
99 | Restricted to areas close to the Pyrenees. A4 slightly longer than A6. Broadened penis valves comparatively longer |
strohmella Stöckhert |
– | Found in mountainous areas across southern and central Spain and northern Portugal. A4 as long as A6. Broadened penis valves comparatively shorter |
icterina Warncke |
100 | Genital capsule with outer margins of the gonostyli concave. Clypeus strongly shagreened and dull | simontornyella Noskiewicz |
– | Genital capsule with outer margins of the gonostyli essentially parallel with the inner margin | 101 |
101 | Clypeus flattened, basally with obscure but distinct longitudinal striations, these covering the basal quarter of the clypeus and are channelled laterally along its margins | 102 |
– | Clypeus typically domed, without longitudinal striations basally | 103 |
102 | Facial pubescence predominantly black. Scutum shagreened with faint and weak greasy metallic reflections. Tergal discs shagreened, weakly shiny. Tergal margins distinctly depressed. Widespread throughout Iberia | nitidula Pérez |
– | Facial pubescence predominantly pale, with scattered black hairs laterally. Scutum more strongly shagreened, matt. Tergal discs with strong microreticulation, dull. Tergal margins weakly depressed. Restricted to the extreme north-east of Spain | distinguenda Schenck |
103 | Genital capsule elongate with penis valves converging apically to form a tapering point (Fig. |
104 |
– | Genital capsule and propodeal triangle otherwise | 105 |
104 | Genital capsule with the inner margins of the gonostyli only weakly produced towards the penis valves; apical spatulate part of the gonostyli comparatively elongate (Fig. |
niveata Friese |
– | Genital capsule with the inner margins of the gonostyli strongly produced towards the penis valves; apical spatulate part of the gonostyli comparatively short (Fig. |
lecana Warncke |
105 | Tergal discs, at least on T1, smooth and shiny, at most with very fine shagreenation | 106 |
– | Tergal discs shagreened or microreticulate, not strongly shiny, most dull and matt | 113 |
106 | Forewing with first submarginal cross vein separated from the stigma by three times its own width (Fig. |
107 |
– | Forewing with first submarginal cross vein separated from the stigma by more or less its own width (Fig. |
108 |
107 | Clypeus medially shagreened. Stigma dark brown. Sterna shagreened with the exception of the marginal zones. Common and widespread across Iberia | nana (Kirby) |
– | Clypeus medially shiny. Stigma bright, yellowish to reddish. Sternal discs shiny. Very rare, restricted to the Pyrenees | floricola Eversmann |
108 | Scutum densely shagreened, with large shallow ‘crater-punctures’. In fresh specimens, T1–5 with broad white apical hair bands that exceed the length of the marginal areas. S2–5 with distinct white apical hair bands. Restricted to mountainous areas of central and northern Spain | argentata Smith (partim, small individuals) |
– | Scutum shiny or obscurely shiny, never densely shagreened, with normal punctures without raised rims. Terga and sterna without clear and dense complete hair bands | 109 |
109 | Dorsal part of gena completely smooth and shiny, without any microsculpture. Discs of T2–4 very densely punctate, punctures separated by 0.5 puncture diameters. Restricted to dry and steppic habitats in central Iberia including northern Portugal | bayona Warncke |
– | Gena uniformly shagreened, never with extensive smooth and shiny areas. Discs of T2–4 less densely punctate, punctures separated by at least 1 puncture diameter | 110 |
110 | Antennae extensively lightened orange ventrally. T1 with very fine and scattered punctation. Rare, restricted to northern Spain, univoltine (August–September) | nanula Nylander |
– | Antennae dark ventrally. T1 with clearer and denser punctation, punctures separated by 1–2 puncture diameters | 111 |
111 | Terga with punctation clearly extending onto the depressed marginal areas. Genital capsule, see Fig. |
fabrella Pérez |
– | Terga with marginal areas essentially impunctate, at most with occasional puncture. Genital capsule otherwise | 112 |
112 | Marginal areas of T2–4 strongly depressed, mirror-smooth, without a single puncture. A4 as long as broad, equalling A3 in length. Found in the Sistema Central to west and north-western Iberia. Associated with Sedum (Crassulaceae) | omnilaevis Wood |
– | Marginal areas of T2–4 comparatively weakly depressed, smooth and shiny, but with occasional punctures. A4 subquadrate, broader than long, shorter than A3 in length. Widespread in Iberia | alfkenella Perkins |
113 | A4 quadrate, as wide as long (Fig. |
semilaevis Pérez |
– | A4 subquadrate, broader than long. Tergal margins depressed or not | 114 |
114 | Propodeal triangle laterally poorly defined, without clear carinae, internal surface with raised rugosity covering only central part in the form of a triangle, lateral parts therefore with granular shagreenation (Fig. |
tenuistriata Pérez |
– | Without this exact combination of characters | 115 |
115 | Scutum strongly shagreened and dull, very obscurely punctate, punctures separated by 2–4 puncture diameters, underlying surface matt; punctures often disappear into background sculpture | 116 |
– | Scutum less strongly shagreened to shiny, with at least some punctures clearly visible against the background sculpture; punctures separated by 1–2 puncture diameters | 117 |
116 | Tergal discs finely shagreened and weakly shiny, with scattered punctures that are nevertheless distinct against the background sculpture, punctures separated by 1–3 puncture diameters. Tergal margins with finer shagreen, impunctate, thus contrasting the tergal discs. Restricted to temperate areas close to the Pyrenees | subopaca Nylander |
– | Tergal discs strongly shagreened and dull, with extremely obscure punctures that are hard to distinguish against the background sculpture, separated by 2–4 puncture diameters. Tergal margins with equally strong shagreen, not strongly contrasting the tergal discs. Iberian distribution unclear, but probably restricted to southern Spain (Cádiz, Málaga) | tiaretta Warncke |
117 | T1–3 with discs strongly and coarsely punctate, punctures clearly visible against the background sculpture (Fig. |
anthrisci Blüthgen |
– | T1–3 without strong and coarse punctures, at most with occasional punctures that typically disappear into the background sculpture. Tergal margins depressed or not, but not noticeably more finely sculptured or more brightly shiny than the tergal discs | 118 |
118 | T2–4 with tergal margins strongly depressed | 119 |
– | T2–4 with tergal margins at most weakly depressed | 120 |
119 | Scutum with scattered punctures, punctures separated by 1–3 puncture diameters. Mesopleuron finely microreticulate, punctate, punctures separated by >1 puncture diameter. Throughout Iberia, common |
spreta Pérez |
– | Scutum with denser punctures, punctures separated by 1–2 puncture diameters. Mesopleuron more coarsely microreticulate, punctate, punctures separated by <1 puncture diameter. Exact Iberian distribution unclear, but currently known only from eastern Spain, typically in mountainous areas |
pauxilla Stöckhert |
120 | Scutum densely punctate, punctures typically separated by 1 puncture diameter. Scutum and scutellum shagreened and dull (1st generation) to weakly shiny (2nd generation). Genital capsule, see Fig. |
minutula (Kirby) |
– | Scutum more sparsely and irregularly punctate, punctures separated by 1–3 puncture diameters. Scutum and scutellum finely shagreened and weakly shiny (1st generation) to polished and shiny (2nd generation). Genital capsule, see Fig. |
minutuloides Perkins |
121 (76) | Mesepisternum and/or dorsolateral parts of the propodeum conspicuously punctate AND ocelloccipital distance at least 3 times the diameter of a lateral ocellus (incisa-group, Pruinosandrena) | 122 |
– | Mesepisternum either impunctate or ocelloccipital distance of less than 3 times the diameter of a lateral ocellus | 126 |
122 | Dorsolateral parts of the propodeum impunctate, with dense network of raised rugosity | 123 |
– | Dorsolateral parts of the propodeum regularly punctate, without a dense network of raised rugosity | 124 |
123 | Face and mesosoma with a mixture of black and white pubescence. T2–4 laterally with widely separated patches of dense white pubescence on their apical margins that obscure the underlying surface. Terminal fringe black | lateralis Morawitz |
– | Face and mesosoma with light brown pubescence. T2–4 laterally without patches of white pubescence, at most with obscure whitish hair bands. Terminal fringe light brown | incisa Eversmann |
124 | A3 exceeding A4+5 in length | parata Warncke |
– | A3 not exceeding A4+5 | 125 |
125 | 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 |
126 (121) | Clypeus flattened over majority of its surface (subgenus Taeniandrena) |
127 |
– | Clypeus not noticeably flattened | 139 |
127 | Genital capsule with pronounced gonocoxal teeth, these clearly projecting anteriorly for a distance greater than the diameter of an antenna (Fig. |
128 |
– | Genital capsule with at most weakly projecting gonocoxal teeth (Fig. |
129 |
128 | Penis valves more or less parallel-sided (Fig. |
lusitania Wood & Ortiz-Sánchez |
– | Penis valves very broad basally, occupying almost entire space between gonostyli (Fig. |
lathyri Alfken |
129 | Genital capsule with penis valves narrow basally, broad medially, and narrowing apically (Fig. |
gredana Warncke |
– | Genital capsule different | 130 |
130 | Penis valves broadened and gonocoxae with their inner margins diverging from their base, gonocoxae therefore forming a broad triangular opening basally (Fig. |
131 |
– | Penis valves not broadened or gonocoxae with their inner margins more or less parallel, sometimes slightly diverging apically | 132 |
131 | Penis valves strongly broadened, occupying the majority of the space between the gonostyli (Fig. |
intermedia Thomson aggregate (this likely represents a complex of an unknown number of species, potentially all of which are undescribed; the true intermedia may be absent from Spain) |
– | Penis valves not so strongly broadened, occupying ½ the space between the gonostyli (Fig. |
levante Wood & Praz |
132 | Genital capsule with weakly produced gonocoxal teeth (Fig. |
contracta Wood |
– | Genital capsule different, species found elsewhere | 133 |
133 | A3 short relative to A4, approximately 0.6–0.7 times as long | 134 |
– | A3 long, more or less as long as A4, typically between 0.8–1.2 times as long | 136 |
134 | Clypeus densely covered with snow-white vestiture, in fresh specimens completely obscuring the underlying surface, particularly of the apical margin. Terga strongly shagreened, punctation obscure, T1 with declivity almost impunctate, any punctures sparse and obscure. Marginal areas of T2–4 almost impunctate. Currently known only from the steppe of central Spain (Guadalajara, Salamanca, Segovia). Univoltine (May–June) | ovata Schenck |
– | Clypeus without such pubescence, the surface of the clypeus is usually clearly visible through the sparser pubescence. Terga with stronger more distinct punctation, including declivity of T1. Marginal areas of T2–4 distinctly punctate, at least basally | 135 |
135 | Terga with weak apical hair bands, widely interrupted on T2–3, complete on T4 (Fig. |
wilkella (Kirby) |
– | Terga with strong and thick apical hair bands, medially interrupted on T2, complete on T3–4 (Fig. |
benoisti Wood & Praz |
136 | Hind tibiae and basitarsi lightened orange. Terga strongly shagreened, usually with punctures obscure and disappearing into underlying sculpture. A3 sometimes a little longer than A4. Larger, 10–12 mm. Univoltine (April–June) | russula Lepeletier sensu lato (including the distinct mitochondrial lineage from southern Portugal) |
– | Hind legs usually dark. Terga variable, often with abundant and distinct punctures. A3 variable. Smaller, 8–10 mm. Bivoltine (March-August) | 137 |
137 | A3 often shorter than A4 (Fig. |
poupillieri Dours |
– | Genital capsule with gonostyli apically truncate, not produced into points | 138 |
138 | A4 usually equal or subequal to A3 in length (Fig. |
afzeliella (Kirby) |
– | A4 often slightly longer than A3 (Fig. |
ovatula (Kirby) |
139 (126) | At least some tergal discs red-marked (Fig. |
140 |
– | Tergal discs never red-marked, at most with tergal margins lightened | 144 |
140 | A3 longer than A4 | 141 |
– | A3 much shorter than A4, at most ¼ the length | 143 |
141 | Head broad, inner margin of compound eyes diverging ventrally (Fig. |
florea Fabricius |
– | Without this combination of characters | 142 |
142 | Clypeus shagreened and matt. Mesonotum and disc of T1 strongly and densely punctate. Hind basitarsi lightened orange. Restricted to the Pyrenees, associated with Campanula (Campanulaceae) | rufizona Imhoff |
– | Clypeus smooth and shiny. Mesonotum and disc of T1 weakly punctate with scattered punctures. Hind basitarsi dark. More widespread across Spain | parviceps Kriechbaumer (partim, light form) |
143 | S8 apically truncate, lacking apical emargination. Mandibles always bidentate | rosae Panzer (partim, light form) |
– | S8 with clear apical emargination. Mandibles unidentate (1st generation) or bidentate (2nd generation) | trimmerana (Kirby) (partim, light form) |
144 (139) | Tergal discs with metallic blue reflections | 145 |
– | Tergal discs dark, without metallic reflections, or at most with greasy greenish or bronzy reflections laterally (if greenish reflections present on clypeus, scutum, and scutellum, go to couplet 182), never with blue metallic reflections | 147 |
145 | Tergal discs with abundant ‘crater punctures’ with raised rims. Pubescence of mesosoma and face light brown, never with abundant black and white hairs | nigroolivacea Dours |
– | Tergal discs with simple punctures, without noticeably raised rims. Pubescence of mesosoma with abundant black and white hairs, on the mesosoma forming a striped pattern; white anteriorly and posteriorly, medially with a uniform black strip | 146 |
146 | A3 0.6 times as long as A4. Facial hair shorter than the width of a compound eye. Antennal scape with intermixed black and white hairs. Bivoltine (April–May; July–August), restricted to mountains in northern Spain and the Pyrenees |
barbareae Panzer |
– | A3 as long or almost as long as A4. Facial hair exceeding the width of a compound eye. Antennal scape with uniformly white hairs. Univoltine (April–May), more widespread across northern Portugal and Spain |
cineraria (Linnaeus) |
147 (144) | Mandibles elongate, sickle-like, strongly crossing apically (Fig. |
148 |
– | Without this combination of characters; mandibles normal, not elongate and strongly crossing apically (Fig. |
167 |
148 | Genital capsule distinctive, gonostyli long and filiform, penis valves grossly expanded, occupying almost the entire space between the gonostyli (Fig. |
bucephala Stephens |
– | Genital capsule and S8 otherwise | 149 |
149 | A3 extremely short, at most ¼ the length of A4 | 150 |
– | A3 longer, never this short. If in doubt, then terga with well-defined and narrow light brown apical hair bands | 153 |
150 | S8 apically truncate, lacking apical emargination. Mandibles always bidentate | rosae Panzer (partim, dark form) |
– | S8 with clear apical emargination. Mandibles either unidentate or bidentate | 151 |
151 | Mandible unidentate, lacking an inner subapical tooth. Gena usually with a long spine. Flying only in the spring (March-May, depending on elevation) | trimmerana (Kirby) (partim, dark form, 1st generation) |
– | Mandible bidentate, with an inner subapical tooth. Gena usually without a spine, sometimes with a very short spine. Flying in the spring or the summer | 152 |
152 | Flying in the spring (usually April to mid-June). Facial pubescence long. Rare, restricted to cooler parts of Iberia | scotica Perkins |
– | Flying in the summer (usually mid-June to July). Facial pubescence short. Widespread throughout Iberia | trimmerana (Kirby) (partim, dark form, 2nd generation) |
153 | Mandible unidentate, without inner subapical tooth. Sometimes there may be a hint of a weakly formed or ancient subapical tooth; this state should be treated as unidentate. Head often grossly enlarged | 154 |
– | Mandible clearly bidentate, with a strong inner subapical tooth. Head not normally grossly enlarged | 156 |
154 | A3 slightly shorter than A4. Clypeus in apical third bulging, surface smooth and shiny with scattered punctures, punctures separated by 1–4 puncture diameters. Hind tibiae and tarsi usually lightened orange. Northern and central Spain, in areas with deciduous forest. Associated with Quercus (Fagaceae) | ferox Smith |
– | A3 clearly longer than A4. Clypeus even, without bulging apical part, underlying surface evenly shagreened and regularly punctate. Hind tibiae and tarsi dark | 155 |
155 | Clypeus densely punctate, punctures separated by 0.5–1 puncture diameter. Clypeus apically occasionally with very small pale marking medially. Tergal discs with scattered punctures, punctures separated by 2–3 puncture diameters, underlying surface smooth and shiny. Bivoltine, flying April-May and again July-August |
nuptialis Pérez |
– | Clypeus with sparser punctures, punctures separated by 1–2 puncture diameters. Tergal discs densely punctate, punctures separated by 1–2 puncture diameters, underlying surface shagreened, weakly shiny. Univoltine, typically flying March-June depending on altitude |
vetula Lepeletier |
156 | A3 distinctly shorter than A4. Genital capsule with gonocoxal teeth pronounced but strongly truncate apically, quadrangular (Fig. |
leptopyga Pérez |
– | A3 at least as long as A4, usually longer. Genital capsule either with less strongly pronounced gonocoxal teeth, or with gonocoxal teeth apically rounded. Terga with or without apical tergal hair bands | 157 |
157 | Marginal area of T2 broad, occupying ½ the length of the segment, strongly depressed, surface smooth and shiny (Fig. |
angustior (Kirby) |
– | Marginal area of T2 never this broad; at most weakly depressed, surface structurally not clearly differentiated from the disc, never smooth and shiny | 158 |
158 | Genital capsule with gonocoxae apically rounded, gonocoxal teeth very slightly protruding at most (Fig. |
lavandulae Pérez |
– | Genital capsule with more strongly projecting gonocoxal teeth, overall capsule more robustly built. Facial pubescence either black or pale, never mixed. Usually larger, 9–12 mm. Typically not found in areas with a Mediterranean climate, species of temperate deciduous forest (subgenus Andrena s. str.) | 159 |
159 | Mandible at its base with a clearly pronounced tooth | 160 |
– | Mandible at its base with either an angulation or straight | 165 |
160 | A3 1.8 times longer than A4. Hind tarsi lightened orange. With a patchy distribution from northern Portugal and Spain to the Pyrenees, with an isolated population in the Sierra de Cazorla (Jáen) | fulva (Müller) |
– | A3 at most 1.3 times longer than A4, usually more or less equal in length. Hind tarsi dark to reddish | 161 |
161 | Propodeum with majority of hairs black, at most with scattered pale hairs | 162 |
– | Propodeum with majority of hairs pale, at most with scattered dark hairs | 163 |
162 | S8 apically emarginate. Slightly smaller, 9–11 mm | praecox (Scopoli) |
– | S8 apically truncate. Slightly larger, 10–12 mm | apicata Smith |
163 | Basal mandibular tooth short. Apical margins of S2–4 with long loose hairs that do not form clear fringes, hairs longer than the length of the hind basitarsis. Restricted to northern Spain. Flying later in the year (June–August), associated with shrubs, particularly Rubus (Rosaceae) | fucata Smith |
– | Basal mandibular tooth long. Apical margins of S2–4 with dense fringes composed of short hairs, these hairs not exceeding the length of the hind basitarsis. Flying earlier in the year (March-June) | 164 |
164 | Hind tarsi reddish. Genital capsule more elongate, gonocoxal teeth comparatively weak. Restricted to areas around the Pyrenees and Cantabrian Mountains. Flying during March-May, associated with Salix (Salicaceae) | mitis Schmiedeknecht |
– | Hind tarsi dark. Genital capsule more compact, gonocoxal teeth strongly produced. More widespread across northern Iberia into northern Portugal. Flying during May-June, associated with Vaccinium (Ericaceae) | lapponica Zetterstedt |
165 | Mandible without any kind of angulation at its base. Mesosoma with bright reddish brown hairs, with some black hairs on the mesepisternum. Associated with Salix (Salicaceae) | clarkella (Kirby) |
– | Mandible at its base with an angulation. Mesosoma never with black hairs laterally. Associated with a wider range of flowering trees and shrubs | 166 |
166 | Mandible at its base with angulation forming a strong 90° angle. In fresh specimens, clypeus with golden hairs. Terga with marginal areas normal, marginal area of T3 occupying at most 30% of segment length. Restricted to the Pyrenees with an isolated population in the Sierra de Cazorla (Jáen) | helvola (Linnaeus) |
– | Mandible at its base with angulation rounded, forming an obtuse angle (c. 120°). In fresh specimens, clypeus with white hairs. Terga with marginal areas broad, marginal area of T3 occupying 60% of segment length. More widespread across central and northern Iberia into northern Portugal | synadelpha Perkins |
167 (147) | Measured along ventral margin, A3 twice as long as A4 | 168 |
– | Measured along ventral margin, A3 longer or shorter than A4, but never twice as long | 193 |
168 | S8 unique, elongate on narrow step, apical portion medially constricted and apically strongly emarginate (Fig. |
paucisquama Noskiewicz |
– | S8 different, not of this shape | 169 |
169 | Fore margin of the clypeus upturned. Propodeal triangle broad and well-defined by raised lateral carinae, margins extending almost to the lateral edges of the metanotum, internal surface covered in raised irregular carinae of a similar width, thus appearing regular and consistent. Pronotum with weak or strong humeral angle. Genital capsule, see Fig. |
170 |
– | Without this combination of characters | 171 |
170 | Larger, 9–10 mm. Tongue with outer surface of galea clearly punctate, punctures separated by 1–2 puncture diameters. Pronotum with humeral angle comparatively weak, and fore margin of the clypeus only weakly upturned. Sterna with weak and sparse fringes on apical margins. Tergal punctation comparatively larger and coarser. Genital capsule, see Fig. |
oviventris Pérez |
– | Smaller, 7–8 mm. Tongue with outer surface of galea more or less smooth and shiny, without obvious punctures. Pronotum with humeral angle comparatively strong, and fore margin of the clypeus more strongly upturned. Sterna with strong and dense fringes on apical margins. Tergal punctation comparatively fine. Genital capsule, see Fig. |
farinosa Pérez |
171 | Clypeus smooth and shiny over almost its entire surface, regularly punctate, punctures separated by 0.5–1 puncture diameters. Metasoma elongate, essentially parallel-sided, surface finely shagreened, more or less smooth and shiny, deeply punctate, punctures separated by 0.5–2 puncture diameters. Genital capsule with gonocoxal teeth apically diverging (Fig. |
alluaudi Benoist |
– | Without this combination of characters | 172 |
172 | Tergal margins distinctly depressed, medially occupying 40% of the visible segment (Fig. |
173 |
– | Tergal margins either not depressed, not impunctate, not lightened hyaline brown, or narrower | 174 |
173 | Tergal discs strongly humped, accentuating contrast with margins (Fig. |
fortipunctata Wood |
– | Tergal discs not strongly humped (Fig. |
hystrix Schmiedeknecht (partim) |
174 | Head elongate, mouthparts extremely long, twice the length of the head, labial palps alone equal the length of the head (Fig. |
solenopalpa Benoist |
– | Head shorter, never with the labial palps equalling the length of the head | 175 |
175 | Gonostyli with outer margin concave, penis valves with weak lateral hyaline extensions (Fig. |
symphyti Schmiedeknecht |
– | Without this combination of characters | 176 |
176 | A4–13 ventrally covered with shiny scales, contrasting A3 which is ventrally dull (c.f. Fig. |
fertoni Pérez |
– | Without this combination of characters | 177 |
177 | Face with pale hairs medially and black hairs along the inner margin of the compound eyes. Genital capsule with surface of the gonocoxae with distinctive latitudinal granular shagreen, this extending onto the basal parts of the gonostyli (Fig. |
178 |
– | Without this combination of characters (for species with contrasting black hairs along the inner margin of the compound eyes, see the next couplet) | 181 |
178 | Genital capsule with outer margins of gonostyli weakly emarginate (Fig. |
179 |
– | Genital capsule with outer margins of gonostyli more or less straight, without emargination (Figs |
180 |
179 | Penis valves comparatively broad (Fig. |
gravida Imhoff (partim) |
– | Penis valves comparatively narrow (Fig. |
soror Dours |
180 | Genital capsule with penis valves comparatively narrow (Fig. |
vulcana Dours |
– | Genital capsule with penis valves comparatively broad (Fig. |
discors Erichson |
181 | Face predominantly white-haired, with black hairs along inner margin of compound eyes. Genital capsule elongate, relatively featureless (Fig. |
182 |
– | Without this combination of characters | 183 |
182 | Tergal discs with uniformly pale pubescence (Fig. |
bicolorata (Rossi) |
– | Tergal discs with intermixed black and white pubescence | florentina Magretti |
183 | Terga densely and regularly punctate, punctures separated by up to 1 puncture diameter, underlying surface weakly shagreened to smooth and shiny. T2–4 with distinct white apical hair bands, often abraded and interrupted medially (remaining Lepidandrena) | 184 |
– | Terga shagreened, with large ‘crater punctures’ with raised rims. Tergal margins without apical hair bands (remaining Chlorandrena) | 186 |
184 | Genital capsule elongate, gonostyli extremely long, many times longer than wide (Fig. |
baetica Wood |
– | Genital capsule with gonostyli compact, not extremely narrow and elongate (Fig. |
185 |
185 | S8 with short hairs that do not noticeably project laterally. Tarsal segment 5 of the hind leg elongate and bent. Slightly larger, 10–11 mm | curvungula Thomson |
– | S8 with long, laterally projecting hairs. Tarsal segment 5 of the hind leg not noticeably bent. Slightly smaller, 9–10 mm | pandellei Pérez |
186 | Process of S8 large, with triangular-shaped lateral projections covered with projecting hair tufts (Fig. |
187 |
– | Process of S8 large or small, but more or less parallel-sided, without triangular lateral projections or hair tufts (Fig. |
189 |
187 | Ventral surface of S8 with long, ventrally projecting hairs, clearly visible in profile. Apex of S8 emarginate | senecionis Pérez |
– | Ventral surface of S8 without ventrally projecting hairs. Apex of S8 rounded, never medially emarginate | 188 |
188 | Apex of gonostyli with dense, deeply impressed punctuation, gonocoxal lobes strongly pronounced, apically pointed. Common throughout Iberia | rhenana Stöckhert |
– | Punctuation of gonostyli scattered, especially on the rim of the inner margin, gonocoxal lobes smaller, apexes broadly rounded. Rare, southern and south-eastern Spain only | curtivalvis Morice |
189 | Process of S8 relatively short, ventral surface apically glabrous, viewed ventrally with hairs covering an area more or less as broad as long (Fig. |
cinerea Brullé |
– | Process of S8 relatively long, ventral surface entirely covered with hairs, viewed ventrally with hairs covering an area clearly longer than broad (Fig. |
190 |
190 | Larger, 11–12 mm. S8 strongly broadened apically, here broader than the stem (Fig. |
191 |
– | Smaller, 9–10 mm. S8 parallel-sided along its entire length, not apically broadened (Fig. |
192 |
191 | Gonostyli with inner margins more or less evenly rounded (Fig. |
livens Pérez |
– | Gonostyli with inner margins strongly flattened and produced into a raised ridge (Fig. |
agnata Warncke |
192 | Clypeus strongly domed, underlying surface weakly shagreened laterally, smooth and shiny over the majority of its area; clypeus largely dark, with at most occasional hints of metallic green reflections. Terga with bases very weakly depressed. Found in dry to steppic areas in central Spain | elata Warncke |
– | Clypeus weakly domed, underlying surface shagreened, weakly shiny; metallic green reflections present. Terga with bases strongly depressed. Found in areas close to or on the coast in southern Portugal and Spain | abrupta Warncke |
193 (167) | Measured along ventral margin, A3 shorter than or as long as A4 | 194 |
– | Measured along ventral margin, A3 slightly longer than A4 | 215 |
194 | A3 extremely short relative to A4, at most ¹/₅ the length (Fig. |
vaulogeri Pérez (partim, normal sized individuals) |
– | Without this combination of characters | 195 |
195 | Fore margin of clypeus upturned. Gena slightly exceeding width of compound eye. T2–4 with dense white apical hair bands (Fig. |
blanda Pérez |
– | Without this combination of characters | 196 |
196 | A4–13 ventrally covered with shiny scales, contrasting A3 which is ventrally dull (Fig. |
197 |
– | A4–13 not noticeably different from A3, without shiny scales | 200 |
197 | Smaller, 7–8 mm. Tergal discs finely shagreened, more or less smooth and shiny, with deep and distinct punctures, punctures on disc of T1 separated by 1–2 puncture diameters, on discs of T2–4 separated by 0.5–1 puncture diameter. Tergal margins broadly lightened yellow-orange hyaline | 198 |
– | Larger, 10–11 mm. Tergal discs shagreened to coarsely microreticulate, at most weakly shiny, with fine regular punctures or coarse ‘crater punctures’. Tergal margins at most with their apical rims narrowly lightened hyaline brown | 199 |
198 | Clypeus, scutum, and scutellum laterally shagreened, medially smooth and shiny (Fig. |
fulvago (Christ) |
– | Clypeus, scutum, and scutellum microreticulate and dull. Widespread across Iberia | hesperia Smith |
199 | Terga shagreened, weakly shiny, with fine and dense punctation. Tergal margins comparatively weakly depressed, with narrow tight apical white hair bands. Hind basitarsi dark. Throughout Iberia | hypopolia Schmiedeknecht |
– | Terga strongly microreticulate, with coarse ‘crater punctures’. Tergal margins comparatively strongly depressed, with long loose white apical hair bands that exceed the length of the marginal areas. Hind basitarsi lightened orange. Restricted to high altitude sites (>1200 m) in the Pyrenees | ranunculorum Morawitz |
200 | Larger species, 12–14 mm, usually with abundant black, white, or brown pubescence, usually with entirely dark black facial hair. Ocelloccipital distance >2 times the diameter of a lateral ocellus. Terga always without apical hair bands, sometimes with lateral hair patches (Melandrena partim) | 201 |
– | Smaller species, 9–10 mm, usually with subdued pubescence, facial hairs often pale, though dark in A. lepida and A. propinqua. Ocelloccipital distance often shorter, <2 times the diameter of a lateral ocellus. Terga with or without hair bands | 207 |
201 | Face and mesepisternum with long white pubescence, mesosoma dorsally with bright light brown pubescence. Univoltine, flying April-June. Restricted to temperate parts of northern Spain |
nitida (Müller) |
– | Without this pattern of pubescence; face either with extensive dark pubescence, or mesepisternum with dark pubescence, or mesosoma dorsally with dark pubescence | 202 |
202 | Body with only black and white pubescence | 203 |
– | Body with at least some brown pubescence | 204 |
203 | Body typically more extensively dark haired, face and mesepisternum normally with entirely black hairs. T2–4 laterally with strongly contrasting dense patches of white pubescence |
albopunctata (Rossi) |
– | Body often with extensive pale hairs, face sometimes white-haired with black hairs laterally, sometimes entirely black-haired. Terga often entirely black haired, laterally without dense patches of white pubescence. In pale forms, if T2–4 have loose white hair fringes laterally, then mesepisternum always white-haired |
morio Brullé |
204 | Terga shagreened, obscurely punctate, with punctures disappearing into the underlying sculpture | assimilis Radoszkowski |
– | Terga at most weakly shagreened, sometimes smooth and shiny, at least weakly shiny. Discs of T2–5 clearly and usually densely punctate, punctures typically separated by 1 puncture diameter | 205 |
205 | Marginal areas of terga with narrow section of apical rim lightened hyaline-brown, tergal discs with weak bronzy reflections. Face with mixture of dark brown and black hairs | nigroaenea (Kirby) |
– | Marginal areas of terga with apical rim dark, not lightened, tergal discs dark, without weak bronzy reflections. Face with uniformly black hairs | 206 |
206 | Disc of T1 with dense punctures, punctures separated by up to 2 puncture diameters |
limata Smith |
– | Disc of T1 with more scattered punctures, punctures usually separated by over 3 puncture diameters |
thoracica (Fabricius) |
207 | A3 extremely short, at most 0.5 times the length of A4 |
208 |
– | A3 longer, at least 0.8 times as long as A4, often as long as A4 | 209 |
208 | Facial hair almost entirely black, with a few scattered light brown hairs around the antennal insertions. Clypeus densely punctate, punctures separated by 0.5 puncture diameters, underlying surface smooth and shiny. Widespread across Iberia | lepida Schenck |
– | Facial hair intermixed light and dark. Clypeus more sparsely and irregularly punctate, punctures separated by 1–2 puncture diameters, underlying surface microreticulate and dull. Very rare, known only from one specimen from Cádiz | rhypara Pérez |
209 | Genital capsule distinctive, elongate, basally narrowed (Fig. |
210 |
– | Genital capsule otherwise. Tergal discs with uniform punctation | 211 |
210 | Face with predominantly dark hairs, with some light hairs intermixed around the antennal insertions. Scutum polished, shiny. Hind tarsi dark. Common throughout Iberia | propinqua Schenck |
– | Face with entirely light hairs. Scutum shagreened and dull. Hind tarsi lightened orange. Restricted to temperate areas in northern Spain | dorsata (Kirby) |
211 | Face with entirely bright pubescence, without any dark hairs laterally. Terga entirely smooth and shiny, without shagreenation | combinata (Christ) |
– | Face with at least some dark hairs laterally. Terga often with shagreenation | 212 |
212 | Terga entirely shagreened and dull to weakly shiny at most. Clypeus with fine granular shagreen, relatively shallowly punctate, medially with weak impunctate longitudinal midline. Discs of T2–4 densely punctate, punctures separated by 0.5 puncture diameters | antigana Pérez |
– | Terga less strongly shagreened, weakly to strongly shiny. Structure of clypeus variable, from shagreened to smooth and shiny, more strongly and deeply punctate. Terga less densely punctate, punctures on discs of T2–4 separated by at least 1 puncture diameter | 213 |
213 | Mesepisternum and propodeum with abundant black-brown hairs; remaining pubescence whitish. Scutum comparatively less strongly shagreened, weakly shiny. Restricted to areas close to the Pyrenees | thomsonii Ducke |
– | Mesepisternum, dorsum of mesosoma, and propodeum with bright yellowish pubescence, at most with occasional scattered black hairs. Scutum comparatively more strongly shagreened, dull | 214 |
214 | Clypeus between the punctures smooth and shiny. Tergal discs without shagreen, smooth and shiny, clearly visible at the base of T2. More widespread across Iberia |
congruens Schmiedeknecht |
– | Clypeus between the punctures shagreened, at most weakly shiny. Tergal discs shagreened, weakly shiny, most clearly visible on the base of T2. Restricted to the Pyrenees and Cantabrian Mountains |
confinis Stöckhert |
215 (193) | Genital capsule unique within the Iberian fauna (Fig. |
polita Smith |
– | Genital capsule otherwise | 216 |
216 | Head elongate, only slightly wider than long. Clypeus shagreened and dull in its basal half, polished and shiny in its apical half (Fig. |
ramosa Wood |
– | Combination of characters otherwise | 217 |
217 | Clypeus domed, covered with dense network of coarse and strongly raised transverse wrinkles. Dorsolateral surface of propodeum with dense network of raised rugosity that is almost indistinguishable from the propodeal triangle, this network extending onto the lateral faces of the propodeum. Terga smooth and shiny with regular deep punctures, without a hint of shagreenation | ampla Warncke |
– | Without this combination of characters | 218 |
218 | Tergal margins distinctly depressed, depressions medially occupying 40% of the visible segment (Fig. |
hystrix Schmiedeknecht (partim) |
– | Without this combination of characters | 219 |
219 | Penis valves narrow, more or less parallel-sided along their length (Fig. |
220 |
– | Penis valves basally broadened, clearly wider basally than medially or apically (Fig. |
225 |
220 | Head relatively elongate. Process of the labrum large, slightly wider than long. Clypeus domed, with large irregular punctures, punctures separated by 0.5–2 puncture diameters, with unclear impunctate longitudinal midline. In fresh specimens, terga with clear unbroken apical hair fringes. Bivoltine, March-April and again in June-July | mucida Kriechbaumer |
– | Head less elongate. Process of the labrum smaller, more clearly wider than long. Clypeus more weakly domed, shiny to shagreened. Terga without apical hair fringes | 221 |
221 | Apical margins of T1–5 widely lightened hyaline-yellow (Fig. |
222 |
– | Tergal margins at most with apical margins narrowly lightened yellowish. Tergal discs comparatively flat. Species not strongly associated with Cistaceae | 223 |
222 | Base of terga and tergal margins comparatively strongly impressed, margins clearly separated from disc by a visible ‘step’ (Fig. |
granulosa Pérez |
– | Base of terga and tergal margins comparatively weakly impressed, margins not clearly separated from discs by a visible ‘step’, almost level with discs medially. Tergal discs less densely punctate, punctures separated by 3–4 puncture diameters |
vulpecula Kriechbaumer |
223 | Gonostyli apically truncate, outer margin forming an acute point. A3 only slightly longer than A4. Pubescence entirely light brown with exception of occasional dark hairs laterally on the face. Restricted to temperate areas close to the Pyrenees | rufula Schmiedeknecht |
– | Gonostyli with outer margin rounded, never forming an acute point. A3 visibly longer than A4 | 224 |
224 | Light hairs on dorsal parts of mesosoma, terga and sterna yellowish white in fresh specimens. Erect hairs on disc of T4 usually predominantly dark. Facial pubescence often entirely dark. Bivoltine, can be recorded between March and August. Widespread across Iberia |
bicolor Fabricius sensu lato (two mitochondrial lineages are present in A. bicolor; to date, only the southern lineage has been found in Iberia) |
– | Light hairs on dorsal parts of mesosoma, terga and sterna snow white, without yellowish hue even in very fresh specimens. Disc of T4 at most with a few isolated, erect dark hairs. Face always with some grey-white hairs medially, at least between and around antennal sockets. Univoltine, April-June. Restricted to mountainous areas in northern and north-western Spain |
allosa Warncke (note, the Spanish taxon may be distinct from populations in Central Europe) |
225 | Face medially with white hairs, laterally with clear line of black hairs along the inner margin of the clypeus. Genital capsule with outer margins of gonostyli weakly emarginate. Rare, restricted to the Pyrenees | gravida Imhoff (partim) |
– | Face entirely pale-haired, at most with scattered black hairs laterally. Genital capsule with outer margins of gonostyli straight, without emargination | 226 |
226 | Terga very weakly shagreened, more or less smooth and shiny. Tergal discs with extremely scattered punctures, punctures separated by 3–4 puncture diameters or more. Genital capsule, see Fig. |
parviceps Kriechbaumer (partim, dark form) |
– | Terga either more strongly microreticulate (at least on discs of T2–3) or clearly and densely punctate, punctures separated by 1–2 puncture diameters | 227 |
227 | Tergal discs finely microreticulate, with obscure and scattered punctures, punctures separated by 2–4 puncture diameters. Tergal margins with narrow broadly interrupted hair fringes apically | barbilabris (Kirby) |
– | Tergal discs at most finely shagreened, more or less smooth and shiny, clearly and regularly punctate, punctures separated by 1–2 puncture diameters. Tergal margins with narrow interrupted or broad uninterrupted hair bands apically | 228 |
228 | Tergal discs with punctures slightly sparser, separated by 2 puncture diameters. T2–4 apically with broad uninterrupted apical white hair bands, these exceeding the length of the tergal margins and obscuring the underlying surface. Ocelloccipital distance equals the diameter of a lateral ocellus. Genital capsule, see Fig. |
argentata Smith (partim, large individuals) |
– | Tergal discs with punctures slightly denser, separated by 1 puncture diameter. T2–4 apically with widely interrupted narrow lateral hair fringes. Ocelloccipital distance equals 2 times the diameter of a lateral ocellus. Genital capsule, see Fig. |
corax Warncke |
Andrena (Opandrena) schencki Morawitz, 1866, male A face, frontal view G genital capsule, dorsal view; Andrena (Oreomelissa) coitana (Kirby, 1802), male B face, frontal view; Andrena (Rufandrena) orbitalis Morawitz, 1871, male C face, frontal view; Andrena (Orandrena) monilia Warncke, 1975, male D face, frontal view; Andrena (Truncandrena) doursana Dufour, 1853, male E face, frontal view; Andrena (Lepidandrena) sardoa Lepeletier, 1841, male F face, frontal view; Andrena (Poecilandrena) labiata Fabricius, 1781, male H genital capsule, dorsal view.
Andrena (Margandrena) marginata Fabricius, 1776, male A apex of clypeus, frontal view; Andrena (Holandrena) flavilabris Schenck, 1874, male B apex of clypeus, frontolateral view; Andrena (Leucandrena) dinizi Warncke, 1975, male C face, frontal view F genital capsule, dorsal view G propodeum, profile view; Andrena (Leucandrena) tunetana Schmiedeknecht, 1900, male D genital capsule, dorsal view; Andrena (Leucandrena) sericata Imhoff, 1868, male E genital capsule, dorsal view; Andrena (Leucandrena) ventralis Imhoff, 1832, male H propodeum, profile view.
Andrena (Parandrenella) taxana Warncke, 1975, male A genital capsule, dorsal view; Andrena (Cryptandrena) ventricosa Dours, 1873, male B genital capsule, dorsal view; Andrena (Truncandrena) minapalumboi Gribodo, 1894, male C terga, dorsolateral view; Andrena (Chlorandrena) rhyssonota Pérez, 1895, male D scutum and scutellum, dorsal view; Andrena (Charitandrena) hattorfiana (Fabricius, 1775), male E genital capsule, dorsal view; Andrena (Orandrena) monilia Warncke, 1975, male F propodeum, dorsal view.
Andrena (Orandrena) monilia Warncke, 1975, male A genital capsule, dorsal view; Andrena (incertae sedis) murana Warncke, 1975, male B face, frontal view D genital capsule, dorsal view; Andrena (incertae sedis) relata Warncke, 1975, male C genital capsule, dorsal view; Andrena (Chlorandrena) humilis Imhoff, 1832, male E genital capsule, dorsal view; Andrena (Truncandrena) nigropilosa Warncke, 1967, male F genital capsule, dorsal view; Andrena (Suandrena) suerinensis Friese, 1884, male G propodeum, dorsal view; Andrena (Plastandrena) pilipes Fabricius, 1781, male H propodeum, dorsal view.
Andrena (Brachyandrena) colletiformis Morawitz, 1873, male A propodeum, dorsal view B T2, profile view; Andrena (Brachyandrena) miegiella Dours, 1873, male C T2, profile view; Andrena (Suandrena) gades Wood & Ortiz-Sánchez, 2022, male D genital capsule, dorsal view; Andrena (Suandrena) cyanomicans Pérez, 1895, male E antennae, frontal view F genital capsule, dorsal view; Andrena (Suandrena) suerinensis Friese, 1884, male G antennae, frontal view H genital capsule, dorsal view.
Andrena (Plastandrena) agilissima (Scopoli, 1770), male A genital capsule, dorsal view; Andrena (Plastandrena) afrensis Warncke, 1967, male B genital capsule, dorsal view; Andrena (Plastandrena) asperrima Pérez, 1895, male C genital capsule, dorsal view; Andrena (Plastandrena) tibialis (Kirby, 1802), male D genital capsule, dorsal view; Andrena (Plastandrena) bimaculata (Kirby, 1802), male E genital capsule, dorsal view; Andrena (Plastandrena) pilipes Fabricius, 1781, male F genital capsule, dorsal view; Andrena (Plastandrena) nigrospina Thomson, 1872, male G genital capsule, dorsal view; Andrena (Melandrena) flavipes Panzer, 1799, male H genital capsule, dorsal view.
Andrena (Avandrena) panurgina De Steffani, 1889, male A propodeal triangle C genital capsule, dorsal view E S8, ventral view; Andrena (Avandrena) melacana Warncke, 1967, male B genital capsule, dorsal view; Andrena (Avandrena) avara liturata Warncke, 1975, male D genital capsule, dorsal view F S8, ventral view.
Andrena (Micrandrena) pandosa trigona Warncke, 1975, male A face, frontal view B genital capsule, dorsal view; Andrena (Graecandrena) verticalis Pérez, 1895, male C genital capsule, dorsal view; Andrena (Graecandrena) impunctata Pérez, 1895, male D genital capsule, dorsal view; Andrena (Graecandrena) nebularia Warncke, 1975, male E genital capsule, dorsal view; Andrena (Micrandrena) longibarbis Pérez, 1895, male F genital capsule, dorsal view.
Andrena (Micrandrena) djelfensis Pérez, 1895, male A genital capsule, dorsal view; Andrena (Micrandrena) falsifica Perkins, 1915, male B genital capsule, dorsal view; Andrena (Micrandrena) saxonica Stöckhert, 1935, male C genital capsule, dorsal view; Andrena (Micrandrena) icterina Warncke, 1974, male D genital capsule, dorsal view; Andrena (Micrandrena) nana (Kirby, 1802), male E forewing, dorsal view F Andrena (Micrandrena) fabrella Pérez, 1903, male F forewing, dorsal view G genital capsule, dorsal view; Andrena (Micrandrena) semilaevis Pérez, 1903, male H antennae, frontal view.
Andrena (Micrandrena) tenuistriata Pérez, 1895, male A propodeum, dorsal view; Andrena (Micrandrena) anthrisci Blüthgen, 1925, male B terga, dorsal view; Andrena (Micrandrena) minutula (Kirby, 1802), male C genital capsule, dorsal view; Andrena (Micrandrena) minutuloides Perkins, 1914, male D genital capsule, dorsal view; Andrena (Pruinosandrena) nilotica Warncke, 1975, male E genital capsule, dorsal view F Andrena (Pruinosandrena) pruinosa Erichson, 1835, male F genital capsule, dorsal view.
Andrena (Taeniandrena) lusitania Wood & Ortiz-Sánchez, 2022, male A genital capsule, dorsal view; Andrena (Taeniandrena) lathyri Alfken, 1900, male B genital capsule, dorsal view; Andrena (Taeniandrena) gredana Warncke, 1975, male C genital capsule, dorsal view; Andrena (Taeniandrena) intermedia Thomson, 1870 aggregate, male D genital capsule, dorsal view; Andrena (Taeniandrena) levante Wood & Praz, 2021, male E genital capsule, dorsal view; Andrena (Taeniandrena) contracta Wood, 2022, male F genital capsule, dorsal view; Andrena (Taeniandrena) wilkella (Kirby, 1802), male G terga, dorsal view; Andrena (Taeniandrena) benoisti Wood & Praz, 2021, male H terga, dorsal view.
Andrena (Taeniandrena) afzeliella (Kirby, 1802), male A antennae, frontal view B genital capsule, dorsal view; Andrena (Taeniandrena) ovatula (Kirby, 1802), male C antennae, frontal view D genital capsule, dorsal view; Andrena (Taeniandrena) poupillieri Dours, 1872, male E antennae, frontal view F genital capsule, dorsal view.
Andrena (Andrena) helvola (Linnaeus, 1758), male A head, frontal view; Andrena (Chrysandrena) fulvago (Christ, 1791), male B head, frontal view; Andrena (Hoplandrena) bucephala Stephens, 1846, male C genital capsule, dorsal view; Andrena (Leucandrena) leptopyga Pérez, 1895, male D genital capsule, dorsal view E terga, dorsal view; Andrena (Euandrena) angustior (Kirby, 1802), male F tergal, dorsal view; Andrena (Euandrena) lavandulae Pérez, 1902, male G genital capsule, dorsal view; Andrena (Lepidandrena) paucisquama Noskiewicz, 1924, male H S8, dorsal view.
Andrena (incertae sedis) alluaudi Benoist, 1961, male A genital capsule, dorsal view; Andrena (Euandrena) fortipunctata Wood, 2021, male B terga, dorsal view; Andrena (Aenandrena) hystrix Schmiedeknecht, 1883, male C terga, dorsal view D Andrena (Euandrena) solenopalpa Benoist, 1945, male D face, frontal view.
Andrena (Euandrena) symphyti Schmiedeknecht, 1883, male A genital capsule, dorsal view; Andrena (Melandrena) gravida Imhoff, 1832, male B genital capsule, dorsal view; Andrena (Melandrena) soror Dours, 1872, male C genital capsule, dorsal view; Andrena (Melandrena) vulcana Dours, 1873, male D genital capsule, dorsal view; Andrena (Melandrena) bicolorata (Rossi, 1790), male E genital capsule, dorsal view; F terga, dorsal view; Andrena (Lepidandrena) baetica Wood, 2020, male G genital capsule, dorsal view; Andrena (Lepidandrena) pandellei Pérez, 1895, male H genital capsule, dorsal view.
Andrena (Chlorandrena) rhenana Stöckhert, 1930, male A S8, dorsal view; Andrena (Chlorandrena) cinerea Brullé, 1832, male B S8, dorsal view; Andrena (Chlorandrena) livens Pérez, 1895, male C S8, ventral view E genital capsule, dorsal view; Andrena (Chlorandrena) abrupta Warncke, 1967, male D S8, ventral view; Andrena (Chlorandrena) agnata Warncke, 1967, male F genital capsule, dorsal view.
Andrena (Simandrena) propinqua Schenck, 1853, male A genital capsule, dorsal view; Andrena (Ulandrena) polita Smith, 1847, male B genital capsule, dorsal view; Andrena (Euandrena) granulosa Pérez, 1902, male C genital capsule, dorsal view H terga, dorsal view; Andrena (Euandrena) bicolor Fabricius, 1775, male D genital capsule, dorsal view; Andrena (incertae sedis) corax Warncke, 1975, male E genital capsule, dorsal view; Andrena (Leucandrena) parviceps Kriechbaumer, 1873, male F genital capsule, dorsal view; Andrena (Leucandrena) argentata Smith, 1844, male G genital capsule, dorsal view.
At 228 species, the Iberian fauna is slightly larger than those of Greece (c. 220) and Israel (c. 220), though the fauna of Israel is likely to eventually be larger than that of Iberia following ongoing revisions (
There are a number of outstanding problems that remain to be resolved, in addition to those highlighted above. Members of the subgenus Avandrena Warncke remain difficult to interpret due to their morphological variation and rarity in collections. Andrena (Avandrena) avara Warncke, 1967 sensu stricto was described from Morocco and also occurs in southern Iberia. Two additional subspecies were described from Spain, A. avara gavia Warncke, 1974 (locus typicus Madrid in central Spain) and A. avara liturata Warncke, 1974 (locus typicus Sierra de Guadarrama in central Spain), as well as two additional subspecies from North Africa. These may all represent valid species; molecular revision of this subgenus across the West Palaearctic is necessary to define species boundaries, and also to establish whether members of the Avandrena that lack spines on the posterior face of the hind femur truly belong here (
A total of 33 of the 228 Andrena species recorded here are endemic to the peninsula (14.5%). Twenty-two of the species are true Spanish endemics (9.6% endemic), whereas no species are endemic to Portugal. Of these 33 species, 31 of them were described after 1967 in just a handful of publications by a very limited number of authors (
It is possible however to comment more broadly on the biogeography of Iberian bees and notable distributional patterns.
The most substantial change since
The elevated plateau of central Spain clearly hosts relictual Andrena faunal elements. This is most clearly seen for two subgenera, Nobandrena Warncke, 1968 and Parandrenella Popov, 1958. Nobandrena consists of 10 species from Central Europe to Central Asia (western limit Switzerland), with one species endemic to central Spain (A. funerea Warncke, 1975), predominantly in the provinces of Ávila, Madrid, Salamanca, and Segovia. Likewise, Parandrenella consists of nine species from eastern Central Europe to Central Asia and Pakistan (European western limit is Slovenia, with an additional species in north-western Africa from Morocco to Tunisia; see
More broadly, Iberia hosts several predominantly eastern species in steppic or dry areas, such as A. lateralis (Spain and Portugal, southern Balkans to the Central Asia), A. (Melandrena) soror Dours, 1872 (Spain, Morocco, Turkey), and A. urdula (Spain, Morocco, and Greece). Iberia has strong faunal links to Morocco, and particularly the Middle Atlas. Due to the nature of European mountain chains, outside of Iberia, raised steppic areas are rare or cover only very small areas. In Central Spain, Morocco, and Turkey, extensive raised areas (the mesetas of Old and New Castille) of steppic habitat can be found. This elevated steppe differs from the Great Eurasian Steppe that runs from the Pannonian basin (predominantly eastern Austria, Hungary, southern Slovakia, western Romania, northern Serbia) to Mongolia and northern China, and hosts a fauna that supports the same evolutionary lineages (e.g. Nobandrena and Parandrenella), but often contains different species, many of which are endemic. This link between Iberia and Morocco can be seen particularly strongly in the Middle Atlas. In addition to the finding of A. relata in the Middle Atlas (
The extraordinary nature of central Spain and its rich Andrena fauna is well-illustrated by both historical (particularly those of Dusmet that were revised by
Away from central Spain, there are more obvious links between southern Spain and the North African Andrena fauna.
Finally, the difference between the size of the Portuguese (128) and Spanish (228) Andrena faunas is large and notable. Though Portugal has a higher density of Andrena species due to its much smaller size, it hosts no endemic Andrena species compared to the 22 species endemic to mainland Spain. As highlighted above, the Portuguese Andrena fauna is so much smaller due to the almost complete absence of high northern mountains and their associated Euro-Siberian fauna (the Serra da Estrella reaches to 1,993 m but is isolated from the high Cantabrian Mountains of northern Spain), the lack of elevated steppe (limited to north-eastern Portugal around Almeida or the Douro valley), the lack of high mountains in the south (the Sierra de las Nieves reaches 1,919 m and the Sierra Nevada 3,479 m), and finally the absence of the very hot and dry Mediterranean habitat from Cádiz to Alicante and Valencia that hosts both North African species as well as restricted endemic species. When taken collectively, it is this enormous variety of habitats, isolated mountains, and Atlantic and Mediterranean influences that has shaped and generated the rich Iberian Andrena fauna, and continues to provide taxonomic surprises and ecological delights.
I am supported by an F.R.S.-FNRS fellowship (Chargé de recherches). This work was also supported by the projects SPRING - Strengthening Pollinator Recovery through Indicators and monitoring (EC DG ENV project Contract No: 09.02001/2021/847887/SER/ENV.D.2.) and ORBIT (Taxonomic resources for European bees (EC DG Env project Contract No 09.029901/2021/848268/SER/ENV.D.2). Support for open-access publication costs was generously provided by the Institut des Biosciences (UMONS). My thanks go to Joseph Monks (
Iberian Andrena species checklist
Data type: National checklist
Explanation note: Iberian Andrena species checklist, with national totals for Portugal and Spain, details on loci typici for species described from Iberia, and dietary niche classifications.