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Monograph
The genus Andrena Fabricius, 1775 in the Iberian Peninsula (Hymenoptera, Andrenidae)
expand article infoThomas J. Wood
‡ University of Mons, Mons, Belgium
Open Access

Abstract

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.

Keywords

Cryptic species, DNA barcoding, Iberian endemic species, solitary bees, taxonomy

Introduction

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 (Lhomme et al. 2020; Ortiz-Sánchez 2020; Orr et al. 2021). Within this large fauna, the most speciose element consists of bees from the genus Andrena which are abundant throughout the peninsula and which are a ubiquitous component of the spring pollinator community. This pattern is typical for the Holarctic, as Andrena are the second largest genus of bees globally with around 1,650 species following recent revisions (Ascher and Pickering 2020; Pisanty et al. 2022a; Wood and Monfared 2022) and are almost always the most speciose genus in regional faunas across this region. Considering that bees emerged in the early to mid-Cretaceous period 110–140 million years ago (Danforth et al. 2013; Branstetter et al. 2017; Murray et al. 2018), as a major bee genus Andrena is relatively young, arising around 25 million years ago (Pisanty et al. 2022b). Given the size of the genus, Andrena has speciated extremely quickly (Bossert et al. 2022), leading to enormous species richness and often extreme taxonomic difficulty in delineating and recognising species. Given the abundance and ubiquity of this genus in the Iberian fauna, it is desirable to have suitable taxonomic resources to enable study of this rich fauna. However, nothing resembling an accessible revision is currently available.

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. Erichson 1841; Lepeletier 1841; Dours 1872, 1873; Pérez 1895, 1902, 1903; Morice 1899; Schmiedeknecht 1900; Saunders 1908; Benoist 1961). Somewhat incredibly, before 2020, only six papers were published that described currently valid Andrena species with a locus typicus in Iberia: Erichson (1835, three species), Dours (1873, one species), Pérez (1895, six species), Pérez (1902, one species), Warncke (1967, 10 species), and Warncke (1975a, 15 species), although some other works described what are now synonymous names (e.g. Spinola 1843; Pérez 1903; Friese 1922). The fauna was therefore quite unstudied before the revisionary work of Warncke (1967, 1975a, 1976), and following these publications almost no additional taxonomic work was carried out on the Iberian fauna until very recently. This attention has come in the form of revisionary taxonomic work and faunal work that has provided new distributional records, demonstrated the presence of doubtful species in Iberia, and recorded new species for the Iberian Andrena fauna (Baldock et al. 2018; Álvarez Fidalgo et al. 2020, 2021a, 2021b, 2022a, 2022b; Wood et al. 2020a, 2021, 2022; Álvarez Fidalgo and Aguado Martín 2022; Wood 2022; Wood and Ortiz-Sánchez 2022).

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. Schwenninger 2015), and treatments on other regional faunas (e.g. Schmid-Egger and Scheuchl 1997; Amiet et al. 2010), as well as in the global revisionary catalogue of Gusenleitner and Schwarz (2002). However, digging through this mountain of information is time consuming and sometimes dispiriting and confusing, in part due to persistent problems with species concepts and taxonomic nomenclature. It is therefore the objective of this present paper to resolve most of the outstanding taxonomic problems with the Iberian Andrena fauna and to present a new synthesis of this information in the form of a comprehensive identification key. Due to the strong faunal links between North Africa and Iberia, it is necessary to revise parts of the North African Andrena fauna in parallel; a subsequent paper dealing with taxonomic revisions exclusive to the North African Andrena fauna will follow shortly. Therefore, several taxonomic changes will be made here that affect the North African fauna, although this is not the principal objective of the current work. Finally, novel ecological data are provided concerning the pollen foraging niche of unstudied Iberian or West Mediterranean Andrena species.

Methodology

Species concepts

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. Gusenleitner and Schwarz 2002). Warncke recognised species solely based on their morphology, and used subspecies to delineate geographical variation. Subsequent workers have treated many of these subspecies as valid species based on morphological, genetic, and ecological data (e.g. Gusenleitner and Schwarz 2002; Schmid-Egger 2005; Schwenninger 2015; Praz et al. 2019; Kratochwil 2021; Wood et al. 2021; McLaughlin et al. 2022; Pisanty et al. 2022a).

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. Williams et al. 2020; Rasmont et al. 2021). Use of a purely biological species concept in Andrena is premature in almost all cases as we have extremely limited understanding of their mating systems and pre- and post-zygotic reproductive barriers, with this only being estimated post hoc by quantifying realised gene flow (e.g. McLaughlin et al. 2022). Given that the study of Andrena taxonomy using genetics is still in its infancy, it is best to consider Andrena species to be evolutionarily independent lineages (de Queiroz 2007), and that through the application of integrative taxonomy (including genetic, morphological, and ecological data), one can arrive at a robust species concept (Schlick-Steiner et al. 2010).

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 Mayr (1963, see Rasmont et al. 2021) and drawing from the taxonomic heritage of Warncke’s West Palaearctic revisions. Where sufficient data are available, these subspecies are validated as species in their own right or synonymised. Where such data are not available or the results are ambiguous, subspecies are retained in order to indicate future avenues of taxonomic study.

Genetic sampling and analysis

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 (Ivanova et al. 2006). Specimens were sequenced following standardised high-throughput protocols. Both Lep1 and BeeCox1F1 primers were used (Hebert et al. 2004; Bleidorn and Henze 2021) to target the COI-5 region. All sequences are published on the Barcode of Life Database (BOLD) website under the public dataset “DS-ANDWMED”.

Phylogenetic trees were supplemented with additional published sequences (e.g. Schmidt et al. 2015) that were downloaded from Genbank and the Barcode of Life Data System. Trees were also significantly enriched with sequences produced by the Centro de Investigação em Biodiversidade e Recursos Genéticos (CIBIO, Portugal) as part of a collaboration to barcode the Portuguese bee fauna that will be published in the near future, and sequences will be available on BOLD. Sequences beginning with the code ‘IBIHM’ were generated by CIBIO. Sequences were aligned using MAFFT (Katoh and Standley 2013). Aligned sequences were analysed in Seaview (Gouy et al. 2010) using a maximum likelihood analysis which was run with 1000 bootstraps. Intra- and interspecific distances were calculated using MEGA-X (Kumar et al. 2018). Outgroups were chosen based on the subgeneric analysis of Pisanty et al. (2022b) in order to ensure that the selected outgroup species is part of a subgenus or species group more basal to those selected for focused study.

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. Toews and Brelsford 2012). Species delineation based solely on differences in mitochondrial DNA is therefore discouraged, with integrative taxonomic preferred (Padial et al. 2010; Gallego-García et al. 2023). More broadly, more powerful genetic techniques are now available such as Ultra Conserved Elements that allow the generation of sequences from thousands of loci. These have been applied to bee taxonomy at the species-level, and whilst they often produce results concordant with COI analysis (e.g. de Oliveira Andrade et al. 2022; McLaughlin et al. 2022), they can also clearly resolve problems for which COI analysis produces ambiguous or inconclusive results (e.g. Gueuning et al. 2020). However, to date, UCEs have predominantly been employed to resolve specific questions, and not to sample across a fauna in order to have the broadest possible taxonomic coverage, as was the objective in this study. Given these limitations, combined with the conceptual points discussed above, analysis of the COI fragment is used as an additional line of evidence to support taxonomic decision making, and is never used as the sole metric.

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 Pisanty et al. (2022b) who identified undescribed clades of Andrena. These clades were not discovered here through analysis of the COI fragment, and these illustrative phylogenetic trees should not be considered strong support for their existence.

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.

Checklist and identification

For updating the Iberian Andrena species total, the checklist of Ortiz-Sánchez (2020) for mainland Spain and Portugal is used as a baseline. As this list is mostly correct, it is not considered necessary to produce a full annotated list for the Iberia Andrena fauna here; instead, a detailed justification will be given for the changes which have been made since 2020 or which are newly made here. A full checklist is provided in Suppl. material 1.

Identification key and geographic scope

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 Schmid-Egger and Scheuchl (1997) and Amiet et al. (2010). Additional characters have also been integrated from published works such as Schönitzer et al. (1995), Schwenninger (2009, 2013, 2015), and Praz et al. (2019, 2022). It is not always easy to remember the precise source of specific characters used here that have been accumulated and integrated over several years of study, as many are scattered throughout the literature in original descriptions, some derive from my own observations, and many are taken from the important diagnostic characters that are embedded throughout Gusenleitner and Schwarz (2002), but which can be somewhat obscure and hidden in individual species accounts. It is my hope that this key can bring this information together as a novel synthesis, becoming a more easily digestible resource.

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 Schmid-Egger and Scheuchl (1997) and Amiet et al. (2010) can be used in conjunction with this Iberian key to cover the vast majority of the Andrena fauna of southern France. The Iberian key should not be used in an Italian context as the Italian fauna is already sufficiently distinct to render the key of limited use, either in northern Italy due to the presence of eastern faunal elements (e.g. Andrena (Aenandrena) bisulcata Morawitz, 1877) or in Sicily due to the large number of non-Iberian North African elements present there, and also because many Iberian taxa are absent from the Italian fauna which has a much lower degree of endemism. Finally, this key should not be used in Corsica or Sardinia, as their island faunas require dedicated study due to the presence of endemic and North African species, as well as local forms or subspecies.

Distribution maps

No distribution maps are presented as part of this work, as they are for other revisions such as that of Ortiz-Sánchez and Pauly (2017). This is because whilst the material that I have examined, validated, and digitised is sufficient to allow a more or less complete taxonomic understanding of the Iberian fauna, it is not sufficiently geographically comprehensive, and hence distribution maps would be incomplete at the scale of the peninsula. Moreover, the distribution maps of Warncke presented by Gusenleitner and Schwarz (2002) are sufficiently correct to be informative to students of the Iberian fauna, with the assumption that the updated taxonomy presented here and by other workers after 1993 is integrated. Important distributional information is given in the text where relevant, and also in the identification key, as many taxa (particularly Euro-Siberian taxa) are geographically limited and the place of capture can strongly inform their identification (e.g. species restricted in an Iberian context to the Pyrenees, to central or southern Spain, etc).

Dietary niches of Iberian Andrena species

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 Wood and Roberts (2018). Dietary classification (polylecty, mesolecty, oligolecty, narrow oligolecty) follows Müller and Kuhlmann (2008).

Morphological terminology

Morphological terminology follows Michener (2007). Specimens were measured from the centre of the clypeus at the front of the head to the apical tip of the metasoma to the nearest 0.5 mm. Photographs were taken using an Olympus E-M1 Mark II with a 60 mm macro lens. Additional close-ups were taken with the addition of a Mitutoyo M Plan Apo 5X infinity corrected objective lens. Photographs were stacked using Helicon Focus B (HeliconSoft, Ukraine) and plates were prepared in GNU Image Manipulation Program (GIMP) 2.10. Post-processing of some images was made in Photoshop Elements (Adobe Systems, USA) in order to improve lighting to highlight specific characters.

The following abbreviations are used in the species descriptions: A = antennal segments, S = metasomal sterna, and T = metasomal terga. Subgeneric concepts follow Pisanty et al. (2022b) with the necessary modifications detailed below. In diagnoses, the defining characters of a species are given, with those of the indicated comparison species given in parentheses.

Collections studied

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;

EBDC EBD-CSIC collection, Estación Biológica de Doñana, Seville, Spain;

MNHN Muséum national d’Histoire naturelle, Paris, France;

MRSN Museo Regionale di Scienze Naturali di Torino, Turin, Italy;

MSC Personal collection of Maximillian Schwarz, Ansfelden, Austria;

MZUR Zoological Museum of Sapienza University of Rome, Rome, Italy;

NHMUK Natural History Museum, London, United Kingdom;

OÖLM Oberösterreiches Landesmusum, Linz, Austria;

RMNH Naturalis Biodiversity Center, Leiden, the Netherlands;

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;

ZISP Zoological Institute of the Russian Academy of Sciences, St. Petersburg, Russia;

ZMHB Museum für Naturkunde, Berlin, Germany.

Results

Genetic results and taxonomic changes

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.

Subgenus Aciandrena Warncke, 1968

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 Wood et al. (2020b) as one of the distinguishing characters was that North African populations had males with the clypeus entirely black, and Iberian populations had males with the clypeus yellow-marked, but males with both colour forms can be found in both geographical regions. There are no clear structural differences in the tergal punctation or the structure of the male genital capsule. Genetically (Fig. 1), Iberian and Moroccan populations differ by an average genetic distance of 1.61% (range 1.31–2.09%). Although Iberian material (including specimens WPATW318-21 and WPATW354-21 which are black-faced males from the Sierra Nevada) forms a clade with bootstrap support of 100, the overall low genetic distance is not considered to be sufficient to justify species status. The synonymy of Wood et al. (2020b) is therefore maintained. The broad A. fulica clade as a whole is well-defined with bootstrap support of 98.

Figure 1. 

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).

Subgenus Aenandrena Warncke, 1968

This subgenus was found to be paraphyletic by Pisanty et al. (2022b), and the same result is found here with a COI-based analysis (Fig. 1). Andrena hystrix Schmiedeknecht, 1883 falls away from A. aeneiventris Morawitz, 1872 and A. hedikae Jäger, 1934. For convenience, these taxa are retained in the same subgenus, but future work will be needed to resolve this issue and probably to describe a new subgenus for the species around A. hystrix (currently three Palaearctic species from Morocco to Central Asia). Type material for A. hystrix was recently rediscovered, and a lectotype for this species is designated below.

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 (Gusenleitner and Schwarz 2002). Barcode analysis (Fig. 1) showed that A. aeneiventris specimens from Austria, Hungary, Israel, and Portugal formed a clade with bootstrap support of 99, and with low average intraspecific genetic distance of 0.78% (range 0.30–1.06%). In contrast, A. hedikae formed two clades. Five specimens from Portugal and Spain formed a clade with bootstrap support of 93 with low differentiation of 0.82% (range 0.16–2.01%). Two specimens from northern Morocco had identical sequences and formed a clade with bootstrap support of 98. These two clades were separated by an average genetic distance of 4.55% (range 4.34–4.72%), and both were well-separated from A. aeneiventris, by an average genetic distance of 5.86% (range 4.98–6.67%) for Iberian specimens and 5.20% (range 4.86–5.75%) for Moroccan specimens. These genetic distances are substantial, but no taxonomic action is taken here, as sequences from south-eastern Europe are needed to i) further understand barcode variation in A. hedikae and ii) identify which of these two clades matches populations from the locus typicus. Moreover, there are no apparent morphological differences between Iberian and Moroccan ‘hedikae’. Further study is required.

Subgenus Chlorandrena Pérez, 1890

This subgenus is clearly supported genetically (Pisanty et al. 2022b) and morphologically by the presence of a row of teeth on the posterior face of the female femur combined with strong ‘crater punctures’ on the terga. Barcodes generally supported all species concepts (Fig. 2), but there are some issues that must be discussed.

Figure 2. 

Phylogenetic tree (maximum likelihood) of Andrena subgenus Chlorandrena 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).

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 (Warncke 1967). Andrena agnata is much closer morphologically to A. livens than the nearest neighbour in the current tree, A. (Chlorandrena) nigroolivacea Dours, 1873; no taxonomic action is taken until such sequences are available, and a broad A. livens concept is maintained for now.

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. Warncke (1967) used a subspecies concept, with A. boyerella s. str. present in North Africa and A. boyerella leucolippa present in Iberia and France, A. leucolippa being described from south-western France, locus typicus Riscle (Fig. 3). It is important to note that Pérez writes in his catalogue under entry 442 that he had males from Algeria; in any case, the designation of the lectotype by Warncke (1967) from France fixes this name unambiguously on European populations. North African and Iberian specimens differ, with A. leucolippa presenting denser tergal punctation, in the male sex the yellow colouration of the clypes extends onto the mandibles and the lower paraocular areas (only the clypeus is yellow-marked in A. boyerella), and there are slight differences in the genital capsule. Gusenleitner and Schwarz (2002) listed A. leucolippa as a distinct species.

Figure 3. 

Andrena (Chlorandrena) leucolippa Pérez, 1895 female lectotype A label details B profile C face, frontal view D dorsal view.

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. Dours (1872) described A. boyerella, and there is no doubt over the morphological identity of this taxon based on the clear description. However, Dours gives a distribution of southern France and Algeria. It is therefore the case that, if there are two different species, then Dours’ type series was polytypic. Warncke (1967) and Gusenleitner and Schwarz (2002) give the locus typicus of A. boyerella as Algeria, but since there is no holotype or designated lectotype, the locus typicus is undefined. Genetically, sequences from four North African specimens from the Middle Atlas in Morocco were identical and formed a clade (Fig. 2) that was strongly separated from two sequences from Estorãos near Fafe in northern Portugal [WPATW078-21] and Balaguer in Catalonia in north-eastern Spain [WPATW111-21] by an average genetic distance of 13.27% (range 13.27–13.27%). The two clades are not sisters, being separated by A. (Chlorandrena) abrupta Warncke, 1967. The matter would appear to be clear, that two taxa are present. However, a specimen from Vieira do Minho in northern Portugal [IBIHM500-21] diverged from both these lineages, being separated from the Moroccan sequences by 8.95% and the two other Iberian sequences by 14.51%. This site is only 16 km from the Estorãos site. Morphologically, there are no obvious differences; all three Iberian specimens are males. Because A. leucolippa was described from south-western France, the sequence from north-eastern Spain is here assumed to be representative of true A. leucolippa. For now, the divergent Portuguese sequence is considered to be aberrant until it can be independently replicated. It is possible that this ‘aberrant’ sequence is a NUMT (nuclear sequences of mitochondrial origin) and represents amplification of nuclear DNA rather than mitochondrial. Since only a single sequence is available, and it is not possible to determine the nature of this sequence, no action is taken. It is not impossible that a third undescribed species is present, but without additional genetic data no further action is taken. Morphology supports the existence of only two species, and in order to fix the name A. boyerella on North African populations, a neotype is designated below from barcoded Moroccan material.

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. 2), though intraspecific variation was high at an average of 4.59% (range 0.00–8.33%). Specimens from Iberian mountain chains (Sistema Central, Sierra de Baza, Sierra Nevada) were the most strongly divergent, differing by 8.02–8.33% from specimens from Austria, Belgium, and Germany. However, multiple clades were found without clear geographic structuring. Therefore, a broad species concept is maintained here, and this overall A. humilis clade has bootstrap support of 98. In Iberia, A. humilis forms a species pair with A. (Chlorandrena) cinerea Brullé, 1832. Andrena cinerea specimens from Morocco, Portugal, and Spain formed a distinct clade with low intraspecific variation of 0.77% (range 0.00–1.54%). This clade was strongly separated from the broad A. humilis clade by an average distance of 15.05% (range 13.89–15.74%). However, a single specimen from Balaguer in north-eastern Spain which was originally identified as A. cinerea diverged from both the broad A. humilis clade by 11.70% (range 11.11–12.65%) and A. cinerea by 12.35% (range 12.35–12.35%). As for the potentially aberrant specimen of A. leucolippa, it is not clear what this sequence represents, though it is not A. (Chlorandrena) kamarti Schmiedeknecht, 1900 from North Africa and Sicily as it also diverges from Moroccan A. kamarti sequences (to be published in the upcoming North African revision). It may also be a NUMT; it is unknown if certain Andrena subgenera are more likely to generate NUMTs, but Chlorandrena may potentially represent one such case, with difficult to interpret results generated in the study of the East Mediterranean fauna (G. Pisanty, unpublished data). More study using additional genetic markers is necessary to understand why two seemingly aberrant strong divergent sequences have been generated from Iberian Chlorandrena specimens.

Finally, the taraxaci-group (see Schwenninger 2015) formed a monophyletic clade, with A. (Chlorandrena) rhenana Stöckhert, 1930 clearly separated from A. (Chlorandrena) taraxaci Giraud, 1861 which has its western range limit in Central Europe. The taxonomic concepts of Schwenninger (2015) are therefore supported and followed.

Material examined. Andrena leucolippa: France: Riscle [43.6564°N, -0.0894°W], 1♀, MNHN (lectotype; Fig. 3).

Subgenus Didonia Gribodo, 1894

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. 4). The spring and summer generation individuals from Guadalajara were separated by 0.15%, strongly supporting the position that A. mucida is a bivoltine species displaying the currently unique trait of intergenerational variation in the structure of the pollen collecting hairs.

Figure 4. 

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 (Warncke 1968a). Genetic evidence instead places this taxon in the subgenus Euandrena (Bossert et al. 2022). This placement is a better fit, as A. solenopalpa has typical Euandrena foveae (narrow and narrowing further ventrally), the male A3 slightly exceeds the length of A4+5, and the process of the labrum is not so strongly thickened and elongate as in A. mucida; this simply represents convergence on a similar morphology. It is possible that the subgenus Didonia is monotypic, containing only A. mucida, but this requires additional study.

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.

Subgenus Euandrena Hedicke, 1933

In comparison to the situation in the Eastern Mediterranean (Praz et al. 2019), Iberian Euandrena are comparatively well-resolved, with lower species diversity, fewer undescribed species, and clearer species boundaries. However, unrecognised montane species are present (Wood et al. 2021), and there are some problems related to the group of species lumped together under a broad A. bicolor Fabricius, 1775. This current work does not deal with the complex issue concerning the two clades of A. bicolor as identified by Praz et al. (2019), but instead with the status of material from southern Spain and North Africa. Material identified as A. bicolor s.l. from the Col du Zad in the Middle Atlas of Morocco at an altitude of 2100 m [WPATW387-22] and the southern slopes of the Sierra Nevada above Trevélez in Spain at an altitude of approximately 1800 m [WPATW368-21] fell close to A. angustior (Kirby, 1802) and far from A. bicolor s.l. (Fig. 5). The specimen from the Sierra Nevada was caught on the same Campanula (Campanulaceae) plant as specimen WPATW290-21 which falls into the A. bicolor s.l. clade. These two Spanish specimens are separated by a genetic distance of 9.80%, and clearly cannot be conspecific despite their strong morphological similarity. More broadly, the Sierra Nevada specimen was separated from the A. bicolor s.l. clade by an average genetic distance of 9.32% (range 7.95–9.80%) and the Moroccan specimen was separated from the A. bicolor s.l. clade by an average distance of 9.06% (range 8.50–9.80%). These two specimens were separated from each other by a distance of 4.99%. The species from the Sierra Nevada is described as new to science below, but the name to apply to the North African taxon (present also in Tunisia based on barcode data, Praz, in litt.) is complex and requires discussion. The nomenclatural and taxonomic status of A. bicolor s.l. will be resolved in a future revisionary work dedicated to the subgenus Euandrena at a West Palaearctic scale.

Figure 5. 

Phylogenetic tree (maximum likelihood) of Andrena subgenus Euandrena Hedicke, 1933 based on the mitochondrial COI gene. Andrena (Pruinosandrena) succinea Dours, 1872 is used as an outgroup. Numbers adjacent to branches represent bootstrap support (values of <75 are omitted).

Andrena (Euandrena) bicolor and associated Euandrena species in North Africa

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.

Warncke (1974) used subspecific concepts for dealing with A. bicolor, with two subspecies used for North African material – A. b. oraniensis Lepeletier, 1841 and A. b. agraria Warncke, 1974 (see also distribution maps in Gusenleitner and Schwarz 2002). However, the use of these names is not immediately straightforward. Andrena oraniensis Lepeletier, 1841 was described from Oran in northern Algeria (Lepeletier 1841) and is a confused and unclear taxon. In the MNHN collection, the type series cannot be located, and it may have been lost as many specimens have been moved between different boxes by past workers and curators. In the Pérez collection, there are long series of Andrena (Melandrena) florentina Magretti, 1883 that were identified by Pérez as A. oraniensis. Warncke (1967) noted this, but argued that this must be incorrect due to size, and instead placed the name in combination with A. bicolor as a subspecies found in North Africa. Warncke (1974) later authored the replacement name A. bicolor agraria Warncke, 1974 for Andrena (Euandrena) nigriventris Pérez, 1902 which was described from northern Algeria and Morocco (locus typicus Tangier, following lectotype designation) but which is a preoccupied name, nec. Apis nigriventris Gmelin, 1790 which is a synonym of A. (Melandrena) nitida (Müller, 1776). Warncke then used both oraniensis and agraria as subspecies, giving an overlapping distribution in Warncke (1974, e.g. both occurring at Oukaimeden in the High Atlas), but in his distribution maps (see Gusenleitner and Schwarz 2002) giving a non-overlapping distribution, with oraniensis more or less occurring only in Algeria and Tunisia and agraria occurring only in Morocco.

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. Pérez (1902, as A. nigriventris) writes: “Abdomen très luisant, particulièrement les depressions, que précèdent les bourrelets très prononcés”. This is suggestive, as the distinct North African taxon has strongly depressed tergal margins. However, barcoded A. bicolor s.l. from Morocco also show this character, so it is not diagnostic in and of itself. Examination of the lectotype specimen of A. nigriventris (designated by Warncke 1967; Fig. 6) shows that the clypeus is apically smooth and shing between the punctures (Fig. 6C), meaning that it cannot belong to the distinct taxon, and it remains a synonym of A. bicolor s.l. One additional name is available, that of A. (Euandrena) fervida Pérez, 1902, a taxon described from Algeria. The lectotype is in poor condition, lacking its metasoma (Fig. 7). Diagnosis is therefore challenging, but the clypeus is shiny between the punctures and it cannot be conspecific with the distinct North African taxon. The position of Warncke (1967) that this name is synonymous with A. bicolor s.l. is maintained. The distinct North African taxon would therefore appear to be undescribed. No taxonomic action is taken in the current work – further genetic data will either demonstrate a closer affinity with the specimen from the Sierra Nevada, in which case this name will be available to apply to North African populations, or these sequences will confirm its distinct nature, and it can be described. In any case, a name is required for the Iberian lineage which is unambiguously undescribed.

Figure 6. 

Andrena (Euandrena) nigriventris Pérez, 1902 (nec. Gmelin, = Andrena bicolor agraria Warncke, 1974) female lectotype A label details B profile C face, frontal view D terga, dorsal view.

Figure 7. 

Andrena (Euandrena) fervida Pérez, 1902, female lectotype A label detail B profile C face, frontal view D face detail, frontal view.

Material examined. Andrena bicolor: Algeria: Theniet El Had [35.8727°N, 2.0007°E] (1♀, MNHN (lectotype of A. fervida); Morocco: Tanger [35.7537°N, -5.7906°W], 1♀, MNHN (lectotype of A. nigriventris Pérez).

Andrena (Euandrena) lavandulae Pérez, 1902, sp. resurr

Andrena (Euandrena) lavandulae Pérez, 1902: 156 ♀♂ [France, lectotype by present designation: MNHN].

Andrena (Euandrena) angustior impressa Warncke, 1967: 234, ♀♂ [Morocco: OÖLM, examined] syn. nov.

Remarks. Warncke (1967) synonymised A. lavandulae with A. bicolor without apparently inspecting the type. This is not explicitly clear in the text of Warncke (1967), but in this publication he typically reported lectotypes in the Paris collection when he designated them, and usually labelled paratypes as well, often taking a selection for his personal collection. There are no specimens of A. lavandulae in the Warncke collection, which combined with the lack of mention of a lectotype in Warncke (1967) leads me to believe that he did not inspect the type series. He then described Andrena angustior impressa Warncke, 1967 in the same publication, a taxon that was subsequently elevated to species status by Wood et al. (2021) based on genetic and morphological evidence.

In the MNHN collection, specimens of A. lavandulae are labelled as lectotype and paralectotype by Teunissen (August 1986; Fig. 8), but these designations were never published. The female specimen is therefore recognised here as a lectotype, by present designation. Examination of these specimens shows that they are clearly the same taxon as A. impressa and distinct from A. bicolor, with the weakly depressed, shagreened, and weakly shing tergal margins, the black terminal fringe and hairs flanking the pygidial plate, the intermixed light and dark hairs on the face, and in the male sex by the broadened gena (broader than the width of the compound eye). The lectotype comes from Banyuls-sur-Mer in the south-west of France, which falls within the distribution of the taxon as presented by Wood et al. (2021). Additional genetic sequences from Spain and Morocco closely match the Portuguese sequence of A. impressa presented by Wood et al. (2021), forming a well-supported clade (Fig. 5; bootstrap support of 93) that is distinct from both A. angustior and A. bicolor. Andrena impressa syn. nov. is therefore synonymised with A. lavandulae sp. resurr. The distribution is Morocco, Algeria, Portugal, Spain, and France (Wood et al. 2021).

Figure 8. 

Andrena (Euandrena) lavandulae Pérez, 1902, female lectotype A label details B profile C face, frontal view D terga, dorsal view; male syntype E label details F profile.

Material examined. France: Banyuls [Banyuls-sur-Mer, 42.5658°N, 2.8658°E], 1♂, 1♀, MNHN (female lectotype, by present designation).

Subgenus Graecandrena Warncke, 1968

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. 1).

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.

Subgenus Melanapis Cameron, 1902

Andrena (Melanapis) fuscosa Erichson, 1835

Andrena fuscosa was described from southern Spain by Erichson (1835). Andrena fuscosa has had a complicated taxonomic history, and has been described many times due to its variation in colouration and size (Gusenleitner and Schwarz 2002). Iberian genetic data are therefore of interest since the peninsula represents the locus typicus. Iberian sequences were identical with each other, and closely matched sequences from Israel and India (the latter identified as A. (Plastandrena) agilissima Scopoli, 1770 but clearly misidentified; Melanapis was actually described as a genus based on material from India), being separated by an average of 1.53% (range 0.88–1.82%; Fig. 9). They were more clearly separated from Moroccan sequences, by an average of 5.43% (range 3.94–5.93%). Given the lack of morphological differences between North African and Iberian specimens, a broad interpretation of this species is taken here. Including all sequences together, average intraspecific distance is 3.14% (range 0.00–6.23%) which is considered to be acceptably small. Additional study is necessary to establish whether North African material is consistently distinct; the first nominally available name would be A. (Melanapis) rutila Spinola, 1838 which was described from Egypt.

Figure 9. 

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).

Subgenus Melandrena Pérez, 1890

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

Andrena (Melandrena) morio Brullé, 1832: 353, ♀♂ [Greece: MNHN, not examined].

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. 10). Confusingly, additional sequences from Greece, Morocco, and Tunisia formed two sister groups to this clade. There is no clear geographic pattern, and so a broad A. morio concept is adopted, including A. hispania syn. nov. as a direct synonym of A. morio. Additional genetic data using more powerful techniques are required before an alternative taxonomic conclusion can be drawn. Andrena morio is known to be highly variable in its colouration, leading to a higher than average number of synonyms for an Andrena species (Gusenleitner and Schwarz 2002). Andrena hispania was classically thought of to have only a single generation in the spring, which may explain why its wings are less infuscate than A. morio, particularly in the summer generation of this species that can be noticeably darker than spring flying individuals.

Figure 10. 

Phylogenetic tree (maximum likelihood) of Andrena subgenus Melandrena Pérez, 1890 based on the mitochondrial COI gene. Andrena (Ovandrena) oviventris Pérez, 1895 is used as an outgroup. Numbers adjacent to branches represent bootstrap support (values of <75 are omitted).

Andrena (Melandrena) nitida (Müller, 1776), Andrena (Melandrena) thoracica (Fabricius, 1775), and Andrena (Melandrena) limata Smith, 1853

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 MNHN collection, but which I could not locate.

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 Wood et al. 2020a). This form can be recognised by the dense punctures of T1, separating it from Iberia A. thoracica; the colour of the hind tibial spur is variable and of little consistent utility. Because of this variation, Warncke later adopted a much broader concept of A. nitida, with A. nitida s. str., A. nitida limata, and A. nitida mixtura (e.g. Warncke 1974). This position was not followed by Gusenleitner and Schwarz (2002) who followed a three distinct species interpretation (nitida, limata including mixtura, thoracica).

Analysis of barcodes does not provide clarity (Fig. 10). Whilst A. thoracica and A. nitida form reciprocally monophyletic clades, three different clades are formed for A. limata. There is no geographic pattern to these clades; clade #1 contains individuals from Austria, Poland (KJ837115; specimen identified as A. nitida but associated photograph on BOLD shows A. limata colour pattern), Spain, and Portugal; clade #2 contains individuals from Morocco, Spain, and Portugal; and clade #3 contains individuals from Finland (MZ625969; specimen identified as A. nitida but associated photograph on BOLD shows A. limata colour pattern), France, and Spain. Confusingly, sequences IBIHM501-21 and WPATW086-21 come from two different female specimens collected from the same field near Pedret i Marzà in north-eastern Spain. As Iberian individuals fall into three of these clades and the power of COI analysis is clearly insufficient to deal with potential hybridisation and introgression, no further action can be taken on the basis of these results. A dedicated study using more powerful genetic markers is necessary to resolve this issue. For now, I maintain the position of Gusenleitner and Schwarz (2002) in that there are three species in this group (nitida, limata including mixtura, and thoracica). It will probably be necessary for a future study to designate a neotype for A. limata if Lepeletier’s original specimens cannot be definitively located.

Andrena (Melandrena) discors Erichson, 1841

Andrena (Melandrena) discors Erichson, 1841: 192, ♀ [Algeria: ZMHB, examined].

Andrena (Melandrena) creberrima Pérez, 1895: 46, ♀♂ [Algeria: MNHN, examined] syn. nov.

Andrena (Melandrena) vachali Pérez, 1895: ♀ [Tunisia, lectotype by present designation: MNHN] syn. nov.

Andrena (Melandrena) bodemeyeri Benoist, 1969: 246, ♀ [Tunisia: MNHN, examined].

Remarks. There has been extensive confusion between A. discors and A. creberrima. Erichson (1841) described A. discors from Algeria; a specimen is preserved in the ZMHB collection which may be automatically the holotype, but this is ambiguous as it is unclear if Erichson described the species from multiple specimens (Fig. 11). In any case, the specimen is a syntype, and Erichson described the species only in the female sex. The specimen conforms to the classical concept of the species, having the typical features of the former members of the subgenus Zonandrena, i.e. a clypeus with punctures forming a pattern of weak longitudinal channels. The pubescence of the face is black (Fig. 11C), and there are numerous intermixed dark hairs on the scutum (Fig. 11D). The tibial scopae are orange (Fig. 11E), and there are weak indications of light tergal hair bands but these are degraded and scarcely visible (Fig. 11F). Warncke (1974) gave numerous localities for A. discors from Morocco, Algeria, and Tunisia. He listed A. aff discors from Madrid based on two specimens (Warncke 1976), but later discarded these records, giving only two points in southern Spain in his distribution maps in addition to the points from north-western Africa (Gusenleitner and Schwarz 2002). I have examined 14 female and six male specimens of A. discors, all from southern parts of Portugal and Spain (Algarve, Alto Alentejo, Cádiz, Huelva, Málaga, Sevilla), where it is active from late February to April.

Figure 11. 

Andrena (Melandrena) discors Erichson, 1841, female holotype/syntype A label details B profile C face, frontal view D scutum, dorsolateral view E tibial scopa, profile view F terga, dorsal view.

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. Warncke (1967) designated a lectotype for A. creberrima (Fig. 12), but did not understand the concept of this species, listing only the type locality of Bône in Warncke (1974). He later gave a distribution map of Algeria (the holotype point), Italy, and Greece (Crete). Examination of the lectotype shows that the specimen is slightly paler than the type of A. discors, with fewer dark intermixed hairs on the scutum (Fig. 12B), and the tergal hair bands in better condition and therefore slightly more pronounced (Fig. 12D). This can be contrasted with the type of A. (Melandrena) bodemeyeri (Fig. 13; known synonym of A. discors, see Warncke 1967) which is comparatively dark. This kind of variation in hair colour is typical for the subgenus Melandrena, and has led to the large number of synonymous names being described, for example for A. (Melandrena) flavipes Panzer, 1799.

Figure 12. 

Andrena (Melandrena) creberrima Pérez, 1895, female lectotype A label details B profile C face, frontal view D terga, dorsal view.

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 (Gusenleitner and Schwarz 2002). Moreover, type material for A. vachali has not actually been designated, as Warncke (1967) could not find material in the MNHN collection. Examination of the MNHN showed several specimens of A. vachali in the Vachal collection from Foum Tatahouine [Tataouine, Tunisia] labelled “Andrena vachali type Pérez” in the handwriting of Vachal (Fig. 14). One specimen was labelled as a lectotype with a handwritten ‘A. vachali’ determination label by H. Teunissen (Fig. 14A), but this designation was never published. Pérez (1895) did not mention the collection locality, and Warncke (1967) indicated that the species was described from Algeria. However, Gusenleitner and Schwarz (2002) give the locus typicus as Tunisia. Examination of the catalogue of Pérez did not reveal an entry for A. vachali, as is often the case, either because Pérez did not write one, or because the information is hidden under a name that differs from the published name (see below for the case of A. (Plastandrena) asperrima Pérez, 1895). It is therefore very difficult to know the type locality with certainty. However, on the basis that the MNHN specimens are labelled as ‘type’ by Vachal, they were present in his collection, and the species was named after Vachal, the position is taken that these represent genuine syntypes. The female specimen labelled as ‘type’ by Vachal is chosen as a lectotype, by present designation. Morphologically, it corresponds to the concept of A. vachali, with black pubescence on the face and mesepisternum, with reddish-brown pubescence on the remaining parts of the mesosoma, and with predominantly reddish-brown tibial scopae, though this specimen shows some darker hairs dorsally (c.f. Fig. 13D); specimens of A. vachali can be found with entirely orange-red tibial scopae. Comparison of sequences from A. discors from southern Portugal and two A. vachali from southern Israel show a very small difference of 0.91% (range 0.91–0.91%; Fig. 10). These three specimens group together and have posterior support of 100, and are clearly separated from the three other former Zonandrena species that group together in this clade, namely A. flavipes, A. (Melandrena) gravida Imhoff, 1832, and A. (Melandrena) vulcana Dours, 1873.

Figure 13. 

Andrena (Melandrena) bodenmeyeri Benoist, 1969, female holotype A label details B profile C face, frontal view D dorsal view.

Figure 14. 

Andrena (Melandrena) vachali Pérez, 1895, female lectotype A label details B profile C face, frontal view D terga, dorsal view.

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. 15). Indeed, the genital capsule of a syntype of A. creberrima (Fig. 15B), the barcoded A. discors from southern Portugal [IBIHM517-21] (Fig. 15C), and A. vachali specimens from Morocco have identical genital capsules (Fig. 15D). Moreover, numerous ‘aberrant’ female individuals can be found across this range with more or less pronounced tergal hair bands and variably dark or light pubescence, particularly in Crete where individuals corresponding in different ways to the concepts of A. creberrima, A. discors, and A. vachali can be found.

Figure 15. 

Andrena (Melandrena) creberrima Pérez, 1895, male syntype A label details B male genital capsule; Andrena (Melandrena) discors Erichson, 1841 C male genital capsule; Andrena (Melandrena) vachali Pérez, 1895 D male genital capsule.

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 (Baldock et al. 2018; Wood et al. 2020a).

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♀, ZMHB (holotype/syntype of A. discors); Bône [=Annaba, 36.9092°N, 7.7264°E], 1♀, MNHN (lectotype of A. creberrima); Alger, Birmandreis [Bir Mourad Raïs], 24.iii.-30.iv.1910, 1♂, 1♀, leg. J. Bequaert, ZMHB; Greece: Krete, Anidri b. Paleochora, 200 m, 31.iii.2002, 1♀, leg. A. Müller, AMC; Krete, Palaeochora, 0–50 m, 29.iii.2002, 5♀, leg. A. Müller, AMC; Kreta, Kavalos [Kavallos], 21.v.1986, 1♀, leg. Brande, OÖLM; Crete, Dept. Heraklion, Moulia, S. of Aghia Varvara, 26.iv.1973, 1♀, leg. v. Ooststroom, RMNH; Italy: Lazio, Roma, Via Falcognana, 8.iv.1990, 1♀, leg. G.G.M. Schulten, RMNH; Roma, Via Falcognana, 15.iv.1993, 1♀, leg. G.G.M. Schulten, RMNH; Israel: Negev, 15 km S of Be’er Sheva, 31.iii.1988, 2♀, leg. R. Leys, RMNH; Jordan: Petra, 800 m, 28.ii.-4.iii.1986, 1♂, 2♀, leg. K.M. Guichard, NHMUK; Malta: Buskett [woodlands, Ħad-Dingli], 7.iii.2018, 1♀, M. Balzan Colln.; Morocco: Drâa-Tafilalet, Tazenakht, Anezal, 5 km NE Ait Igga, 21.iv.2022, 1♂, 4♀, leg. T.J. Wood, TJWC; Souss-Massa, R105, Tizirt, 8 km N, Ighir Ifran env., 12.iii.2022, 1♂, 1♀, leg. T.J. Wood, TJWC; Portugal: Foros de Vale Figueira, Montemor-o-Novo, 20.iii.2019, 1♂, leg. T.J. Wood, TJWC; Montemor-o-novo, Foros de Vale de Figuera, 24.iv.2011, 1♀, leg. A. Albernaz-Valente, TJWC; Spain: Canary Islands, Lanzarote, Haria, 6.ii.1979, 1♂, leg. Ellis & Simon-Thomas, RMNH; Bollullos Par del Condado (Huelva), 160 m, 28.iii.2009, 1♂, leg. F.J. Ortiz-Sánchez, FJOS; Puerto de Gáliz, P.N. Alcornocales (Cádiz), 425 m, 28.iv.2007, 1♀, leg. F.J. Ortiz-Sánchez, FJOS; Cádiz, Zahara de los Atunes, 12.iii.1977, 1♀, leg. Madey, NHMUK; Tunisia: Foum Tatahouine [32.9361°N, 10.4458°E], 1♀, MNHN (lectotype of A. vachali, by present designation); Oasis Gafsa [34.4350°N, 8.7678°E], 1928, 1♀, leg. B. v. Bodemeyer, MNHN (holotype of A. bodemeyeri); Fritissa, 17–19.iv.1983, 1♀, TJWC.

Subgenus Micrandrena Ashmead, 1899

This is by far the most species-rich subgenus in Iberia, containing 37 species. Pisanty et al. (2022b) took the decision to synonymise the subgenera Distandrena Warncke, 1968, Fumandrena Warncke, 1975, and Proxiandrena Schmid-Egger, 2005 with a broad concept of Micrandrena, as morphological differences such as the shape of the foveae, propodeal triangle, and presence or absence of striations on the clypeus are inconsistent and insufficient to allow unambiguous differentiation between the groups, as well as lacking phylogenetic support. This broad Micrandrena definition is followed here.

Andrena (Micrandrena) obsoleta Pérez, 1895 and Andrena (Micrandrena) mariana Warncke, 1968 sensu lato

The treatment of A. obsoleta has a long and confused history that has caused many problems. Warncke (1967) designated a female lectotype from Babor in northern Algeria (Fig. 16). However, this lectotype designation is incorrect. In the catalogue of Pérez, under entry 1506 (p. 214), Pérez writes: “Sicile, ♀ - M’pellier ♀. – Tebessa, avril, ♀. – Biskra, mars, 3♀, rares. – Bône, ♀. – Marseille, ♂. - Bône, 27/iv ♂♀ in cop. Souci et autres [illegible – presumably a type of flower]”. Babor is not mentioned, and so Warncke’s lectotype designation is rejected. Examination of specimens labelled as obsoleta by Pérez from the localities indicated shows that they belong to a different taxon to that of Warncke’s false lectotype. Warncke’s false lectotype is a female Distandrena (=Micrandrena) with a clearly striate clypeus, and T1 is polished. This conforms to the concept of A. (Micrandrena) nitidula Pérez, 1903, and indeed Warncke (1967) synonymised A. nitidula with A. obsoleta. Only female specimens of A. obsoleta from Bône [=Annaba] in northern Algeria could be found in the Pérez collection, and almost all were in very poor condition, missing their metasomas. A single female from Bône was however in good condition, and it is here designated as a new lectotype (Fig. 17). This new lectotype specimen (i.e. the true A. obsoleta) morphologically falls within the A. (Micrandrena) mariana Warncke, 1968 complex within the former Distandrena, as it lacks striations on the clypeus. The larger A. (Micrandrena) merimna Saunders, 1908 can be excluded as this species has a very long A3 that clearly exceeds the length of A4+5, whereas members of the A. mariana complex have A3 at most slightly exceeding A4+5, A4 and A5 are sub-quadratic and slightly shorter than wide, A6–12 are as long as wide.

Figure 16. 

Andrena (Micrandrena) obsoleta Pérez, 1895, false female lectotype, =Andrena (Micrandrena) nitidula Pérez, 1903 A label details B profile C face, frontal view D dorsal view.

Figure 17. 

Andrena (Micrandrena) obsoleta Pérez, 1895, true female lectotype A label details B profile C face, frontal view D scutum, dorsal view E propodeal triangle, dorsal view F terga, dorsal view.

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 (Warncke 1968b). He then described five subspecies in subsequent publications (Warncke 1974, 1975a): A. m. mica Warncke, 1974, A. m. leptura Warncke, 1974, A. m. solda Warncke, 1974, A. m. alma Warncke, 1975, and A. m. tenostra Warncke, 1975. For North African taxa, Warncke (1974) described A. mariana solda from Morocco (Tangier) and Algeria (Algiers, including Husseyn-Dey [Hussein Dey] the locus typicus, Constantine, Bone, Blida) and Tunisia (Carthago), and noted that this species corresponds to the Andrena forms from Bone identified as iota, kappa, and lambda by Saunders (1908), though this is clearly wrong for lambda as the genital capsule illustrated by Saunders shows A. (Micrandrena) djelfensis Pérez, 1895. The A. mariana solda taxon differs from A. mariana s. str. in the female sex by the darker tergal margins, the darker antennae, and the more extensively shagreened clypeus. Warncke (1974) described A. mariana mica from southern Algeria (locus typicus Ghardaia) and Morocco, giving it a more arid distribution. It also differs from A. mariana s. str. in the female sex by the darker tergal margins and darker antennae, but the clypeus is more extensively shiny than in A. mariana solda, and the nervulus is said to be more antefurcal. Andrena m. leptura is a more eastern taxon, being described from Libya and Egypt, and having a broader process of the labrum than any of these species.

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 (Warncke 1974, as A. m. solda). Pérez’s reference to specimens from France indicates that his original syntypic series would have been polytypic, or he may have changed his mind between writing this catalogue entry and his 1895 publication, or even after this point. Warncke (1974: 40) indicated that A. m. solda is present in Sicily which is supported by a newly examined specimen (see below), though he did not designate any Italian specimens as paratypes. As no specimens from France labelled as obsoleta could be found in Pérez collection, the conservative position is taken here that this species does not occur in France. The implications for Warncke’s misinterpretation of A. obsoleta are discussed below.

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. 18). Contrary to the position of Warncke (1968b), examination of material from Morocco, Algeria, and Tunisia shows that A. mariana is actually widely distributed across this region, predominantly in southern, more arid areas. Genetically, A. mariana s. str. was more closely related to A. (Micrandrena) abjecta Pérez, 1895, separated by an average genetic distance of 5.55% (range 5.49–5.61%), and was strongly separated from A. m. mica by 12.73% (range 12.50–12.95%) and A. m. alma by 10.94% (range 10.65–11.39%). Andrena abjecta + A. mariana s. str. therefore formed a clade with bootstrap support of 99. Andrena m. mica and A. m. alma formed a clade, but were strongly separated from each other by 11.26% (range 10.97–11.41%). The conclusion therefore is that each of these taxa is distinct, and so A. mica stat. nov. and A. alma stat. nov. are raised to species status. Morphologically, all three taxa can clearly be separated by the structure of the female clypeus, the colour of the antennae, the extent of lightening on the tergal margins, the strength of the scutal punctures, and the shape of the process of the labrum.

Figure 18. 

Phylogenetic tree (maximum likelihood) of Andrena subgenus Micrandrena Ashmead, 1899 based on the mitochondrial COI gene. Andrena (incertae sedis) murana Warncke, 1975 is used as an outgroup. Numbers adjacent to branches represent bootstrap support (values of <75 are omitted).

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 (Warncke 1976). The status of A. mariana leptura will be dealt with in a future publication, as its exact species concept and its eastern and western limits are unclear, though it will be a valid species as it clearly differs from A. mariana s. str.; its relationship with A. mariana s.l. specimens from southern Israel must also be clarified (Pisanty, in litt.). In Iberia, A. m. tenostra is very poorly known. The distinguishing features given by Warncke (1975a) are that the process of the labrum is narrowly trapezoidal and that the clypeus is arched and somewhat flattened in the middle, in contrast to A. alma where the clypeus is largely flattened. Additional female specimens were found in Granada which highlight a difference not noted by Warncke, which is that the scutellum is polished and shiny, whereas it is dull in A. alma, and more broadly the scutal punctures are less strongly pronounced. Given that subtle differences within this group represent large genetic differentiation, A. tenostra stat. nov. is raised to species status.

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. Warncke (1976) did not record this species from Iberia, and only his distribution maps (Gusenleitner and Schwarz 2002) indicate the presence of this species in northern and eastern Spain. This distribution does not make sense given that all other known records of A. abjecta come from north-western Africa (Morocco, Algeria, Tunisia), so the species would naturally be expected to occur in southern Iberia. Dardón (2014) listed two female specimens from Huelva (22.iv.1987) and Zaragoza (9.iii.1991). The specimen from Huelva may represent the undescribed aff mica taxon, but it is more difficult to conclude on the specimen from Zaragoza. For now, A. abjecta is retained on the Iberian list, but I have not examined any specimens and I consider its presence in Iberia doubtful.

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, RMNH; 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; Castelo Branco, Fundão Vale de Prazeres, 17.iv.2021, 1♀, leg. C. Siopa, FLOW; Spain: El Soldado, Córdoba [38.3747°N, -5.0001°E], 1♀, leg. Seyrig, OÖLM (holotype); 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, EBDC.

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, RMNH; 23 km NE of Ras El Ma, st. 38, 26.iv.1983, 1♀, leg. R. Leys & P. v. d. Hurk, RMNH; Dayet el Kerch [Daïet el Kerch], st. 5, 5.iv.1983, 2♀, leg. R. Leys & P. v. d. Hurk, RMNH; Morocco: Guelmim-Oued Noun, Guelmim, Asrir, 1 km W, 20.iii.2022, 1♀, leg. T.J. Wood, TJWC; Spain: Fuerteventura, Valley Granadillos [28.3864°N, -14.0865°W], 17.iv.1934, 1♀, OÖLM (holotype); Tunisia: Ksar Hadada, 4–5.iv.1996, 46♀, leg. K. Deneš, OÖLM; M’saken, 20–21.iv.1998, 2♀, leg. K. Deneš, OÖLM; Wadi Ram, 45 km E Douz, 4.iv.2006, 1♀, leg. J. Straka, OÖLM.

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, RMNH; Tunisia: Hammamet env, 15.iii.1996, 1♀, leg. K. Deneš, OÖLM; Kasserine, 13.iv.1998, 1♀, leg. K. Deneš, OÖLM; Nefta [Naftah], 20.v.1993, 1♀, leg. J. Batelka, OÖLM.

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♀, MNHN (false lectotype of A. obsoleta).

Andrena obsoleta: Algeria: Bône [=Annaba, 36.9092°N, 7.7264°E], 1♀, MNHN (lectotype of A. obsoleta, by present designation); 23 km NE of Ras El Ma, st. 38, 26.iv.1983, 1♀, leg. R. Leys & P. v.d. Hurk, TJWC; 5 km N of Mecheria, Djebel Antar, st. 10, 7.iv.1983, 1♀, leg. R. Leys & P. v.d. Hurk, RMNH; Italy: Sicilia orient., Taormina, 200 m, Ghardino [Giardini Naxos], 25.iii.1950, 1♂, Hartig & Grisham, MZUR; Tunisia: Tunis, 1911, 3♀, leg. O. Schmiedeknecht, SMFD.

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.

Andrena (Micrandrena) distinguenda Schenck, 1871 and Andrena (Micrandrena) nitidula Pérez, 1903

The status of these two species has been extensively argued over, and despite much attention the position remains somewhat unclear. Warncke (1967) argued that A. distinguenda Schenck, 1871 was an unavailable name because it was preoccupied by A. distinguenda Schenck, 1853. He then authored the replacement name A. obsoleta spongiosa Warncke, 1967, using the incorrect concept that A. obsoleta was the oldest available name for this group of species, as discussed above. Schönitzer et al. (1992) clarified the situation, and argued that A. distinguenda Schenck, 1853 was a nomen nudum because it appears without a description in a discussion concerning species related to A. bicolor. As a nomen nudum, the name is available and cannot preoccupy its later usage. Therefore, the name A. distinguenda Schenck, 1871 is both available and valid as the senior synonym; A. obsoleta spongiosa is consequently an unnecessary replacement name.

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 Warncke 1967, Fig. 19). As discussed above, Warncke (1967) incorrectly recognised a specimen that morphologically corresponds to A. nitidula as the lectotype of A. obsoleta, and then synonymised A. nitidula with A. obsoleta. He then treated A. distinguenda Schenck, 1871 as a subspecies of A. obsoleta in the form A. obsoleta spongiosa. He then later adopted a three subspecies model, A. o. obsoleta (North Africa), A. o. nitidula (Western Europe), and A. o. spongiosa (Central Europe) (see Schönitzer et al. 1992), finally introducing a fourth subspecies A. o. puella Alfken, 1938 in Sardinia and Sicily (see Gusenleitner and Schwarz 2002). Schönitzer et al. (1992) argue that both A. distinguenda and A. nitidula can be morphologically separated in north-western Italy (Bordighera). They argue that A. nitidula occurs in North Africa, citing Pérez (1903) and Alfken (1914). They also argue that specimens from southern Spain (Tarifa) appear to be somewhat intermediate between A. distinguenda and A. nitidula, concluding that the question as to whether there are two species or simply a very variable single species remains open.

Figure 19. 

Andrena (Micrandrena) nitidula Pérez, 1903, female lectotype A label details B profile C face, frontal view D terga, dorsal view.

Burger and Herrmann (2003) revised European material from this species pair using the names A. distinguenda and A. nitidula, after having examined syntypes of A. nitidula in the ZMHB collection. Using morphological characters, they conclude that (in a European context) A. distinguenda has a predominantly eastern distribution, from Central Europe to Greece, with populations also in the south of France and a few records from north-eastern Spain. In contrast, A. nitidula is a western species, from Iberia to the south and east of France and into south-western Germany and the north-west of Italy. They reach the same conclusion as Schönitzer et al. (1992), that where these two forms occur in sympatry (Germany, France, Italy) they can be morphologically separated. They do not conclude on the status of A. obsoleta, but note that A. nitidula occurs in North Africa, and that Warncke’s treatment is unclear. This problem arises because it does not appear that any of the authors of Schönitzer et al. (1992) or Burger and Herrmann (2003) inspected the lectotype of A. obsoleta designated by Warncke in the MNHN. This lectotype bears a handwritten label that the lectotype designation is incorrect, and that the specimen corresponds to A. distinguenda Schenck (Fig. 16A). However, it is unclear who wrote this, as neither Schönitzer et al. (1992) nor Burger and Herrmann (2003) mention this explicitly, and I do not recognise the handwriting on the label.

Barcode analysis complicates this matter further (Fig. 18). Analysis of sequences from France, Germany, Morocco, Portugal, and Spain shows the presence of two clear clades. One clade is composed of specimens identified as A. nitidula from France, Portugal, and Spain, including specimens from the south of Iberia (Algarve, Málaga, Murcia). This clade would appear to be the true A. nitidula. However, the second clade contained sequences from specimens identified as A. distinguenda from central Germany, but also from a specimen identified as A. nitidula from central Spain (Guadalajara) and two specimens identified as A. nitidula from northern Morocco (Fès-Meknes, Oriental). These two clades were consistently separated genetically, by an average genetic distance of 7.08% (range 5.58–8.77%). The nominally true A. nitidula clade showed low intraspecific variation of 0.26% (range 0.00–0.63%). However, the situation in the clade containing A. distinguenda sequences from Germany showed intraspecific variation of 2.84% (range 0.00–4.10%), with the 4.10% genetic distance recorded between specimens from Germany and the Middle Atlas (Bakrit, near Azrou) in Morocco.

Morphologically, both Spanish and Moroccan specimens falling into the A. distinguenda clade conform to the concept of A. nitidula using the criteria specified by Burger and Herrmann (2003). Indeed, all material I have identified from north-western Africa morphologically conforms to the concept of A. nitidula sensu Burger and Herrmann. Although the genetic data strongly supports the existence of two species, the conflict between the genetic and morphological results suggests that, at the moment, they cannot be consistently separated morphologically across their range. No taxonomic action is taken here, and the morphological criteria of Burger and Herrmann (2003) are followed for the purpose of the identification key. In Iberia, A. nitidula is considered to be present throughout, but A. distinguenda is limited to the extreme north-east of Spain. However, it is clear that there is a major disagreement between the morphological and genetic methods, and these two species may ultimately not be consistently separable. Further analysis using more powerful genetic techniques is necessary.

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 (Gusenleitner and Schwarz 2002). Samples collected from the Sierra Nevada at high altitude (2000–2100 m) during 2021 are strongly differentiated from A. rugulosa, separated by an average genetic distance of 16.01% (range 15.81–16.41%; Fig. 18). Instead, they form a clade with bootstrap support of 96 with sequences of A. (Micrandrena) atlantea Wood, 2021 (High and Middle Atlas in Morocco) and a morphologically similar undescribed Micrandrena from high altitude (1800–1900 m) in the Moroccan Middle Atlas.

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 (Micrandrena) niveata Friese, 1887 sensu lato

Andrena niveata was described from Germany and Hungary, without further information (Friese 1887). The exact type locality is unclear, as well as the type depository, with no clearly identifiable syntypes located during searches in the SMFD and ZMHB collections, the suggested possible depositories (Gusenleitner and Schwarz 2002). Nevertheless, the concept of A. niveata in Central Europe is clear, and A. niveata is a well-defined species in this region (Schmid-Egger and Scheuchl 1997; Gusenleitner and Schwarz 2002; Amiet et al. 2010). Warncke described three subspecies; A. n. lecana Warncke, 1975 (Spain; locus typicus: Rivas-Vaciamadrid), A. n. haloga Warncke, 1980 (Italy; locus typicus: Lecce), and A. n. bubulca Warncke, 1975 (Turkey; locus typicus: Erzurum); the status of the Italian and Turkish subspecies is not addressed here. Unfortunately, all nine freshly collected A. n. lecana specimens from central and southern Spain (including from the locus typicus) sent for genetic analysis failed to produce sequences. More specific primers will be required to address this question using genetic evidence.

Warncke (1975a) described A. n. lecana extremely briefly, noting that in both sexes the first tergum was finely but clearly shagreened, and finely but more strongly punctured than in A. niveata s. str., with the following terga also more clearly and strongly punctured. He then gave a distribution of A. n. lecana across much of central Spain south to hilly parts of the provinces of Granada (Huéscar) and Almería (Tíjola). In contrast he gives a distribution for A. niveata s. str. of the whole of Iberia, though his distribution maps (Gusenleitner and Schwarz 2002) show a distribution covering eastern, central, and south-eastern Spain. There is therefore the possibility that the two subspecies exist in sympatry, suggesting that they may be distinct.

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. 20A). In contrast, in A. n. lecana, the inner margins of the gonostyli are strongly and clearly produced towards the penis valves (Fig. 20B); the apical spatulate part of the gonostyli are also comparatively shorter, thus the gonostyli appear shorter and more compact in A. n. lecana and longer and more elongate in A. niveata s. str. The structure of the genital capsule of A. niveata s. str. is consistent across Spain to the province of Málaga (the most southerly A. niveata s. str. male examined). Furthermore, the male scutum is shagreened in A. niveata s. str., whereas it is extensively shiny in A. n. lecana, and the terga are also more strongly shagreened in A. niveata s. str. compared to A. n. lecana (Fig. 20C, D). Separation of females (recognised by their wide and poorly defined propodeal triangle, Fig. 20E, F, in combination with their depressed tergal margins) is more challenging and may not be possible in all situations. Typically, A. niveata s. str. has the terga shagreened, with punctures on the tergal discs partially disappearing into this shagreenation (Fig. 20G). In contrast, A. n. lecana has the terga shiny, almost without shagreen, with the dense punctures on the tergal discs strong and clearly visible, not disappearing due to the absence of background shagreenation (Fig. 20H). The scutum and scutellum are also more extensively shiny, whereas in A. niveata s. str. these areas usually show at least some shagreenation and are never polished and smooth. The problem comes that some individuals nominally assigned to A. niveata s. str. have partially shing terga, and without genetic sequences it is not possible to say with complete confidence if they belong to A. niveata s. str. or A. n. lecana. Moreover, all Iberian A. niveata s. str. have tergal shagreenation that is weaker than that of A. niveata s. str. populations in Central Europe.

Figure 20. 

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, RMNH; Cádiz, Grazalema, Río Guadalete, 28.v.2021, 1♀, leg. T.J. Wood, TJWC; Cádiz, Embalse de Barbate Sw, 1♂, 3♀, 6.v.2017, leg. Barták & Kubik, OÖLM/TJWC; Granada, Sierra de Baza, Prados del Rey, 2000 m, 19.vi.2021, 11♀, leg. T.J. Wood, TJWC; Málaga, 5 km W of Alhaurín de la Torre, 4.vi.1962, 1♀, leg. Jeekel & Wiering, RMNH; Málaga, Cortes de la Frontera, path to Llano de las Labores, 26.v.2021, 3♀, leg. T.J. Wood, TJWC; Málaga, Estepona, 21.iv.1983, 1♀, leg. H. Teunissen, RMNH; Navarra, Tudela, 16.iv.1978, 1♀, leg. C. Gielis, RMNH; Gerona, Figueras, 15.iv.1971, 1♀, leg. J. Leclercq, UMONS.

Andrena (Micrandrena) spreta Pérez, 1895, Andrena (Micrandrena) curtula Pérez, 1903, Andrena (Micrandrena) pusilla Pérez, 1903, and Andrena (Micrandrena) pauxilla Stöckhert, 1935

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 Warncke (1967) designating a lectotype from Biskra (Fig. 21A; Pérez also mentions a female from Constantine in his catalogue). Andrena curtula was described from north-eastern Spain, with Warncke (1967) designating a lectotype from Barcelona (Fig. 21B). Andrena pusilla was described from the south of France (Bordeaux, Nantes) and Spain, with Warncke (1967) designating a lectotype from Nantes (Fig. 21C). Finally, A. pauxilla was described from the Upper Rhine valley in south-western Germany, with Stöckhert (1935) designating a female from Karlsruhe as the holotype.

Figure 21. 

Andrena (Micrandrena) spreta Pérez, 1895, female lectotype A label details B profile; Andrena (Micrandrena) curtula Pérez, 1903, female lectotype C label details D profile; Andrena (Micrandrena) pusilla Pérez, 1903, female lectotype E label details F profile.

Warncke (1967) recognised three species, A. spreta, A. curtula, and A. pauxilla, treating A. pusilla as a subspecies of A. spreta. Warncke (1974) recognised only one taxon in North Africa (A. spreta spreta), but Warncke (1976) surprisingly recognised only A. spreta pauxilla in Iberia, not mentioning A. curtula as a species despite its description from Iberia, only referring to its listing in Ceballos (1956) but indicating that this referred to A. (Micrandrena) tenuistriata Pérez, 1895. This treatment is difficult to understand. Warncke later adopted an extremely broad position (see distribution maps in Gusenleitner and Schwarz 2002), lumping all members of the group into a broad A. spreta. In this model, A. spreta s. str. was restricted to North Africa, A. spreta curtula was distributed across Iberia to southern France, A. spreta pusilla was found in northern and Central Europe, with two more subspecies in i) Italy and ii) south-eastern Europe to the Levant.

This position was not adopted by subsequent authors. Schwarz et al. (1996) accepted A. pusilla as a valid species, as well as A. curtula. However, they synonymised A. pauxilla with A. curtula. Schmid-Egger and Scheuchl (1997) did not follow this interpretation, treating A. pauxilla as a distinct species, though Amiet et al. (2010) followed the position of Schwarz et al. (1996). Dardón (2010) and Dardón et al. (2014) accepted only a broad A. spreta taxon, including A. curtula, A. pusilla, and A. pauxilla, recognising A. spreta s. str. and A. spreta pusilla in an Iberian context. The situation is therefore unclear, both for the number of species present in this complex, and their distributions.

Analysis of barcodes provides unambiguous support for the existence of three distinct species (Fig. 18), A. spreta including A. curtula, A. pusilla, and A. pauxilla sp. resurr. Specimens of A. spreta from Israel, Morocco, Portugal, Spain formed a clear clade with bootstrap support of 85. These sequences were separated by an average genetic distance of 0.45% (range 0.00–1.07%). They were clearly separated from a sister clade of A. pusilla sequences from Belgium, France, and Germany by an average genetic distance of 4.33% (range 3.88–4.84%). The A. pusilla clade had low intraspecific genetic distance of 0.29% (range 0.00–0.71%). Andrena pauxilla was strongly differentiated and was found as sister to A. (Micrandrena) rugulosa, showing average genetic differentiation of 10.96% (range 10.38–11.85%) from A. spreta and 11.69% (range 10.75–12.59%) from A. pusilla. This A. pauxilla clade includes a specimen from the south of France (KJ837178, Lac St. Croix) that was identified as A. curtula, the genetic results suggest that it is misidentified. Morphologically, A. pauxilla is clearly differentiated from A. spreta and A. pusilla due to the absence of a gradulus at the base of the terga. Confusion is impossible in the female sex after recognition of this character; the foveae are also longer and narrower and filled with white rather than light brown hairs, the scutum is more densely and uniformly punctate when compared to A. spreta. Examination of the lectotypes of A. spreta and A. curtula shows no clear structural difference. The degree of shagreenation varies, but this is typical for A. spreta across its range; both lectotype specimens show the presence of a gradulus at the base of the terga, excluding their conspecificity with A. pauxilla. Due to these genetic results combined with the morphology of the lectotype specimens, the synonymy of A. curtula with A. spreta as proposed by Dardón (2010) and Dardón et al. (2014) is therefore followed, though their synonymies of A. pusilla and A. pauxilla with A. spreta are rejected.

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. 22A, B), Sierra Cazorla, Sistema Central, and Sistema Ibérico (Fig. 22C, D). Andrena pauxilla would therefore have a currently known distribution of Morocco, Spain, France, and Germany. On the basis of these results, A. spreta would appear to be a pan-Mediterranean species, and A. pusilla its predominantly northern counterpart (though the species descends south into Italy). The two species can be found in sympatry in the south of France (Praz, in litt.), but to date I have examined no A. pusilla specimens from Iberia. Due to the confusion in this group, it must be considered absent until it can be positively demonstrated to be present south of the Pyrenees.

Figure 22. 

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♀, MNHN (lectotype; Fig. 21B).

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♀, MNHN (lectotype; Fig. 21C).

Andrena spreta: Algeria: Biskra [34.8600°N, 5.6995°E], 1♀, MNHN (lectotype; Fig. 21A).

Andrena (Micrandrena) strohmella Stöckhert, 1928 and Andrena (Micrandrena) icterina Warncke, 1974

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 (Gusenleitner and Schwarz 2002). Warncke (1974) later described A. icterina from northern Algeria. He later indicated the presence of this species in south-eastern Spain in his distribution maps (Gusenleitner and Schwarz 2002), though he did not list its present in Iberia (Warncke 1976). Wood et al. (2020b) later reported the presence of this species in northern Morocco. In his description of A. icterina, Warncke draws comparison with A. strohmella, noting that the process of the labrum is half as broad, the clypeus is more sparsely punctate and lacking an unpunctured midline, that the terga are more strongly shagreened, and that the carinae on the dorsolateral corners of the first tergum are more weakly pronounced. Dardón (2010) and Dardón et al. (2014) did not actually examine any material of Iberian A. icterina, but inspection of new Iberian material has demonstrated that A. icterina is more widespread in Iberia than previously thought, being found from the Los Alcornocales to the Sierra de las Nieves, the Sierra Nevada, Sierra de Baza, Sierra Cazorla, and Sistema Central, and is also newly reported from northern Portugal (see below). In all of these localities it is found in mountainous areas, generally at high altitude in excess of 1000 m.

Dardón (2010) and Dardón et al. (2014) reported A. strohmella from north-eastern Spain, from Tarragona: Alcanar, v.2002, leg. Kadlec, OÖLM. I have not been able to locate and examine this specimen for myself in the OÖLM collection, but it is slightly problematic for ecological regions. Andrena strohmella is clearly a species of deciduous temperate woodland; this is reflected in its distribution across Central Europe, and in its association with flowering trees such as Crataegus (Rosaceae), Prunus (Rosaceae), and Salix (Salicaceae). Andrena strohmella has therefore not previously been reported from Mediterranean regions. The record from Alcanar does not therefore match the ecology of this species, Alcanar being a low-elevation (c. 70 m) village close to the Mediterranean coast. However, A. strohmella is present in north-eastern Catalonia based on recently collected specimens (Álvarez Fidalgo, in litt.), and so the species is retained on the Spanish and Iberian lists.

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, RMNH; La Cabrilla, Sierra Cazorla (Jaén), 1600 m, 3.vi.2022, leg. C.M. Hererra, CMHC; Málaga, PN Sierra de las Nieves, mountain peak S of Pinsapo Escalereta, 1600 m, 30.v.2021, 3♀, leg. T.J. Wood, TJWC; Sierra Cazorla, Puerto Llano, 1800 m, 11.vi.2022, 1♀, leg. J. Valverde, CMHC; Puerto Crucetillas, Riópar (Albacete), 1340 m, 30.iv.2022, 1♂, leg. F.J. Ortiz-Sánchez, FJOS; Sierra de Cazorla, Nava de las Correhuelas, 1.v.2021, 1♂, leg. C.M. Hererra, CMHC.

Subgenus Notandrena Pérez, 1890

The definition of this subgenus was expanded by Pisanty et al. (2022b) to include Carandrena Warncke, 1968 (type species Andrena aerinifrons Dours, 1873). It is important to note that several species formerly placed within the Carandrena fall elsewhere into an undescribed subgenus, but these are desert-living species, none of which are present in Iberia. In the context of this revision, Carandrena is treated as a strict synonym of Notandrena.

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 Gusenleitner and Schwarz (2002: 1201) actually refers to A. (Notandrena) ungeri Mavromoustakis 1952; the distribution map for A. ungeri (p. 1200) is actually the distribution map for A. urdula. A barcoded specimen from south of Madrid [WPATW351-21] does not clearly fall into the Notandrena group (Fig. 23); its true placement is unclear because it does not have any strong characters that lead to obvious affinities with specific groups. It is best treated as incertae sedis until more genetic data are available.

Figure 23. 

Phylogenetic tree (maximum likelihood) of Andrena subgenus Notandrena Pérez, 1890 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).

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. 23). Moreover, A. reperta and A. reperta varuga do not form a monophyletic clade, being rendered paraphyletic by A. nigroviridula. On this basis, A. varuga stat. nov. is raised to species status. Andrena varuga is endemic to Spain, and A. reperta is found in Morocco, Algeria, and Tunisia.

Dours (1873) described A. aerinifrons from Algeria, and A. aerinifrons levantina Hedicke, 1938 was later described from the Levant. Barcode analysis shows that specimens from Israel, Morocco, Portugal, and Spain show high average intraspecific variation of 8.64% (range 0.00–11.67%). The Moroccan specimen is intermediate between the Iberian and Levantine sequences, being separated from the Iberian sequences by an average genetic distance of 9.92% (range 9.73–10.12%) and from the Levantine sequences by 10.12% (range 10.12–10.12%). Given that these sequences form a monophyletic clade, a broad interpretation of A. aerinifrons is taken here; a neotype can be designated for A. aerinifrons at a later date when the situation becomes clearer, as Dours’ collection was destroyed in a fire and is not available for study (see below).

Andrena (Notandrena) fulvicornis Schenck, 1861 has been recognised as distinct from A. (Notandrena) nitidiuscula Schenck, 1853 (Schmid-Egger and Doczkal 1995; Schmid-Egger and Scheuchl 1997; Schwenninger 2013; Bénon and Praz 2016). Barcode analysis shows that specimens from Germany, Israel, and Turkey are indeed clearly distinct from A. nitidiuscula, not forming a sister clade and showing average separation of 3.79% (range 3.50–3.89%), with bootstrap support of 96. However, sequences of putative A. fulvicornis from Spain fall into a clade of A. nitidiuscula and A. (Notandrena) chrysosceles (Kirby, 1802). Spanish specimens are separated from A. nitidiuscula by 1.95% (range 1.95–1.95%), from A. chrysosceles by 4.67% (range 4.28–5.06%), and from the other clade of A. fulvicornis by 4.18% (range 3.89–4.28%). This Iberian ‘fulvicornis’ is also well-supported, with bootstrap support of 96. At the present time, no taxonomic action can be taken until more genetic data are available. Morphologically, Iberian ‘fulvicornis’ do not appear to be different from Central European/Turkish/Levantine specimens. Additional sequences from Iberia and France are needed; this barcode difference may be the result of an isolated population in the Iberian glacial refugium.

Subgenus Plastandrena Hedicke, 1933

Pisanty et al. (2022b) synonymised the subgenus Agandrena with Plastandrena, and this broader definition is used here. Some taxonomic problems within this subgenus were discussed by Wood (2023a), who found support for the species-status of A. (Plastandrena) nigrospina Thomson, 1872. Standard barcodes have limited utility for this group for unclear reasons, so care must be taken when interpreting results, as morphologically distinct taxa can form unclear clades, for example A. (Plastandrena) tibialis (Kirby, 1802). This result was reproduced here (Fig. 9), and no further discussion or action is taken; in an Iberian context, members of this subgenus appear to be identifiable using barcodes, but work is needed to revise this group at a Palaearctic scale using more powerful genetic techniques.

Andrena (Plastandrena) pilipes Fabricius, 1781 and Andrena (Plastandrena) nigrospina Thomson, 1872

Ortiz-Sánchez et al. (2022) reported A. nigrospina as new for Spain from the Sierra de Cazorla based on male specimens with their distinctive genital capsule. Wood (2023a) further reported two barcoded specimens from Spain (Sistema Central and Sistema Ibérico) belonging to A. nigrospina. This genetic result was further supported with the addition of more sequences from Morocco (Fig. 9), revealing that A. nigrospina is also present in the Middle Atlas. The A. nigrospina clade maintained a low intraspecific genetic distance of 0.41% (range 0.00–1.06%) from Morocco to Kyrgyzstan, and average separation from A. pilipes was 1.63% (range 1.06–2.35%). As this difference is still small in absolute terms, bootstrap support was moderate, with 77 for A. pilipes and 84 for A. nigrospina, but both are considered to be distinct species. Females cannot currently be consistently separated morphologically, so no characters are given in the identification key. In Iberia, A. nigrospina is newly reported for northern Portugal from close to the Peneda-Gerês National Park. The species is likely to be restricted to cooler parts of northern, central, and eastern Iberia where it will probably be found only in mountainous regions, or at least at elevation. More sampling and genetic study is required to define its exact range limits.

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 (Plastandrena) agilissima Scopoli, 1770 and Andrena (Plastandrena) asperrima Pérez, 1895

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 Pérez (1895) were described based on North African material. Warncke (1967) designated a male lectotype for A. asperrima using a specimen from Biskra from the collection of Pic (Fig. 24A, B). This is unjustified, as it is not part of the original syntypic series, and Warncke’s designation is here rejected. In Pérez’s catalogue, under entry ‘1030 Andrena trachodes J.P.’ [an unpublished name; Pérez sometimes changed his mind and therefore a different name to the published name can be present in the catalogue] Pérez writes that the species comes from Constantine [in Algeria], drawing comparison with A. flessae Panzer, 1805 (= A. agilissima (Scopoli, 1770)) and arguing it differs by the stronger punctation of the metasoma. Inspection of the Pérez collection shows the presence of a female specimen from Constantine which bears the label ‘1030’, the code used by Pérez for this species. This specimen is designated as a new lectotype (Fig. 24C–F); it conforms to the concept of A. asperrima used by subsequent authors.

Figure 24. 

Andrena (Plastandrena) asperrima Pérez, 1895, false male lectotype A label details B profile; true female lectotype C label details D profile E scutum, dorsal view F terga, dorsal view.

Andrena atricapilla Pérez, 1895 was also described from Algeria, but only in the male sex. Warncke (1967) listed this taxon as a synonym of A. asperrima, though he did not mention a lectotype. A male specimen labelled as atricapilla J.P. in the handwriting of Pérez is separated in the MNHN collection and labelled as a lectotype, probably by Teunissen as one of his determination labels is present. This lectotype designation was never published, and so it is here designated as a lectotype (Fig. 25). The synonymy with A. asperrima is maintained, as the genital capsule is typical for the species concept.

Figure 25. 

Andrena (Plastandrena) atricapilla Pérez, 1895, male lectotype A label details B profile C terga, dorsal view D genital capsule.

Genetic analysis of specimens of A. agilissima and A. asperrima from Croatia, Morocco, Portugal, and Spain showed two clear clades (Fig. 9). One clade contained only specimens determined as A. agilissima from Croatia, Portugal, and Spain. These sequences showed average intraspecific variation of 0.07% (range 0.00–0.25%), and were strongly separated from the second clade by an average genetic distance of 9.33% (range 8.74–10.20%). This second clade contained specimens identified as A. asperrima from Spain and Morocco, but also three specimens identified as A. agilissima from the Middle Atlas. There can be no doubt as to the identity of these specimens, as two are males, and their genital capsule is that of A. agilissima and not A. asperrima. However, the genetic differentiation within this clade is very low at an average of 0.42% (range 0.00–0.91%). As for A. bimaculata and A. tibialis, there is clearly genetic complexity here, as two species with distinct genital capsules have barcodes that form intermixed clades. No taxonomic action is taken on the basis of these barcoding results.

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 Warncke (1967; 1974) and Gusenleitner and Schwarz (2002) regarding A. asperrima is maintained. The full synonymy is given below.

Andrena (Plastandrena) asperrima Pérez, 1895

Andrena (Plastandrena) asperrima Pérez, 1895: 33, ♀♂ [Algeria, lectotype by present designation: MNHN].

Andrena (Plastandrena) atricapilla Pérez, 1895: 33, ♂ [Algeria, lectotype by present designation: MNHN].

Andrena (Plastandrena) flessae var. elcheensis Friese, 1922: 211, ♀ [Spain: ZMHB, not examined].

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♀, MNHN (lectotype of A. asperrima, by present designation; Fig. 24C–F); Biskra, v.1885, 1♂, leg. Blause, MNHN (false lectotype of A. asperrima; Fig. 24A, B); Biskra [34.8600°N, 5.6995°E], 1♂, MNHN (lectotype of A. atricapilla, by present designation; Fig. 25); Morocco: Tizi-n-Talrhemt nr. Midelt [32.6821°N, -2.9344°E], 1900 m, 1.vi.1968, leg. M.A. Lieftinck, OÖLM (holotype of A. asperrima alascana).

Subgenus Poecilandrena Hedicke, 1933

This subgenus is strongly polyphyletic (Pisanty et al. (2022b), but these classification issues relate to eastern taxa; in Iberia, the species belong to Poecilandrena s. str. Warncke (1976) listed only one Poecilandrena species in Iberia, A. (Poecilandrena) labiata Fabricius, 1781. Ceballos (1956) and Pérez-Íñigo (1984) also listed A. (Poecilandrena) potentillae Panzer, 1809, though this species was not listed as present by Warncke and his distribution maps (Gusenleitner and Schwarz 2002) indicate that this species has a western range limit in central France.

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. 1). The four A. labiata sequences from Belgium, Portugal, and Spain showed average intraspecific variation of 2.52% (range 0.78–3.92%), much lower than the average separation from A. potentillae sequences from Belgium, France, Germany, and Hungary of 11.81% (range 11.23–12.27%). The position is taken here that true A. potentillae is not present in Iberia, even though some females of A. labiata can resemble it morphologically. These specimens conform to the subspecific concept of A. labiata bellina Warncke, 1967 that was described from Madrid. Warncke (1967) noted that this form had weaker shagreenation and finer punctation than the nominate form, and I believe that this is the source of the confusion and the erroneous reports of A. potentillae in Iberia. Andrena potentillae is therefore not included in the identification key as typical female characters that can be used to separate the two species in Central Europe do not appear to work universally in Iberia.

Finally, Ortiz-Sánchez (2011, 2020) lists A. (Poecilandrena) viridescens Viereck, 1916 as part of the Spanish fauna. The distribution maps of Warncke (Gusenleitner and Schwarz 2002) indicate the possible presence of this species on the Spanish side of the Pyrenees. I have not examined any specimens of A. viridescens from Iberia, but its presence in the Pyrenees is plausible and so it is retained on the list. Several Andrena taxa with apparent range limits in the Pyrenees have recently been confirmed to occur in northern Spain (e.g. A. (Ulandrena) polita Smith, 1847 and A. (Micrandrena) nanula Nylander, 1848, see Álvarez Fidalgo et al. 2022), and so additional searches may well confirm the presence of A. viridescens as well.

Subgenus Simandrena Pérez, 1890

Barcode analysis returned Simandrena as paraphyletic (Fig. 4), but this means very little, as the subgenus is very well characterised morphologically and genetically based on UCE analysis (Pisanty et al. 2022b). No major changes in Simandrena taxonomy are made here, but there are a number of issues to discuss.

Warncke (1967) described A. (Simandrena) combinata crudelis Warncke, 1967 from Spain. Comparison of sequences shows that Iberian material is only weakly differentiated, being separated from A. (Simandrena) combinata (Christ, 1791) sequences from Germany by an average genetic distance of 1.12% (range 1.06–1.29%). Based on this evidence, a subspecific status is not justified.

Andrena (Simandrena) vetula Lepeletier, 1841 was recently placed in the Simandrena, as its unusual male morphology had led to confused previous placement (Pisanty et al. 2022b). Genetically there was a large difference between Iberian and North African sequences, separated by an average genetic distance of 6.33% (range 6.08–6.57%). Andrena vetula has an enormous range, from Morocco and Iberia to Central Asia (Wood and Monfared 2022). Sequences are required over this range before any taxonomic changes can be made, as morphologically there are no obvious differences between Iberian and North African specimens.

The species pair of A. (Simandrena) confinis Stöckhert, 1930 and A. (Simandrena) congruens Schmiedeknecht, 1884 continues to pose problems. Warncke (1967) treated A. confinis as a synonym of A. congruens, but others have not followed this interpretation. The two taxa can be separated morphologically in Central Europe (Schmid-Egger and Scheuchl 1997), and there are also ecological differences with A. confinis preferring cooler northern climates and A. congruens preferring warmer and drier southern climates, with overlap in Central Europe. However, in southern Europe the situation is complex and confused. Wood et al. (2021) added A. confinis to the Iberian list on the basis of a specimen from the Picos de Europa Mountains. Genetically, this specimen clusters with A. confinis sequences from Germany. However, sequences from specimens provisionally identified as A. congruens from northern Portugal as well as a specimen of A. congruens from Bulgaria render A. congruens paraphyletic. The genetic distance between all sequences is low. The Portuguese specimens are strongly divergent morphologically from the Spanish A. confinis specimen, with dense and obvious tergal punctation compared to terga which are almost impunctate, but the average genetic differentiation is low at 0.88% (range 0.60–1.15%). No major taxonomic decision is made here; much more genetic data are required for a dedicated study focused on this group. Both nominal taxa are included in the identification key.

Andrena (Simandrena) cilissaeformis Pérez, 1895, sp. resurr

Andrena (Simandrena) cilissaeformis Pérez, 1895: 42, ♀ [Spain, lectotype by present designation: MNHN]

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. 26) and listed the taxon as a synonym of A. numida Lepeletier, 1841 (Warncke 1967). However, just a few years later he listed A. breviscopa as a valid taxon in the subgenus Simandrena (Warncke 1974). Examination of the female lectotype designated by Warncke shows that his original synonymy was correct; the taxon is clearly not a Simandrena, and is indeed a synonym of A. numida.

Figure 26. 

Andrena (incertae sedis) breviscopa Pérez, 1895, female lectotype A label details B profile C face, frontal view D terga, dorsal view.

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 Warncke (1967) and Gusenleitner and Schwarz (2002). This is because Pérez (1895) does not directly state the type locality (or indeed, any information about the collecting locality of any of the species described in this work), but this information is included in his personal catalogue. Warncke (1967) did not examine material of A. cilissaeformis, stating that whilst the label was present, material was missing. Examination of material in the MNHN has located a specimen labelled with ‘Esp’ [Espagne = Spain] in Pérez’s handwriting (Fig. 27). This specimen was separated by Teunissen, but he never published this information. This specimen is badly damaged, but it is a Simandrena and conforms to A. breviscopa auctorum sensu Warncke (1974) and subsequent publications. It is designated as a lectotype to fix the name on the Iberian population. Andrena breviscopa is returned to its original synonymy with A. numida.

Figure 27. 

Andrena (Simandrena) cilissaeformis Pérez, 1895; female lectotype A label details B profile C face, frontal view D terga, dorsal view.

Distribution. Spain, Morocco, Algeria.

Material examined. Andrena breviscopa: Algeria: Ghardaia [32.5047°N, 3.6419°E], 1♀, MNHN (lectotype; Fig. 26); (Andrena cilissaeformis): Spain: no collection information, 1♀, MNHN (lectotype by present designation; Fig. 27).

Subgenus Taeniandrena Hedicke, 1933

Large parts of this subgenus have been revised recently by Wood et al. (2021), Praz et al. (2022), Wood (2022), and Wood and Ortiz-Sánchez (2022). There is relatively little new information to present here, other than to revise the status of A. (Taeniandrena) poupillieri Dours, 1872, and so a reduced phylogenetic tree is presented given the results presented in these previous publications (Fig. 28). However, it is clear that there is still unfinished work to be completed in this subgenus in an Iberian context. Specifically, Praz et al. (2022) identified an unclear lineage “sp. nov. 2” from northern Portugal. This specimen is a male that has a genital capsule that diverges from any known Iberian species. Furthermore, I have an unbarcoded male from Guadalajara province that also possesses a genital capsule that does not match any known species. These may represent additional undescribed Taeniandrena species endemic to the peninsula. No further action is taken until more genetic samples are available; it is extremely likely that additional barcoding will discover more diversity in this challenging and speciose subgenus.

Figure 28. 

Phylogenetic tree (maximum likelihood) of Andrena subgenus Taeniandrena Hedicke, 1933 based on the mitochondrial COI gene. Andrena (Taeniandrena) lathyri Alfken, 1900 is used as an outgroup. Numbers adjacent to branches represent bootstrap support (values of <75 are omitted).

Andrena (Taeniandrena) poupillieri Dours, 1872

Two further issues require discussion. The first is the identity of A. poupillieri. Praz et al. (2022) identified two potential mitochondrial lineages that could correspond to this species. Additional sampling in Morocco has clarified the situation; poupillieri 1 sensu Praz et al. (2022) corresponds to A. (Taeniandrena) gregaria Warncke, 1974 and poupillieri 2 sensu Praz et al. (2022) corresponds to the concept of A. poupillieri used by Warncke. As the type of A. poupillieri is lost, it is beneficial to designate a neotype in order to fix the concept of this species in line with the existing literature; a specimen from Algeria is chosen, as this is the original locus typicus. As A. gregaria does not occur in Iberia, it is not included in the phylogenetic tree presented here; it will be dealt with in the upcoming North African Andrena revision. The species A. poupillieri is commonly encountered in Morocco, and is clearly identifiable from barcodes, forming a clade with bootstrap support of 100. Separation of females from A. (Taeniandrena) ovatula (Kirby, 1802) can be made by the tergal punctation, this being much stronger in A. ovatula. Generally, the two taxa are well-separated by an average genetic distance of 6.12% (range 5.86–7.79%). Males of A. poupillieri can be recognised based on the genital capsule, in which the gonostyli are apically produced into acute points.

Warncke changed his mind about the status of A. poupillieri – in Warncke (1967), he lists the taxon as A. ovatula poupillieri, but then in Warncke (1975a) he described A. poupillieri incana from the Balearic islands, listing A. poupillieri s. str. from southern Iberia (Warncke 1976). In his distribution maps (Gusenleitner and Schwarz 2002), he gives a distribution of southern Iberia, the Balearic islands, north-western Africa, Libya, and Crete. Records from Crete are likely to refer to unrecognised A. (Taeniandrena) ovata Schenck, 1853 (Wood, unpublished data). Examination of male specimens from the extreme south of Spain shows that A. poupillieri is present based on the distinctive genital capsule, though as females cannot be separated morphologically from A. ovatula the exact range of the two species is unclear. Andrena ovatula reaches at least as far south at the Sierra de las Nieves in Málaga province [IBIHM1045-22], though this far south it may be restricted to mountainous areas whereas the two examined male A. poupillieri specimens come from the coast. More collection is required. For now, A. poupillieri is considered to be a rare and little-collected taxon in Iberia, probably restricted to the coast in the south and south-east of the peninsula. A single sequence for A. p. incana was available from Mallorca that unambiguously nests within the North African A. poupillieri sequences; it is identical to 12 of the North African sequences, and differs from the thirteenth by just 0.15%. As such, A. poupillieri including A. p. incana shows extremely low intraspecific variation of 0.03% (range 0.00–0.15%), and subspecific status is unnecessary for the population on the Balearic islands.

Material examined. Spain: Málaga, Estepona, 24.iii.1986, 1♂, leg. J. van Oosterhout, RMNH; Málaga, San Pedro de Alcántara, 15.iii.1986, 1♂, leg. C. v. Achterberg, RMNH.

Andrena (Taeniandrena) russula Lepeletier, 1841

Praz et al. (2022) synonymised A. (Taeniandrena) similis Smith, 1849 with A. russula, taking a broad, pan-Mediterranean approach. In an Iberian context, sequences from central and northern Iberia clearly fall into a broad A. russula clade along with sequences from Morocco to Cyprus and Israel and north to the United Kingdom and Finland (Fig. 28). However, in the extreme south-west of Iberia in southern Portugal, sequences from specimens that are morphologically indistinguishable from A. russula from the rest of Iberia form a distinct clade that falls closest to A. (Taeniandrena) afzeliella (Kirby, 1802). These specimens come from the south of Baixo Alentejo (Cercal) and the Algarve (Aljezur, Benagil, Moncarapacho). These sequences are consistently separated from the broad A. russula clade by 5.50% (range 3.68–11.11%). Two specimens from Morocco that correspond to A. russula form a third clade of A. russula s.l. At the present time, no taxonomic action is taken; these three lineages may represent different isolated populations. More powerful genetic techniques are required to resolve this problem, as for Iberian members of the gelriae-group (see Praz et al. 2022).

Andrena (Taeniandrena) gredana Warncke, 1975

Wood et al. (2021) elevated A. gredana to species status, and gave a distribution across the Sistema Central, central and northern Portugal, and northern Spain across to the Pyrenees in the province of Huesca. Examination of material from the Hautes-Pyrénées in France approximately 80 km north-east of the Huesca site revealed the presence of A. gredana in France. The species is therefore not endemic to the Iberian Peninsula, though it is likely to have a French range restricted to high altitude sites in the Pyrenees.

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, RMNH (see also records in Wood et al. 2021).

Subgenus Truncandrena Warncke, 1968

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.

Andrena (Truncandrena) doursana Dufour, 1853 sensu lato

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 Warncke (1975a, 1975b, 1980) described the subspecies A. d. citreola Warncke, 1975 from Spain, A. d. agadira Warncke, 1980 from southern Morocco, A. d. bengasia Warncke, 1980 from Libya, and A. d. mizorhina Warncke, 1975 from Turkey. Pisanty et al. (2022b) elevated the eastern subspecies A. (Truncandrena) doursana mizorhina to species status, leaving populations in Iberia and north-western Africa to Libya.

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. Wood et al. (2020b) found the undescribed female of a similar species endemic to Morocco, A. (Truncandrena) alchata Warncke, 1974, which is structurally very similar to the female of A. d. agadira, but these authors were unable to conclude if the two taxa were synonymous.

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. 29). Light forms from southern Portugal [IBIHM1223-22] and the Moroccan Anti-Atlas near Tazenakht [WPATW505-22] were separated, and dark forms from south-western Morocco near Guelmim [WPATW506-22] and Tiznit [WPATW503-22] were clustered closer to the light specimen from Tazenakht. The light specimen from Portugal was clustered close to a specimen of A. doursana s. str. from the Middle Atlas near Taza [WPATW509-22]. This pattern strongly suggests simple separation by distance, with geographically closer specimens displaying more similar barcode sequences. Accepting a broad A. doursana species concept, these sequences showed an average intraspecific distance of 2.79% (range 0.29–5.93%).

Figure 29. 

Phylogenetic tree (maximum likelihood) of Andrena subgenus Truncandrena Warncke, 1968 based on the mitochondrial COI gene. Andrena (Limbandrena) limbata Eversmann, 1852 is used as an outgroup. Numbers adjacent to branches represent bootstrap support (values of <75 are omitted).

More broadly, this A. doursana clade had bootstrap support of 78, and was sister to the A. alchata sequences generated by Wood et al. (2020b), a species that has a clearly different male morphology. Andrena alchata has bootstrap support of 91, and was separated from A. doursana by an average of 6.29% (range 5.62–8.03%). Finally, the single sequence of A. mizorhina was strongly separated from A. doursana by an average of 10.45% (range 10.18–10.94%). These genetic results justify the decision of Pisanty et al. (2022b) to elevate A. mizorhina to species level, the findings of Wood et al. (2020b) who identifed the female of A. alchata, and the original description of A. d. agadira by Warncke as a subspecies, correctly associating dark specimens from south-western Morocco with A. doursana. Given the variation in colour forms observed here, no subspecies framework is employed, and so the Iberian taxon is referred to simply as A. doursana.

Andrena (Truncandrena) medeninensis Pérez, 1895 sensu lato

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. Warncke (1967, 1974, 1980) described several subspecies, A. m. donata Warncke, 1967 from Spain, A. m. abunda Warncke, 1974 from Morocco, A. m. tiznita Warncke, 1980 from south-western Morocco, and A. m. usura Warncke, 1967 from Turkey. Wood (2023b) synonymised A. m. usura with A. pareklisiae Mavromoustakis, 1957, leaving the North African and Iberian populations.

Sequences of A. medeninensis s. str. and A. m. tiznita formed a clade with bootstrap support of 91 (Fig. 29). However, the single available sequence of A. medeninensis s. str. differed from A. m. tiznita sequences by an average of 4.80% (range 4.70–4.85%), A. m. tiznita forming a subclade with bootstrap support of 94. However, A. m. abunda was strongly separated from this clade of A. medeninensis s.l. by an average genetic­ distance of 10.19% (range 9.97–10.57%). This A. m. abunda clade had bootstrap support of 100, and was sister to an A. medeninensis s.l. + A. tscheki Morawitz, 1872 s.l. clade. Andrena abunda stat. nov. is therefore raised to species status; it restricted to Algeria and Morocco. Morphologically, it is extremely similar to A. medeninensis s.l., but has much darker pubescence. In the female sex (the male is unknown), the only clear structural difference is that A3 exceeds the length of A4+5+6, whereas in A. medeninensis s.l. A3 is shorter than A4+5+6.

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 (Truncandrena) truncatilabris Morawitz, 1877, Andrena (Truncandrena) truncatilabris espanola Warncke, 1967, and Andrena (Truncandrena) truncatilabris nigropilosa Warncke, 1967

Andrena truncatilabris is a widespread species that was originally described from the Caucasus from what is today Armenia (Astafurova et al. 2021). The species is nominally distributed across the Mediterranean basin, from Morocco and Iberia to the Urals and Iran (Gusenleitner and Schwarz 2002). However, in the east it descends only to the Levant and does not enter the eastern part of North Africa. In the west, Warncke (1967) described two subspecies: A. t. espanola from Spain and A. t. nigropilosa from Algeria that differed from A. truncatilabris s. str. in the structure of their clypeus and their male genital capsule, though Warncke noted that the difference between the two subspecies was minimal, pointing to the darker pubescence of North African specimens as a point of difference.

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. 29). They were strongly separated from A. truncatilabris s. str. sequences from northern Israel by an average genetic distance of 9.66% (range 9.12–10.12%). Andrena truncatilabris s. str. formed a clade with bootstrap support of 99, and was sister to A. (Truncandrena) ferrugineicrus Dours, 1872, whereas A. t. nigropilosa + A. t. espanola formed a clade with bootstrap support of 100 that was sister to the A. truncatilabris s. str. + A. ferrugineicrus clade.

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) nigropilosa Warncke, 1967, stat. nov

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.

Andrena (Truncandrena) villipes Pérez, 1895

Pérez (1895) described A. villipes from north-eastern Spain (Fig. 30), later describing the synonymous A. (Truncandrena) squalida Pérez, 1903 from south-western France. It has a restricted distribution, known from Cistaceae-rich habitats from northern Morocco, Portugal, Spain, southern France, and north-western Italy (Gusenleitner and Schwarz 2002; Lhomme et al. 2020). Searches in the Sierra de las Nieves in southern Spain produced two particularly large and dark specimens provisionally identified as A. villipes. A sequence from one of these specimens is separated from A. villipes sequences from southern Portugal by an average genetic distance of 11.85% (range 11.85–11.85%). This new species is described and diagnosed below.

Figure 30. 

Andrena (Truncandrena) villipes Pérez, 1895; female lectotype A label details B profile C face, frontal view D dorsal view.

Material examined. Spain: Barcelona [41.4028°N, 2.1332°E], 1♀, MNHN (lectotype of A. villipes; Fig. 30).

Undescribed subgenera

As a result of the ground-breaking analysis of Pisanty et al. (2022b), we now have unprecedented phylogenetic resolution for the genus Andrena, and are now able to seriously deal with the problems inherent to the subgeneric classification system largely solidified by Warncke (1968a). Building on the work of Pisanty et al. (2022b), the Iberian fauna contains representatives of 44 described subgenera, but also representatives for nine additional clades that are currently undescribed. Some of these are dealt with here, though the aegyptiaca-group (Iberian representative: A. alluaudi Benoist, 1961) is not treated here; this lineage will be described in a planned future comprehensive revision of the subgenus.

The former Poliandrena Warncke, 1968

Pisanty et al. (2022b) demonstrated that Andrena polita Smith, 1847 falls into an expanded Ulandrena, and that Poliandrena is a strict synonym of this subgenus, and additionally that Poliandrena sensu Warncke contains at least five different polyphyletic clades, with the possibility of additional distinct clades when taxonomic sampling and phylogenetic analysis is more complete. Warncke, who described the subgenus Poliandrena, understandably used it as a ‘waste-basket’ for unclear taxa that displayed no clear defining characters. It is therefore necessary to split up this old grouping and to describe new subgenera.

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 Pisanty et al. (2022b) which is based on UCE analysis. UCE analyses are based on thousands of loci, and hence offer a high degree of confidence that groups are or are not related. Whilst single locus COI analyses cannot compare to those based on UCEs, they can demonstrate if individual species are closely related, and hence complement a UCE-informed phylogeny. No members of the oviventris-group were included in the analysis of Pisanty et al. (2022b), but analysis of barcodes (Fig. 31) places A. farinosa Pérez, 1895, A. farinosoides Wood, 2020, and A. oviventris Pérez, 1895 in a single clade with bootstrap support of 99, well-separated from A. corax Warncke, 1975, A. murana Warncke, 1975, and A. relata Warncke, 1975 which belong to the relata-group and which are morphologically the most similar to members of the oviventris-group. The two clades are separated by A. limbata Eversmann, 1852. On the basis of the analysis of Pisanty et al. (2022b) combined with these barcodes, subgenera are described for the blanda-group, florea-group, limbata-group, and oviventris-group.

Figure 31. 

Phylogenetic tree (maximum likelihood) of Andrena from currently undescribed subgenera 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).

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.

The caroli-group

Members of this group of species have been placed in the subgenus Campylogaster Dours, 1873 (Warncke 1968a) that has the type species A. erberi Morawitz, 1871. Pisanty et al. (2022b) identified that Campylogaster is polyphyletic, with A. caroli Pérez, 1895 falling far away from A. erberi. Campylogaster therefore can probably only be applied to the species around A. erberi that have strongly depressed tergal margins, e.g. A. iranella Popov, 1940 (Iran, Turkemenistan, ?Arabian Peninsula), A. nanshanica Popov, 1940 (China, Mongolia), A. chengtehensis Yasumatsu, 1935 (China, South Korea). This subgenus therefore appears to be eastern and predominantly Asian, with a western limit of A. erberi in the southern Balkans. In contrast, A. caroli has the tergal margins flat and not noticeably depressed. Based on the analysis of Pisanty et al. (2022b) combined with this morphological difference, a new subgenus Pruinosandrena subgen. nov. is described below for the species around A. caroli. Importantly, A. lateralis Morawitz, 1876 and A. incisa Eversmann, 1852 were also placed into Campylogaster by Warncke (1968a). Although not sampled by Pisanty et al. (2022b), A. lateralis falls far away from the Pruinosandrena based on COI analysis (Fig. 32). Morphologically, A. lateralis and A. incisa form a species pair (that can be referred to at the incisa-group) that lacks many of the characters shared by members of the Pruinosandrena (see below), and they probably represent an additional undescribed subgenus. Without additional genetic data, no further steps are taken for this species pair at the present time.

Figure 32. 

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. Erichson (1835) described A. pruinosa from southern Spain (Andalusia); a single female labelled as ‘type’ is conserved in the ZMHB which may automatically be the holotype (Fig. 33), but this is ambiguous as it is unclear if Erichson described the species from multiple specimens or not. Dours (1872) described A. succinea from Algeria, noting the clear red colouration of the metasoma (hence the species name, succin = amber). Warncke (1967) used A. succinea as a subspecies in combination with A. pruinosa, arguing that males from North Africa could not be clearly separated from Spanish males. He then described A. pruinosa parata from south-eastern Spain, giving characters related to colouration and antennal ratios.

Figure 33. 

Andrena (Pruinosandrena) pruinosa Erichson, 1835, female holotype/syntype A label details B head, dorsal view C mesosoma, dorsolateral view D dorsal view.

Genetic analysis of members of the Pruinosandrena demonstrates that the broad concept of A. pruinosa used by Warncke was overly conservative (Fig. 32). A single red-marked female specimen initially identified as A. pruinosa was separated by 7.45% from three additional A. pruinosa s. str. specimens. Inspection of Warncke’s original description of A. pruinosa parata and comparison of the antennal ratios demonstrates that this divergent specimen has A3 clearly longer than A4+5, whereas A3=A4+5 in A. pruinosa s. str. The other characters mentioned by Warncke relating to colour do not work consistently, as one of the barcoded A. pruinosa s. str. has the terga partially red-marked and the hind tibiae lightened orange. Though small, this morphological difference is meaningful, as the two taxa are found essentially in direct sympatry; the distance between the sampling location of barcoded specimens of A. pruinosa s. str. from Pinto was approximately 11 km from the Camino de Uclés sampling site for A. pruinosa parata. In this context, a genetic separation of 7.45% combined with the morphological difference is highly significant, and A. parata stat. nov. is treated as a valid species.

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 Wood et al. 2020b), with the yellow markings sometimes extending onto the lower part of the paraocular areas (uniformly black in A. pruinosa s. str. and A. parata) and, viewed ventrally, A4 is short, as long as broad (A4 is elongate and clearly longer than broad in A. pruinosa s. str.). Andrena parata males can easily be separated as A3 is longer than A4+5, whereas A3 is shorter than A4+5 in both A. pruinosa s. str. and A. succinea.

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. 34A, B), whereas it has never been recorded from the more humid parts of the Middle Atlas that have grasslands resembling those that can be found in central Iberia (e.g. Madrid, north of Chinchón, Fig. 34C, D).

Figure 34. 

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 MNHN collection (A. sitifensis Pérez, 1895; Fig. 35), A. succinea is the name that is established in the literature, either as a species itself or in combination with A. pruinosa (Benoist 1961; Warncke 1967; Warncke 1974; Gusenleitner and Schwarz 2002; Wood et al. 2020b; Dermane et al. 2021). In order to conserve this use, a neotype is designated below for A. succinea. As a result of these numerous changes, the updated statuses and synonymies are given here:

Figure 35. 

Andrena (Pruinosandrena) sitifensis Pérez, 1895, female lectotype A label details B profile C scutum, dorsal view D terga, dorsal view.

Andrena (Pruinosandrena) parata Warncke, 1967, stat. nov

Andrena pruinosa parata Warncke, 1967: 233, ♀♂ [Spain: OÖLM, examined].

Remarks. Though described from south-eastern Spain, Warncke (1976) noted that he had examined a single specimen of A. parata (as A. pruinosa parata) from Madrid: Ribas de Jarama, but he expressed doubts as to whether or not it had been correctly labelled given the collecting localities of all other known specimens. The contemporary Camino de Uclés site is approximately 7 km from Ribas de Jarama, confirming the presence of this taxon in central 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 (Pruinosandrena) pruinosa Erichson, 1835

Andrena pruinosa Erichson, 1835: 104, ♀ [Spain: ZMHB, examined].

Andrena lanuginosa Spinola, 1843: 137, ♀ [Spain, lectotype by present designation: MRSN].

Remarks. Examination of the type material of both A. pruinosa and A. lanuginosa (Fig. 36) showed that both were female specimens with dark terga, conforming to the classical concept of this species. Neither represent A. parata, and hence the synonymy of A. lanuginosa with A. pruinosa is maintained. Spinola (1843) did not specify how many specimens he described the species from. The specimen examined here may be automatically the holotype, but as this is not clear from the original description, it is here designated as the lectotype.

Figure 36. 

Andrena (Pruinosandrena) lanuginosa Spinola, 1843, female lectotype A label details B profile C head, dorso-frontal view D terga, dorsal view.

Distribution. Spain.

Material examined. Spain: Andalusia, 1♀, leg. Waltl, ZMHB (holotype/syntype; Fig. 33); Andalusia, 1♀, leg. Ghilinni, MRSN (lectotype of A. lanuginosa, by present designation; Fig. 36); Madrid, Madrid, Chinchón, 6 km N, M-311, 14.v.2021, 2♂, 1♀, leg. T.J. Wood, TJWC; Madrid, Madrid, Pinto, 1 km E, Carr. la Marañosa, 13.v.2021, 1♂, 1♀, leg. T.J. Wood, TJWC.

Andrena (Pruinosandrena) succinea Dours, 1872, stat. nov

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: MNHN, examined]

Andrena fulvisquama Popov, 1940: 260, ♀ [Algeria: ZISP, not examined]

Remarks. The synonymy of A. mayeti Pérez, 1895 (described from Tunisia) with A. succinea reported by Warncke (1967, see also Benoist 1961) is incorrect. Examination of the female lectotype (Fig. 37) shows that A. mayeti syn. nov. is a synonym of A. caroli Pérez, 1895 (described from Algeria; Fig. 38) because the foveae are wide and there are well-developed hair bands on the tergal margins (see identification key for Pruinosandrena below).

Figure 37. 

Andrena (Pruinosandrena) mayeti Pérez, 1895, female lectotype A label details B profile C face, frontal view D terga, dorsal view.

Figure 38. 

Andrena (Pruinosandrena) caroli Pérez, 1895, female lectotype A label details B profile C face, frontal view D terga, dorsal view.

Distribution. Morocco, Algeria, Tunisia, Libya, Egypt, Israel and the West Bank, Jordan, Syria, Saudi Arabia, Iran (Wood and Monfared 2022).

Material examined. Algeria: Setif [36.2059°N, 5.3965°E], 1♀, MNHN (lectotype of A. sitifensis; Fig. 35); 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] (neotype of A. succinea, see below).

Andrena (Pruinosandrena) caroli Pérez, 1895

Andrena (Pruinosandrena) caroli Pérez, 1895: 47, ♀ [Algeria: MNHN, examined]

Andrena (Pruinosandrena) mayeti Pérez, 1895: 47, ♀ [Tunisia: MNHN, examined] syn. nov.

Distribution. Morocco, Algeria, Tunisia, Egypt, Israel.

Material examined. Algeria: Biskra [34.8600°N, 5.6995°E], 1♀, MNHN (lectotype of A. caroli; Fig. 38); Tunisia: Chott el Djerid [33.8806°N, 8.1435°E], 1♀, MNHN (lectotype of A. mayeti; Fig. 37).

The numida-group

This group of Palaearctic species was previously placed in the subgenus Thysandrena Lanham, 1949 by Warncke (1968a). However, true Thysandrena occur only in North America, and Palaearctic species fall elsewhere (Pisanty et al. 2022b). Four species are recognised in the Palaearctic; A. hypopolia Schmiedeknecht, 1884, A. numida, A. ranunculorum Morawitz, 1877, and A. lunata Warncke, 1975, though additional genetic work is required to ensure that they all belong together; no subgenus is therefore described in the current work.

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 Gusenleitner and Schwarz 2002), southern Italy (ssp. syracusae Strand, 1921), and Turkey (ssp. albiscopa Warncke, 1967, see Gusenleitner and Schwarz 2002). No members of this group are known from the Levant.

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 RMNH collection, see below), and the type of A. numida can also not be located at the MNHN. Furthermore, A. inconspicua Morawitz, 1871 was described from Calabria in southern Italy. Warncke (1967) treated this as A. (Taeniandrena) russula Lepeletier, 1841 based on the description; Gusenleitner and Schwarz (2001) then listed it as a synonym of A. numida, before finally Gusenleitner and Schwarz (2002) listed it as a synonym of A. hypopolia despite the fact that it is an older name. This is because, under the broad treatment of Warncke, A. inconspicua would be junior to A. numida. However, Gusenleitner and Schwarz (2002) treated A. hypopolia as a distinct species, and did not resolve the status of A. inconspicua, even though it is an older name than A. hypopolia. The lectotype of A. inconspicua was recently illustrated by Astafurova et al. (2021), confirming its affinity with the numida-group, and not the subgenus Taeniandrena. The use of the name A. inconspicua therefore depends on two things; 1) whether European and North African material are distinct, and 2) whether material from southern Italy shows a stronger affinity with the European or North African taxon.

Genetically, barcoded specimens from Iberia showed almost no differentiation from specimens from Germany (average genetic distance 0.26%; Fig. 1), thus demonstrating that the name A. hypopolia can be applied to both the south-western and Central European populations. However, there was a consistent separation between Moroccan and German/Iberian sequences, these separated by an average genetic distance of 3.05% (range 2.87–3.13%). A single sequence was available from Sicily. This sequence differed from German/Iberian sequences by an average genetic distance of 6.11% (range 3.87–6.82%) and from Moroccan sequences by an average of 2.39% (range 2.17–2.64%). It clustered with Moroccan sequences, with the clades Morocco+Sicily showing bootstrap support of 92 and Germany+Iberia showing bootstrap support of 86. Examination of these barcoded specimens shows that there is a subtle but consistent morphological difference, which is the density of punctures on T3 of females. In Iberian specimens, the disc of T3 is densely punctate, with punctures separated by <1 puncture diameter. In contrast, in Moroccan and Sicilian specimens, the disc of T3 is shallowly and obscurely punctate, with punctures separated by >1 puncture diameter. Examination of the female lectotype of A. inconspicua (Astafurova et al. 2021: fig. 24) shows that this specimen morphologically conforms to A. numida, with weak and obscure punctures on the disc of T3. On this basis, A. inconspicua is synonymised syn. nov. with A. numida, and A. hypopolia is maintained as the senior name for populations in Iberia and Central Europe. A barcoded neotype of A. numida is designated from Moroccan material (see below).

This action largely maintains the status quo of Gusenleitner and Schwarz (2002), and A. numida is considered to have a distribution of Morocco, Algeria, Tunisia, Libya, and Sicily and the extreme southern parts of Italy (Calabria, Campania). The inclusion of Campania derives from treatment of the taxon A. syracusae. This taxon was described by Strand in the male sex only as a variety of A. (Simandrena) propinqua Schenck, 1853. Warncke (1967) treated this as a subspecies of A. numida with dark facial hair. The barcoded specimen from Sicily is a female, and whilst it should represent A. syracusae since it was collected only 150 km west of the locus typicus of Syracuse, since the female of A. syracusae is unknown and undescribed this cannot be said for certain. I have not seen male A. numida material from Sicily, or the type material of A. syracusae itself. However, I have seen males from the island of Ischia by Naples. These conform to the concept of A. syracusae, with extensive dark facial pubescence, and show extremely weak and sparse punctation on T3. The island of Ischia and the nearby Sorrento Peninsula on the Italian mainland are unusual and host an Andrena fauna with several species that are otherwise found in Italy only in Sicily, such as A. (Micrandrena) fumida Pérez, 1895. On the basis of its morphology, A. syracusae is considered to represent only a form of A. numida. This should ultimately be confirmed through the direct barcoding of males with dark facial pubescence; these are expected to fall into the A. numida clade.

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. 39; des. Warncke 1967) reveals that it is not a synonym of A. numida hypopolia as given by Warncke (1967), but a synonym (syn. nov.) of Andrena ranunculorum. The specimen comes from Arreau in France which is in the Hautes-Pyrénées department, and is situated immediately adjacent to mountains that ascend to 3,000 m. Andrena ranunculorum is known from alpine habitats in the central and eastern Pyrenees on both the French and Spanish sides, so this synonymy makes both morphological and ecological sense.

Figure 39. 

Andrena (incertae sedis) setosa Pérez, 1903, female lectotype A label details B profile C face, frontal view D terga, dorsal view.

Material examined. Andrena ranunculorum: France: Arreau [42.9064°N, 0.3557°E], 1♀, MNHN (lectotype; Fig. 39).

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, MRSN; Sorrento [Naples], 3.v.1970, 1♂, leg. J.P. van Lith, RMNH.

Description of new subgenera

Blandandrena subgen. nov.

Type species

Andrena blanda Pérez, 1895.

Diagnosis

Blandandrena is monotypic, and hence diagnosis of A. blanda (Fig. 40) is de facto diagnosis of the subgenus. Through the combination of slightly upturned fore margin of the clypeus, fovea broad and occupying over 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), lack of squamous hairs, 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 Ovandrena subgen. nov. that were formerly lumped together under the subgenus Poliandrena (see above). Andrena blanda females can provisionally be separated from the relata-group by the weakly punctate terga, punctures shallow and somewhat obscure, separated by 1–2 puncture diameters (Fig. 40D) whereas in the relata-group, tergal punctures are typically much stronger, clear and dense, separated by 1–2 puncture diameters but often by only 1 puncture diameter. This character works for West Mediterranean members of the relata-group, but additional work is needed to define this subgenus in the east, and so it is not defined and described here. Andrena blanda females can be separated from the Ovandrena by their simple scutal hairs (Fig. 40A–C; hairs semi-squamous in Ovandrena) and poorly delineated propodeal triangle that lacks lateral carinae (Fig. 40C; propodeal triangle clearly defined by presence of lateral carinae in Ovandrena; see below for additional detail).

Figure 40. 

Andrena (Blandandrena) blanda Pérez, 1895, female A profile B face, frontal view C propodeum, dorsal view D terga, dorsal view; male E profile F face, frontal view G terga, dorsal view H genital capsule.

Andrena blanda males can be separated by their black clypeus (Fig. 40F) in combination with their distinctive genital capsule (Fig. 40G). Most members of the relata-group have a yellow-marked clypeus, but for those with a black clypeus (e.g. A. corax), the genital capsule allows recognition. In A. blanda, the genital capsule is short and compact, more or less round, with gonocoxae with apically diverging inner margins and which are produced into short pointed teeth, and the gonostyli are apically strongly broadened and flattened, the disc being slightly broader than long. In the relata-group, the genital capsule is typically simple, without strongly apically broadened gonostyli, or when this is the case (e.g. A. murana Warncke, 1975a) then the clypeus is yellow-marked and the gonocoxae are not produced into pointed teeth. Andrena blanda can be separated from the Ovandrena by the genital capsule, as in Ovandrena the gonostyli are apically flattened and spatulate, but the disc is always longer than broad. The gonocoxae are also produced into apically projecting teeth, but the inner margins of the gonocoxae are parallel and do not diverge. The propodeal triangle of Ovandrena is also triangular and strongly defined by lateral carinae, whereas in A. blanda it is poorly defined and lacks lateral carinae.

Description

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.

Etymology

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.

Included species

Andrena blanda (Spain, including mainland Spain and newly recorded for Fuerteventura), Morocco, Algeria, Tunisia; Gusenleitner and Schwarz 2002).

Material examined (illustrative)

Algeria: Biskra [34.8600°N, 5.6995°E], 1♀, MNHN (lectotype des. Warncke 1967); 5 km N of Mecheria, st. 9, 7.iv.1983, 1♀, leg. R. Leys & P. v. d. Hurk, RMNH; 5 km SE of Sfissifa, st. 8, 6.iv.1983, 1♀, leg. R. Leys & P. v. d. Hurk, RMNH; Morocco: Drâa-Tafilalet, Tazenakht, 1 km W Anezal, c. 1600 m, 15.iv.2022, 1♀, leg. T.J. Wood, TJWC; Foum Zguid, 50 km N, 30.iii.1986, 1♂, 16♀, leg. M. Schwarz, MSC; Ifkern, 25 km E Boulemane, 25.v.1995, 10♀, leg. Mi. Halada, OÖLM; Spain: Fuerteventura, Costa Calma, ESE Montaña Pelada, 29.iii.2015, 3♀, leg. A. Müller, AMC/TJWC; Fuerteventura, S Costa Calma, Montañeta de los Verdes, 1.iv.2015, 5♀, leg. A. Müller, AMC; Sierra de María, 25 km W Lorca, 10.v.2003, 1♀, leg. J. Halada, OÖLM; Granada, Pantano de los Bermejales, 26.v.1982, 1♀, leg. R. Leys, RMNH.

Bryandrena subgen. nov.

Type species

Andrena florea Fabricius, 1793.

Diagnosis

Bryandrena is monotypic, and hence diagnosis of A. florea is de facto diagnosis of the subgenus. The combination of broad head (Fig. 41B, F; 1.3–1.4 times broader than long), the inner margins of the compound eyes diverging ventrally, humeral angle (stronger in male), punctate clypeus with underlying surface shiny over the majority of its area, propodeal triangle not defined by lateral carinae and comparatively smooth relative to the microreticulate dorsolateral parts of the propodeum (Fig. 41C), terga that are always at least partially red-marked (Fig. 41D, G), unmodified posterior face of the hind femur (without teeth, carinae, or spines), lack of squamous hairs, simple hind tibial spur (not broadened basally or medially), black male clypeus, and unique genital capsule (Fig. 41H; see description below) allows separation from any other Andrena species.

Figure 41. 

Andrena (Bryandrena) florea Fabricius, 1793, female A profile B head, frontal view C propodeum, dorsal view D dorsal view; male E profile F face, frontal view G terga, dorsal view H genital capsule.

Description

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.

Etymology

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.

Included species

Andrena florea (West Palaearctic, from Morocco and Iberia to Iran and the Ural Mountains; Gusenleitner and Schwarz 2002).

Limbandrena subgen. nov.

Type species

Andrena limbata Eversmann, 1852 (illustrated by Astafurova et al. 2022).

Remarks

Historically, A. toelgiana Friese, 1921 has been considered the sister species to A. limbata, differing by the yellow clypeus in the female sex (see Gusenleitner and Schwarz 2002). However, the male is unknown. Structurally, there are no differences. Examination of specimens from Bulgaria and Turkey show a gradient of yellow colouration on the clypeus, from entirely black, with a narrow longitudinal yellow strip, predominantly yellow-marked, and entirely yellow-marked (Fig. 42). Across this gradient there is also variation in the strength of the tergal hair bands, with material nominally conforming to A. toelgiana falling closer to A. limbata s. str. in displaying clear hair bands, whereas the subspecies A. limbata dusmeti Warncke, 1975 (Portugal, Spain, southern France, north-western Italy) has hair bands that are almost absent in fresh specimens (Figs 43C–H, 44A, B, E, F). On the basis of this colour gradient, absence of structural characters, and lack of a described male that could offer a distinctive difference, A. toelgiana syn. nov. is synonymised with A. limbata.

Figure 42. 

Andrena (Limbandrena) limbata Eversmann, 1852 female head in frontal view on west-east gradient A Spain B Bulgaria C Turkey (Gürün) D Turkey (Bingöl) E Turkey (Bingöl) F Turkey (Diyarbakır).

Figure 43. 

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.

Diagnosis

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. 43A), the long ocelloccipital distance equal to three times the diameter of a lateral ocellus (Fig. 43B), the posterior face of the hind femur which lacks a transverse carina and transverse row of raised teeth or spines, the weakly and shallowly punctate dorsolateral surfaces of the propodeum, the laterally clearly delineated and internally finely rugose propodeal triangle (not rugosely areolate), the pronotum with at most a weak lateral angle, the more or less squarish head which is only 1.1 times wider than long, and the simple hind tibial spur that is not broadened basally or medially.

Males can be recognised by most of the same characters: the yellow clypeus (Fig. 44C, D), the relatively rounded head which is only 1.1–1.2 times broader than long (Fig. 44C, D), the long ocelloccipital distance equal to three times the diameter of a lateral ocellus, the pronotum with at most a weak lateral angle, the weakly and shallowly punctate dorsolateral surfaces of the propodeum, the laterally clearly delineated and internally finely rugose propodeal triangle (not rugosely areolate), and the genital capsule with weakly produced rounded gonocoxal teeth, gonostyli with weakly raised and rounded projection on inner margin, and penis valves with rounded lateral hyaline extensions (Fig. 44G, H). No other Andrena species shows this combination of characters.

Figure 44. 

Andrena (Limbandrena) limbata Eversmann, 1852, Turkish male A profile C face, frontal view E terga, dorsal view G genital capsule; Andrena limbata dusmeti Warncke, 1967, Spanish male B profile D face, frontal view F terga, dorsal view H genital capsule.

Description

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.

Etymology

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.

Included species

Andrena limbata (Europe from Portugal and Spain to Turkey, Israel, northern Iran, and the Ural Mountains; Gusenleitner and Schwarz 2002).

Material examined (illustrative)

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, RMNH; Georgia: E Tbilisi, lori river S Sagarejo, 19.vi.2015, 3♀, leg. M. Snižek, OÖLM; Greece: Kos I. Kefalos, 4.v.1989, 2♀, leg. K.M. Guichard, NHMUK; Peloponnese, 12 km NWW Sparti, 26.v.2005, 1♀, leg. M. Kadlecová, OÖLM; Italy: Arezzo, Oliveto, 28.v.1989, 3♀, leg. H. & J.E. Wiering, RMNH; Piemonte, S. Benefetto Belbo, 10.vi.1979, 1♀, leg. Pagliano, MRSN; Portugal: Manteigas, Serra da Estrela [40.4009°N, -7.5396°E], 28.iv.–9.vii.1929, 1♀, leg. Kricheldorf, OÖLM (holotype of A. limbata dusmeti); Spain: Segovia, Madrona, 500 m NE, Arroyo del Hocino, 15.v.2021, 1♂, 2♀, leg. T.J. Wood, TJWC; Ávila, Hoyocasero, 1350 m, 20.v.1995, 1♂, 2♀, leg. H. & J.E. Wiering, RMNH; Turkey: Tanin-Tanin-Pass, 1700 m, 12.vi.1984, 1♀, OÖLM; Karatepe/Adana, 17.iv.1984, 3♂, 1♀, leg. K. Warncke, OÖLM; Akyaka, Yeşilova, 20.vi.2016, 1♀, leg. M. Kasparek, OÖLM.

Ovandrena subgen. nov.

Type species

Andrena oviventris Pérez, 1895.

Diagnosis

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. 45A, B). Ovandrena species have scutal hairs that are not as short and clearly squamous as in members of the Aenandrena, Limbandrena, or Lepidandrena Hedicke, 1933, but they are partially squamous in some species (Fig. 45C, D), whereas the scutal hairs in the relata-group and Blandandrena are typically simple, though in some species such as A. corax the hairs can be semi-squamous. The structure of the propodeal triangle is therefore the only character that allows consistent separation. In Ovandrena, the propodeal triangle is a broad and well-defined triangle (lateral margins extending almost to the lateral edges of the metanotum) that is defined by raised carinae; the internal surface is covered in raised irregular carinae of a similar width, therefore appearing homogenous and consistent (Fig. 45A, B). In the relata-group and Blandandrena, the propodeal triangle varies from smooth and undefined to weakly defined with at most irregular rugosity on its internal surface, but never approaches the condition in Ovandrena. The combination of these two characters therefore always allows recognition. They can also be partially recognised by the extremely dense and fine punctation of the terga, punctures separated by <0.5 puncture diameters; in comparison groups, the punctures are never this fine or this dense.

Figure 45. 

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. 45E, F). Members of the relata-group often have a yellow-marked clypeus, but for species showing a dark clypeus (e.g. A. corax), the clearly defined triangular propodeal triangle allows separation (smooth to at most weakly defined in the relata-group). Separation can be made from Blandandrena by the genital capsule and the propodeal triangle, as detailed in the diagnosis section for that subgenus.

Description

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. 45C, D). Pronotum laterally with weak to strong humeral angle. Dorsolateral surface of propodeum with regular and dense network of raised reticulation; propodeal triangle wide, lateral margins extending almost to lateral edges of metanotum, laterally defined by straight raised carinae, thus strongly triangular (Fig. 45A, B). Internal surface of propodeum covered by raised irregular carinae of a similar width, therefore homogenous and consistent. Forewing with nervulus interstitial to weakly antefurcal. Hind tibial spurs simple, not broadened basally or medially. Terga densely punctate, in females punctures extremely dense and fine, separated by <0.5 puncture diameters, in males punctures slightly sparser and coarser, separated by 0.5–1 puncture diameters. Male genital capsule compact, consistent across species, with gonocoxae strongly produced into apical rounded teeth (Fig. 45E, F). Gonostyli apically spatulate, with a more or less strongly raised inner margin. Penis valves basally narrow or moderately broad, in both cases becoming narrow medially.

Etymology

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.

Included species

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 Pisanty et al. 2022a).

Material examined

Andrena oviventris: Algeria: Teniet el Had [35.8727°N, 2.0007°E], 1♀, MNHN (lectotype; Fig. 46).

Figure 46. 

Andrena (Ovandrena) oviventris Pérez, 1895, female lectotype A label details B profile C face, frontal view D terga, dorsal view.

Andrena farinosa: Spain: Murcie [Murcia], 1♀, MNHN (lectotype).

Identification key to subgenus Ovandrena subgen. nov

1 Females 2
Males1 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)2 marsae
Tergal and sternal margins dark to hyaline, never extensively lightened reddish orange-brown (Morocco only)2 farinosoides

Pruinosandrena subgen. nov.

Type species

Andrena pruinosa Erichson, 1835.

Diagnosis

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.

Description

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.

Etymology

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.

Included species

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.

Identification key to subgenus Pruinosandrena subgen. nov

Note, the male of A. sparsipunctata is unknown. The males described by Wood et al. (2020b) are actually misidentified A. mediovittata Pérez, 1895; A. sparsipunctata is expected to be restricted to south-western Morocco. See Wood et al. (2020b) for additional useful images.

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. 37C). Terga with strong apical pale hair bands that uniformly cover the entirety of the tergal depressions and obscure the underlying surface (Figs 37D, 38D; North Africa to Israel) caroli Pérez
Fovea narrow, occupying at most half the distance between the lateral ocellus and the compound eye (Figs 33B, 36C). Terga with or without clear hair bands; if with hair bands, then hair bands typically do not uniformly cover the tergal depressions, either weaker basally or stronger apically 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. 59D). Fovea not strongly separated from the inner margin of the compound eye 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 33C, 59C). 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 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 33D, 36D). Found in Spain pruinosa Erichson
Terga always extensively or entirely red-marked (Figs 35D, 59D). Found outside of Spain 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. 85E). Scutum less densely punctate, punctures separated by 0.5 puncture diameters, with shiny interspaces, predominantly shiny (eastern and south-eastern Spain only) nilotica Warncke
Genital capsule with clear kink in the inner margins of the gonostyli (Fig. 85F). Scutum more densely punctate, punctures almost confluent, predominantly dull (central and southern Spain) pruinosa Erichson

Description of new species

Andrena (Avandrena) juliae Wood, sp. nov.

Type material

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.

Description

Female. Body length: 9 mm (Fig. 47A). Head: Dark, 1.4 times wider than long (Fig. 47B). Clypeus weakly domed, with fine granular shagreen, dull, overlain by weak raised rugosity, rugosity forming weak, obscure, and interrupted raised latitudinal striations; remaining surface impunctate. Process of labrum rounded rectangular, twice as broad as long, surface smooth and shiny. Gena equalling width of compound eye; ocelloccipital distance equals diameter of lateral ocellus. Foveae moderately broad, occupying half space between compound eye and lateral ocellus, short, ventrally extending to level of antennal insertions; foveae filled with black hairs (Fig. 47C). Face medially with light brown-white hairs covering clypeus and antennal insertions, intermixing with black hairs, becoming entirely black haired along inner margin of compound eyes and frons; gena and vertex with long light brown hairs, longest hairs equalling length of scape. Antennae dark, A7–12 ventrally slightly lightened grey by presence of scales; A3 equalling A4+5+6. Mesosoma: Scutum and scutellum with fine granular shagreen, very weakly shiny, predominantly dull, underlying surface shallowly and obscurely punctate, punctures separated by 1–2 puncture diameters (Fig. 47D). Pronotum rounded. Mesepisternum and dorsolateral parts of propodeum with fine granular shagreen, predominantly dull, shagreenation overlain by fine network of raised rugosity, not forming a linked network. Propodeal triangle short and broad, slightly depressed below level of dorsolateral parts of propodeum, with extremely fine granular shagreen, basal 2/3rds covered with raised longitudinal rugae, propodeal triangle thus strongly contrasting dorsolateral parts of propodeum (Fig. 47E). Mesepisternum with long strongly plumose hairs, hairs predominantly pale with some intermixed black hairs, hairs clearly exceeding length of scape. Scutum and scutellum with strongly plumose light brown hairs, intermixed with black hairs medially. Propodeum with long strongly plumose light brown hairs, propodeal corbicula incomplete, dorsal fringe weak and poorly defined, internal surface with numerous pale simple hairs. Legs dark, tarsi obscurely lightened reddish brown; pubescence light brown. Flocculus complete, strong, composed of strongly plumose light brown hairs; femoral and tibial scopa composed of light brown simple hairs. Hind tibial claws with strong inner tooth. Wings hyaline, stigma and venation dark orange, nervulus interstitial. Metasoma: Terga dark, apical rim of marginal areas narrowly lightened hyaline-yellow (Fig. 47F). Tergal discs with scattered and small hair-bearing punctures, punctures separated by 3–4 puncture diameters, underlying surface finely shagreened, silky smooth, shiny. Tergal discs with sparse upstanding pale hair, T2–4 with dense apical whitish hair bands that obscure underlying surface. Apical fringe of T5 and hairs flanking pygidial plate dark brown; pygidial plate apically rounded, surface flat and featureless.

Figure 47. 

Andrena (Avandrena) juliae sp. nov. female A profile B face, frontal view C head, dorsal view D scutum, dorsal view E propodeum, dorsal view F terga, dorsal view.

Male. Unknown.

Diagnosis

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. 73E). The propodeal triangle is clearly delinated by raised carinae and with the entire surface covered with longitudinal carinae. In A. melacana, the dorsolateral parts of the propodeum are entirely covered with fine granular shagreen, without an overlay of raised rugosity, with the propodeal triangle not depressed and with even finer granular shagreen, not strongly differentiated from the surrounding areas.

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).

Remarks

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. 47A). Across sampling locations in Cádiz province during March 2023, four species of Avandrena were active, specifically A. avara sensu stricto, A. juliae, A. melacana, and A. panurgina. All species appear to be narrowly oligolectic on Erodium (see below), and A. juliae could be found in direct sympatry with A. avara sensu stricto, A. melacana, and A. panurgina at the Las Algamitas site and with A. melacana and A. panurgina at the Tarifa site. Andrena erodiorum is currently known only from south-eastern Spain (Albacete). One A. juliae female showed abraded body hairs, and no males were captured, implying that the flight season may begin in early March or even late February, perhaps offering a reason as to why this species has been overlooked until now.

Etymology

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.

Distribution

Spain (Cádiz province).

Other material examined

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.

Andrena (Euandrena) isolata Wood, sp. nov.

Type material

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].

Description

Female. Body length: 9.5 mm (Fig. 48A). Head: Dark, 1.2 times wider than long (Fig. 48B). Clypeus weakly domed, densely and evenly punctate, punctures separated by <0.5 puncture diameters, interspaces shiny. Process of labrum trapezoidal, twice as broad as long, apical margin with very weak emargination medially. Gena slightly exceeding width of compound eye; ocelloccipital distance 1.5 times diameter of lateral ocellus; vertex behind ocellar triangle densely punctate, punctures separated by 0.5 puncture diameters. Fovea dorsally narrow, occupying ⅓ of space between lateral ocellus and compound eye, ventrally reaching level of antennal insertions, here narrowing to ½ dorsal width; fovea filled with black hairs. Face, gena, and scape covered with uniformly short black hairs, vertex dorsally with predominantly golden-brown hairs with scattered intermixed black hairs. Antennae dark, A3 slightly exceeding A4+5, shorter than A4+5+6. Mesosoma: Scutum and scutellum densely punctate, punctures predominantly separated by <0.5 puncture diameters, at most by 1 puncture diameter medially, underlying surface finely shagreened and weakly shiny (Fig. 48C). Pronotum rounded. Mesepisternum microreticulate, weakly shiny to dull, with obscure raised reticulation. Dorsolateral parts of propodeum microreticulate, weakly shiny, densely and shallowly punctate, punctures separated by 0.5–1 puncture diameter. Propodeal triangle narrow, internal surface finely granulate and impunctate, basally with obscurely and weakly raised rugosity, propodeal triangle thus strongly contrasting punctate dorsolateral parts of propodeum. Mesepisternum ventrally with moderately long black hairs, becoming intermixed with golden-brown hairs medially, with entirely golden-brown hairs dorsally, hairs at most equalling length of scape. Scutum and scutellum dorsally with short golden-brown hairs, scutum medially with very short and obscure black hairs. Propodeum with long golden-brown hairs, propodeal corbicula incomplete, dorsal fringe composed of long plumose golden-brown hairs, internal surface with numerous plumose light hairs, becoming darker ventrally. Legs predominantly dark, hind femorae ventrally and posterior face of hind tibiae obscurely lightened reddish-orange; pubescence black to orange-brown. Flocculus incomplete, short, composed of weakly plumose orange hairs; femoral and tibial scopa orange, tibial scopa with at most occasional brown hairs dorso-basally. Hind tibial claws with strong inner tooth. Wings weakly infuscate, stigma and venation dark brown, nervulus interstitial. Metasoma: Terga dark, apical rim of marginal areas very narrowly lightened hyaline-brown (Fig. 48D). Tergal discs regularly punctate, punctures separated by 1–2 puncture diameters, underlying surface shagreened, weakly shiny; punctures extending onto marginal areas, here weak and obscure, separated by 2–3 puncture diameters. T1–2 with orange-brown hairs basally, laterally, and apically; T2–3 with orange-brown hairs forming weak apical hair bands, not obscuring underlying surface, T4 with weak apical hair band black; discs of T3–4 with short black hairs. Apical fringe of T5 and hairs flanking pygidial plate black; pygidial plate rounded triangular, with slightly depressed and densely punctate area medially, lateral margins impunctate.

Figure 48. 

Andrena (Euandrena) isolata sp. nov. female A profile B face, frontal view C scutum, dorsal view D terga, dorsal view.

Male. Unknown.

Diagnosis

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. 48B). In A. bicolor s.l., the clypeal punctures are separated by an average of at least 1 puncture diameter, sometimes more, and the clypeus is therefore more strongly shiny due to the larger interspaces. The dense clypeal punctures place A. isolata close to A. fortipunctata, but the two species can be separated by the tergal structure (see illustrations in Wood et al. 2021), with the tergal margins weakly depressed with at most the apical rim lightened hyaline yellow (tergal margins strongly depressed and extensively lightened semi-translucent brown in A. fortipunctata), terga punctate with some punctures extending onto tergal margins, here separated by 2–3 puncture diameters (tergal margins impunctate in A. fortipunctata), margins of T2–3 with weak light brown hair fringes apically, T4 with entirely black hairs (T2–4 with white hair fringes emerging from junction between disc and margin, these hair fringes overlying marginal areas in A. fortipunctata), terminal fringe of T5 and hairs flanking pygidial plate black (terminal fringe and hairs flanking pygidial plate dark brown in A. fortipunctata). Andrena isolata is almost identical to the probably undescribed taxon from north-western Africa, but can also be separated by the structure of the clypeus. Andrena isolata has the clypeus densely punctate with punctures separated by <0.5 puncture diameters and with the interspaces shiny, whereas in the taxon from north-western Africa, the punctures are equally dense but the interspaces are shagreened and dull. Furthermore, this latter taxon has a short and shallow longitudinal furrow at the apex of the clypeus that is absent in A. isolata.

Remarks

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 Praz et al. 2019), but this requires dedicated study.

Etymology

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.

Distribution

Spain (Sierra Nevada).

Andrena (Micrandrena) ortizi Wood, sp. nov.

Type material

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, RMNH; Granada: Órgiva N, 1300 m, 26.vi.1988, 1♀, leg. M. Schwarz, OÖLM.

Description

Female. Body length: 7 mm (Fig. 49A). Head: Dark, 1.1 times wider than long (Fig. 49B). Clypeus long, apical margin clearly ventrally exceeding level of line between lower margins of compound eyes, surface weakly domed, irregularly punctate, punctures separated by 1–3 puncture diameters, impunctate longitudinal midline present; underlying surface basally finely shagreened and weakly shiny, shagreenation becoming weaker apically, here almost smooth and shiny. Process of labrum narrowly trapezoidal, slightly broader than long, apical margin narrowly truncate. Gena slightly exceeding width of compound eye; ocelloccipital distance equalling diameter of lateral ocellus. Fovea dorsally narrow, occupying ¼ space between lateral ocellus and compound eye, ventrally extending below level of antennal insertions, consistently wide along its length, filled with white hairs. Face, gena, vertex, and scape covered with sparse whitish hairs, none equalling length of scape. Antennae dark, A3 equalling length of A4+5. Mesosoma: Scutum and scutellum regularly punctate, punctures separated by 1 puncture diameter, underlying surface shagreened and weakly shiny (Fig. 49C). Pronotum rounded. Mesepisternum microreticulate, weakly shiny with finely raised reticulation. Dorsolateral parts of propodeum with dense network of reticulation; propodeal triangle laterally delineated with raised carinae, internal surface with dense network of raised rugosity. Mesepisternum, scutum, and scutellum with long whitish hairs, none equalling length of scape. Propodeal corbicula incomplete, dorsal fringe composed of long whitish plumose hairs, internal surface with scattered long simple whitish hairs. Legs dark, pubescence whitish to light brownish. Flocculus complete, composed of short weakly upturned plumose hairs; flocculus, femoral and tibial scopae white, tibial scopae with some brown hairs dorso-basally. Hind tibial claws with inner tooth. Wings hyaline, stigma dark brown, venation brown, nervulus interstitial. Metasoma: Terga dark, apical rim of marginal areas very narrowly lightened hyaline-brown. Tergal discs densely and clearly punctate, disc of T1 with punctures separated by 1 puncture diameter, T2–3 with punctures separated by 0.5 puncture diameters, underlying surface shagreened and weakly shiny (Fig. 49D). Marginal areas impunctate, with rectangular shagreen; marginal areas long, on T1 occupying ¼ of tergum, on T2 occupying ½ of tergum, on T3 occupying 2/3rds of tergum, on T4 occupying almost all visible tergum. T2–4 with long sparse plumose hairs arising from base of marginal area, covering but not obscuring marginal area; T3–4 apically with dense short apical fringe of white hairs laterally, obscuring underlying surface. Apical fringe of T5 and hairs flanking pygidial plate dark brown with occasional whitish hairs laterally; pygidial plate rounded triangular, with obscurely raised medial ridge, otherwise featureless.

Figure 49. 

Andrena (Micrandrena) ortizi sp. nov. female A profile B face, frontal view C scutum, dorsal view D terga, dorsal view.

Male. Unknown.

Diagnosis

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.

Remarks

At the Mirador Monte Ahí de Cara (Fig. 50A, see also Fig. 22A), this species was collected foraging for pollen on the spiny yellow species Vella spinosa (Brassicaceae) in open habitat just above the tree line. Andrena ortizi appears to be endemic to the Sierra Nevada. Searches at elevations lower than 2000 m did not detect the species during 2021 surveys, so the collection nominally made at 1300 m near Órgiva may not be representative – the slopes above Órgiva (southern slopes of the Sierra Nevada) ascend rapidly, the collector may well have climbed to a higher altitude when collecting this specimen. The morphological similarity between A. rugulosa, A. ortizi, and A. atlantea suggests a pattern of geographical isolation in montane habitats, though only A. ortizi and A. atlantea show a close genetic relationship based on the COI sequences.

Figure 50. 

Habitat context A Andrena (Micrandrena) ortizi sp. nov., Granada, Sierra Nevada, Mirador Monte Ahí de Cara, 2100 m, 12.vi.2021 B Andrena (Truncandrena) ghisbaini sp. nov., Málaga, PN Sierra de las Nieves, mountain peak S of Pinsapo Escalereta, 1600 m, 30.v.2021.

Etymology

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.

Distribution

Spain (Sierra Nevada).

Andrena (Truncandrena) ghisbaini Wood, sp. nov.

Type material

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, RMNH.

Description

Female. Body length: 15–16 mm (Fig. 51A). Head: Dark, 1.2 times wider than long (Fig. 51B). Clypeus weakly domed, clearly punctate, punctures separated by 0.5–1 puncture diameters with exception of median longitudinal impunctate line, narrow basally, broadening apically, thus elongate triangular; underlying surface finely shagreened, weakly shiny. Process of labrum broadly trapezoidal, twice as broad as long, apical margin clearly emarginate. Gena broad, almost two times width of compound eye; ocelloccipital distance 1.5 times diameter of lateral ocellus. Fovea dorsally occupying slightly more than ½ space between lateral ocellus and compound eye, extending ventrally to lower margins of antennal insertions, filled with black hairs. Gena ventrally and laterally with long light brown hairs, longest equalling length of scape, hairs becoming black on vertex, scape, and majority of face, with shorter light brown hairs around antennae insertions. Antennae basally dark, A4 apically, A5–12 ventrally extensively lightened orange; A3 exceeding A4+5, shorter than A4+5+6. Mesosoma: Scutum and scutellum with extremely shallow and obscure punctures, punctures separated by 0.5–1 puncture diameters, punctures disappearing into underlying fine granular shagreen, surface dull to very weakly shiny. Pronotum rounded. Mesepisternum and dorsolateral parts of propodeum with fine granular shagreen, weakly shiny, with fine and scattered raised hair-bearing punctures, punctures separated by 2–3 puncture diameters; propodeal triangle broad, with extremely fine granular shagreen, without hair-bearing punctures, propodeal triangle thus defined by change in surface sculpture from dorsolateral parts of propodeum. Mesepisternum and propodeum with long finely plumose light brown hairs, clearly exceeding length of scape; scutum and scutellum medially with long black hairs and occasional isolated pale hairs covering majority of disc, laterally becoming intermixed with light brown hairs. Propodeal corbicula incomplete, very weakly defined, dorsal fringe not differentiated from hairs of internal surface, both parts composed of long finely plumose long light brown hairs. Legs dark, apical tarsal segments lightened dark reddish, pubescence dark brown. Flocculus complete, composed of long weakly plumose and upturned light brown hairs; femoral scopae composed of light brown simple hairs; tibial scopa long, hairs exceeding apical width of hind tibia, hairs dorsally dark brown, ventrally golden orange (Fig. 51D). Hind tarsal claws with inner tooth. Wings hyaline, stigma dark brown, venation dark brown to orange, nervulus interstitial. Metasoma: Terga dark, apical rim of marginal areas narrowly lightened hyaline-yellow; discs with extremely fine granular shagreen, weakly shiny, with fine and scattered hair-bearing punctures, punctures separated by 3–4 puncture diameters (Fig. 51C). Disc of T1 with long light brown hairs, exceeding length of scape, decreasing in length over T2–3, disc of T3 with intermixed short light brown and black hairs, becoming predominantly black on discs of T4–5. T2–4 with weak apical fringes of short light brown hairs, not obscuring underlying surface. Apical fringe of T5 and hairs flanking pygidial plate dark brown, pygidial plate rounded triangular, with weakly raised medial area, otherwise featureless.

Figure 51. 

Andrena (Truncandrena) ghisbaini sp. nov. female A profile B face, frontal view C terga, dorsal view D tibial scopa, profile view.

Male. Body length: 13–14 mm (Fig. 52A). Head: Dark, 1.3 times wider than long (Fig. 52B). Clypeus weakly domed, entirely yellow-marked with exception of two dark rounded spots medio-laterally. Clypeus punctured, punctures separated by 0.5–1 puncture diameters with exception of median longitudinal impunctate line, essentially non-existent basally, broadening apically, thus elongate triangular; underlying surface finely shagreened, weakly shiny. Process of labrum broadly rectangular, 2.5 times wider than long, apical margin weakly emarginate, surface smooth and shiny. Gena broad, 2 times width of compound eye; ocelloccipital distance 1.5 times diameter of lateral ocellus. Gena ventrally and laterally with long light brown hairs clearly exceeding length of scape, becoming intermixed with black hairs on vertex. Face medially with extensive whitish to light-brownish hairs on clypeus, antennae insertions, and scape intermixed with black hairs along inner margins of compound eyes and scape. Antennae basally dark, A4–13 ventrally lightened orange; A3 longer than A4, shorter than A4+5; A4 rectangular, longer than broad, slightly shorter than A5. Mesosoma: Mesosoma structurally as in female; pubescence as in female. Legs basally dark, apical tarsal segments and hind tibiae lightened dark reddish-brown, pubescence dark brown to orange brown. Hind tarsal claws with inner tooth. Wings hyaline, stigma orange, venation dark brown to orange, nervulus interstitial. Metasoma: Terga structurally as in female. Discs of T1–4 with long light brown hair, on T1 clearly exceeding length of scape, becoming progressively shorter to T4; T5–6 with short black hairs on disc (Fig. 52C). T2–4 with weak apical hair fringes apically, not obscuring underlying surface. S8 columnar, apex rounded, ventral surface covered with short brown hairs. Genital capsule slightly elongate, gonocoxae produced into strong apical teeth, teeth pointed with apical margins diverging (Fig. 52D, E). Gonostyli basally narrow, apically produced and flattened into rounded triangular plates, internal margin strongly raised and reflexed. Penis valves basally broad, occupying more than ½ space between gonostyli, with narrow laterally produced hyaline extensions; penis valves strongly narrowing apically.

Figure 52. 

Andrena (Truncandrena) ghisbaini sp. nov. male A profile B face, frontal view C terga, dorsal view D genital capsule, dorsolateral view E genital capsule, dorsal view; Andrena (Truncandrena) villipes Pérez, 1895 male F genital capsule, dorsal view.

Diagnosis

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. 30) due to the antennae that are ventrally extensively lightened orange, the intermixed light and dark pubescence of the face, with pale hairs medially and dark hairs laterally, the intermixed light and dark pubescence of the scutum, with dark hairs medially and lighter hairs laterally, by the entirely yellow-marked male clypeus, and by the penis valves basally broad with lateral hyaline extensions, valves strongly narrowing apically.

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. 51D; A. villipes with scopa unicolourous orange, Fig. 30B), the longer face, clypeus clearly projecting ventrally well below a line drawn between the lower margins of the compound eyes (A. villipes with face shorter, clypeus only slightly projecting below this line in direct comparison), the clypeus clearly punctured with a longitudinal impunctate midline that broadens apically (A. villipes with clypeus obscurely and shallowly punctate, without obvious impunctate midline), and the reduced pale pubescence of the face, with light hairs restricted to the area around the antennal insertions (A. villipes with extensive pale hairs covering majority of face and clypeus, with black hairs predominantly along the inner margins of the compound eyes).

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. 52D, E; in A. villipes with the flattened apical part of the gonostyli more rounded, about as long as broad, Fig. 52F), the inner margins of these parts more strongly and acutely raised, slightly reflexed (in A. villipes with the inner margin less strongly raised and not reflexed).

Remarks

The two females from the Sierra de las Nieves (Fig. 50B) were collected from Cistus albidus Linnaeus (Cistaceae). Like A. villipes (Table 1), this species is likely to be oligolectic on Cistaceae. Additional surveys are needed to clarify the limits of its range. Male specimens from Elvira in the Naturalis collection were incorrectly determined by Teunissen as A. maroccana Benoist, 1950 which is a synonym of A. leptopyga Pérez, 1895.

Table 1.

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)

Etymology

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.

Distribution

Spain (Málaga province).

Description of missing sexes

Andrena (Micrandrena) alma Warncke, 1975

Description. Male. Body length 6.5–7 mm (Fig. 53A). Head: Dark, 1.2 times wider than long. Clypeus flattened, unevenly punctate with large punctures, punctures separated by 0.5–2 puncture diameters, underlying surface shagreened in basal half, polished and shiny in apical half. Process of labrum trapezoidal, slightly wider than long, apical margin slightly thickened. Gena 1.3 times width of compound eye (Fig. 53B, C); ocelloccipital distance 0.5 times diameter of lateral ocellus. Face medially with whitish hairs, scape with mixture of black and white hairs, inner margin of compound eyes with black hairs, gena ventrally with white hairs, becoming black dorsolaterally, vertex and frons with mixture of black and pale hairs, none exceeding length of scape. Antennae dark, A3 exceeding length of A4, shorter than A4+5. Mesosoma: Scutum and scutellum finely granularly shagreened and weakly shiny, shallowly and obscurely punctate, punctures separated by 2–3 puncture diameters. Pronotum with clear humeral angle. Mesepisternum and dorsolateral surfaces of propodeum with fine granular microreticulation, with regular slightly raised hair bearing punctures. Propodeal triangle with regular granular shagreen, basally and medially with obscure and finely raised rugosity, propodeal triangle thus defined by change in surface sculpture compared to dorsolateral parts of propodeum (Fig. 53D). Mesosoma laterally with long white pubescence, exceeding length of scape, pubescence becoming brownish and shorter dorsally, not exceeding length of scape. Legs dark, pubescence whitish. Hind tarsal claws with inner tooth. Wings hyaline, venation and stigma dark brown, nervulus interstitial. Metasoma: Terga dark, marginal areas obscurely lightened dark hyaline brown apically (Fig. 53E). Tergal discs with obscure and weak punctures that disappear into even underlying microreticulation, surface weakly shiny. Tergal discs with sparse long light brown hairs, T2–4 laterally with weak, widely interrupted apical hair fringes. S8 columnar, slightly broadened apically, ventrally covered with short yellowish hairs. Genital capsule compact, gonocoxae apically weakly produced into short rounded teeth; gonostyli parallel-sided, spatulate; penis valves slightly broadened basally (Fig. 53F).

Figure 53. 

Andrena (Micrandrena) alma Warncke, 1975 male A profile B head, dorsal view C head, profile view D propodeal triangle, dorsal view E terga, dorsal view F genital capsule, dorsal view.

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 53B, C), flattened and striation-free clypeus, simple genital capsule (Fig. 53F), and smooth and granulate propodeal triangle with obscure and finely raised rugosity basally (Fig. 53D). This smooth propodeal triangle places it close to former Distandrena species, but the flattened and striation-free clypeus excludes A. longibarbis Pérez, 1895 (clypeus domed, obscurely striate) and A. orana Warncke, 1974 (clypeus domed and striate). The evenly shagreened terga and obscure basal rugosity on the largely granularly shagreened propodeal triangle place it superficially close to A. djelfensis, but this species can easily be separated by the distinctive genital capsule with elongate and strongly medially bent gonostyli, whereas the genital capsule is simple and lacking distinctive features in A. alma. Finally, A. alma has a distinctly broadened gena that slightly but distinctly wider than the width of the compound eye, an unusual character in Micrandrena. This allows separation from A. abjecta which has the gena equalling the width of the compound eye. Collectively, these characters make recognition of A. alma straightforward, though as the males of A. tenostra and A. aff mica are unknown, diagnosis may become more challenging in the future. A level of caution should therefore be applied when identifying material from the extreme south and south-east of Spain.

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, EBDC.

Andrena (?Euandrena) ramosa Wood, 2022

Description. Male. Body length 8–10 mm (Fig. 54A). Head: Dark, 1.05 times wider than long (Fig. 54C). Clypeus long, weakly domed, unevenly punctate, punctures separated by 0.5–2 puncture diameters, underlying surface strongly shagreened to microreticulate in basal half, becoming smooth and shiny in apical half. Process of labrum trapezoidal, 3 times wider than long, ventral surface smooth and polished. Gena equalling width of compound eye; ocelloccipital distance 1.5 times diameter of lateral ocellus. Face medially and gena ventrally with long yellowish hairs, face laterally, frons, and scape with long black hairs, mixing medially on face with yellowish hairs, longest exceeding length of scape. Antennae dark, A3 exceeding length of A4, shorter than A4+5, A4 slightly longer than wide, A5–13 elongate, clearly longer than wide. Mesosoma: Scutum and scutellum obscurely punctate, punctures separated by 1–2 puncture diameters, disappearing into extremely strong underlying granular microreticulation, surface dull (Fig. 54D). Pronotum rounded. Mesepisternum and dorsolateral surface of propodeum with fine granular microreticulation, with finely raised network of reticulation that gives impression of large shallow punctures. Propodeal triangle narrow, surface with fine granular shagreen, basally and medially with finely raised rugosity, propodeal triangle thus defined by change in surface sculpture compared to dorsolateral parts of propodeum. Mesosoma with long, densely branches and plumose yellowish hairs clearly exceeding length of scape, black plumose hairs intermixed around wing bases, on scutum, and propodeum. Legs dark, pubescence brownish to black. Hind tarsal claws with inner tooth. Wings hyaline, stigma dark brown, venation dark to light brown, nervulus weakly antefurcal. Metasoma: Terga dark, apical rim of marginal areas very narrowly lightened hyaline brown (Fig. 54E). Tergal discs with obscure hair-bearing punctures, disappearing into background sculpture, becoming more strongly defined laterally, underlying surface shagreened and weakly shiny. T1–3 with long but loose plumose yellowish-brown hairs, these becoming black on T4–5. S8 relatively short, rectangular, apically truncate, ventrally covered with dark brown hairs. Genital capsule moderately elongate, gonocoxae apically weakly produced into short rounded projections, gonostyli more or less parallel-sided, spatulate (Fig. 54F). Penis valves occupying ½ space between gonostyli, slightly narrowing towards their apexes.

Figure 54. 

Andrena (?Euandrena) ramosa Wood, 2022 male A profile B head, profile view C head, frontal view D scutum, dorsal view E terga, dorsal view F genital capsule.

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. 54B), whereas in A. solenopalpa the mouthparts are extremely long, the labial palpi alone exceed the length of their head, the part of the mouthparts protruding in front of the head therefore collectively greatly exceed the length of the head. The clypeus of A. solenopalpa is also more extensively shiny, the gonostyli have their outer margin emarginate and are apically produced into narrow points, and A3 slightly exceeds the length of A4+5, whereas in A. ramosa the clypeus is shiny only in its apical half, the gonostyli are spatulate and apically truncate, and A3 only slightly exceeds A4, and is shorter than A4+5. The two species do not occur in sympatry, with A. ramosa restricted to south-western Spain, whereas A. solenopalpa is found in central and eastern Spain to southern France.

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 (Pisanty et al. 2022b), and subgeneric classification in this clade has been highly problematic due to the lack of clear delineating characters. Andrena ramosa does not belong to Margandrena Warncke, 1968 due to the lack of a strong humeral angle on the pronotum. It does not belong to the crocusella-group due to the lack of a humeral angle and the lack of lateral projections on the male penis valves (in addition to the lack of barcode similarity), and whilst it has strongly plumose pollen collecting hairs comprising the propodeal corbiculae and femoral scopae, those of the tibial scopae are simple, and the foveae do not narrow below which makes placement in Chrysandrena Hedicke, 1933 difficult. Andrena ramosa is therefore best considered to be affiliated with the subgenus Euandrena, pending investigation with more powerful genetic techniques. It clearly possesses a unique and unusual morphology within the West Palaearctic Andrena fauna.

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 Wood et al. 2022a) would not seem to be an adaptation to the collection of small Ericaceae pollen grains, and indeed this adaptation is absent in the Ericaceae specialist A. (Cnemidandrena) fuscipes (Kirby, 1802) which has simple pollen collecting hairs. Further study is required; the assumption remains that A. ramosa collects pollen from a plant species with large pollen grains, thus necessitating these branched and plumose hairs.

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.

Additional designation of lectotypes

Andrena (Aenandrena) hystrix Schmiedeknecht, 1883

Andrena (Aenandrena) hystrix Schmiedeknecht, 1883: 618, ♀ [France, lectotype by present designation: RMNH].

Remarks. Schmiedeknecht (1883) described several Andrena species from material that had been sent to him by Pérez. In each case, he indicated this in his title, e.g. ‘Andrena hystrix Perez in litt’ [sic]. For several of these species, the location of type material has been unclear, as they mostly did not seem to be in the MNHN in the Pérez collection (see Le Divelec 2021), or elsewhere (see Gusenleitner and Schwarz 2002). Searches in the RMNH unexpectedly uncovered specimens of four species described by Schmiedeknecht with labels written in Pérez’s distinctive handwriting. The exact providence of these specimens is unclear, but the RMNH collection is known to contain material from Schmiedeknecht’s collection (F. Bakker, pers. comm.). A lectotype was designated for one of these species (A. (Andrena) mitis Schmiedeknecht, 1883) in a previous publication (Wood 2023a); the others are designated here.

Schmiedeknecht (1883) described A. hystrix from female specimens from Hungary (Hungaria) and southern France (Gallia meridionali). He did not specify a type, though Gusenleitner and Schwarz (2002) list south France as the locus typicus. A specimen of A. hystrix from Marseille is labelled with Pérez’s handwriting, and this is considered to be part of the original syntypic series; it is here designated as a lectotype (Fig. 55A, B). No specimens from Hungary s.l. could be found.

Figure 55. 

Andrena (Aenandrena) hystrix Schmiedeknecht, 1883, female lectotype A label details B profile; Andrena (Notandrena) ranunculi Schmiedeknecht, 1883, female lectotype C label details D profile; Andrena (Euandrena) symphyti Schmiedeknecht, 1883, female lectotype E label details F profile.

Material examined. France: Marseille [43.3612°N, 5.3942°E], 1♀, RMNH (lectotype by present designation; Fig. 55A, B).

Andrena (Notandrena) ranunculi Schmiedeknecht, 1883

Andrena (Notandrena) ranunculi Schmiedeknecht, 1883: 617, ♀♂ [France, lectotype by present designation: RMNH].

Remarks. Schmiedeknecht (1883) described A. ranunculi, comparing it to A. ranunculorum. He gave the habitat as Russia (referring to A. ranunculorum) and southern France (referring to the specimens received from Pérez). A series of males and females labelled by Pérez as being from Bordeau [sic, = Bordeaux] were found in the RMNH. These conform to Schmiedeknecht’s description, and the concept used by subsequent authors. A female is here designated as a lectotype (Fig. 55C, D).

Material examined. France: Bordeaux [44.8352°N, -0.5888°E], 1♀, RMNH (lectotype by present designation; Fig. 55C, D); Bordeaux, 8♂, 5♀, RMNH.

Andrena (Euandrena) symphyti Schmiedeknecht, 1883

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 RMNH collection. Two females and one male labelled as being from Bordeau [sic, = Bordeaux]. Schmiedeknecht (1883) specifically states that the type material comes from Bordeaux, writing “In Gallia prope Bordeaux a Dom. Illustrissimo Perez detecta”. A female is here designated as a lectotype (Fig. 55E, F).

Material examined. France: Bordeaux [44.8352°N, -0.5888°E], 1♀, RMNH (lectotype by present designation; Fig. 55E, F); Bordeaux, 1♂, 1♀, RMNH.

Designation of neotypes

Andrena (Chlorandrena) boyerella Dours, 1872

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. 56).

Figure 56. 

Andrena (Chlorandrena) boyerella Dours, 1872, male neotype A profile B face, frontal view C terga, dorsal view D genital capsule dorsal view.

Remarks. As discussed above, Dours (1872) described A. boyerella from southern France and Algeria. No material is available for study, as all of Dours’ types were destroyed in a fire. Given that two taxa are present in these regions, in order to fix the name A. boyerella on the North African population, a barcoded neotype is designated from Moroccan material (Fig. 56).

Distribution. Morocco, Algeria, Tunisia, Italy (Sicily).

Andrena (Notandrena) griseobalteata Dours, 1872

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, RMNH (Fig. 57).

Figure 57. 

Andrena (Notandrena) griseobalteata Dours, 1872, female neotype A profile B face, frontal view C scutum, dorsal view D terga, dorsal view.

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. Warncke 1967), but the lectotype of A. erythrocnemis is actually A. chrysosceles (see Proshchalykin et al. 2017; Astafurova et al. 2021). Gusenleitner and Schwarz (2002) resolved this issue by resurrecting A. griseobalteata to species status. Finally, Wood and Monfared (2022) removed A. emesiana Pérez, 1911 (southern Turkey, Syria, Iran) from synonymy with this taxon.

Although Gusenleitner and Schwarz (2002) would seem to have resolved the issue, there is no type specimen for A. griseobalteata due to the loss of Dours’ collection. In the original description, Dours (1872: 428) listed the species from Saint-Sever in the department of Landes in south-western France, but also from Algeria. This is peculiar, because A. griseobalteata is not known from North Africa. The original description also does not allow for completely unambiguous recognition of the species, though the dense punctation of the scutum is mentioned. In the interests of nomenclatural stability, a neotype from Bérenx in south-western France (36 km from Saint-Sever) is designated in order to fix the species concept for the future (Fig. 57).

Finally, though listed from Spain by Ortiz-Sánchez (2011, as A. griseobalteata; 2020, as A. erythrocnemis), the presence of this taxon in Spain is somewhat doubtful. The distribution maps of Warncke (Gusenleitner and Schwarz 2002) show dots around south-western France into the Pyrenees, but there are no unambiguous dots for Spain. However, I have been able to examine one specimen of A. griseobalteata from Spain, from the Sistema Central. The species is also expected to occur in the western Pyrenees; more recent material should be found to establish whether this taxon remains present in Iberia.

Material examined. Spain: Sierra de Gredos, 12 km SSW Hoyos del Espino, 1950–2100 m, 4.vii.1972, 1♀, leg. J.A.W. Lucas, RMNH.

Distribution. Spain, France, Italy, Croatia, Hungary, Albania, Romania, North Macedonia, Bulgaria, Greece, Turkey (western and northern Turkey; Gusenleitner and Schwarz 2002). The species is not considered to be present in North Africa, or in the Levant, as it is not found in very dry environments.

Andrena (Taeniandrena) poupillieri Dours, 1872

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. 58).

Figure 58. 

Andrena (Taeniandrena) poupillieri Dours, 1872, female neotype A profile B face, frontal view C dorsal view D terga, dorsal view.

Remarks. This is the taxon referred to as ‘A. poupillieri 2’ by Praz et al. (2022). The taxon A. poupillieri incana Warncke, 1975 which was described from and restricted to the Balearic Islands is considered a simple synonym of A. poupillieri due to the lack of genetic differentiation observed (see above), even though the tergal punctation is slightly reduced compared to the nominate taxon. The specimen used in the analysis of Praz et al. (2022) is designated as a lectotype (Fig. 58).

Distribution. Morocco, Algeria, Tunisia, Spain (mainland and Balearic Islands). Records (Gusenleitner and Schwarz 2002) from Libya must be confirmed, though they probably do refer to true A. poupillieri. Records from Crete probably refer to unrecognised A. ovata specimens, and so A. poupillieri is not considered to be present there until definitive material is located.

Andrena (Pruinosandrena) succinea Dours, 1872

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. 59).

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. 34B is selected as a neotype (Fig. 59) in order to fix the species concept for the future.

Figure 59. 

Andrena (Pruinosandrena) succinea Dours, 1872, female neotype A profile B face, frontal view C mesosoma, dorsolateral view D terga, dorsal view.

Distribution. Morocco, Algeria, Tunisia, Libya, Egypt, Israel and the West Bank, Jordan, Syria, Saudi Arabia, Iran (Wood and Monfared 2022).

Andrena (incertae sedis) numida Lepeletier, 1841

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. 60).

Figure 60. 

Andrena (incertae sedis) numida Lepeletier, 1841, female neotype A profile B face, frontal view C dorsal view D T2–5, dorsal view detail.

Remarks. As discussed above, it is beneficial to designate a neotype for A. numida since the original type series cannot be located in the MNHN, and so that the name and genetic identity of North African populations can be fixed.

Distribution. Morocco, Algeria, Tunisia, Libya, Italy (Sicily, Calabria, Campania).

Updated faunal list

Ortiz-Sánchez (2020) lists 212 species of Andrena from Spain. However, given the taxonomic and faunal work that has been conducted since then, this total is slightly too low and contains mistakes resulting from taxonomic confusion and the use of variable species concepts in the literature.

Species removed from baseline list

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 Ortiz-Sánchez (2020). Four further species must also be removed, A. similis Smith, 1849, A. enslinella Stöckhert, 1924, A. simillima Smith, 1851, and A. chrysopyga Schenck, 1853. Andrena (Taeniandrena) similis was shown to be a synonym of A. (Taeniandrena) russula Lepeletier, 1841 by Praz et al. (2022). Ortiz-Sánchez (2020) listed both species as present.

Dardón (2010) and Dardón et al. (2014) listed A. (Micrandrena) aff enslinella as present in Spain based on the listing of Warncke (1976) and the specimen in his collection. I do not consider this to be ecologically possible, as A. enslinella is a species of Central and Eastern Europe to the Caucasus and Iran, having its western limit in Germany and Austria. The species appears to be absent from France (Le Divelec 2021). Given the highly interesting Andrena fauna in the Sistema Ibérico that contains relictual North African taxa such as A. (Cordandrena) vaulogeri Pérez, 1895 and the endemic A. (Parandrenella) taxana Warncke, 1975, further study is required, as this specimen may represent an undescribed species, as suggested by Dardón (2010).

Andrena (Cnemidandrena) simillima was listed by Ortiz-Sánchez (2020), but not by Warncke (1976). The distribution maps of Warncke (Gusenleitner and Schwarz 2002) give a single isolated dot for this species in north-eastern Spain, presumably in the eastern Pyrenees. The next closest record comes from western France (c. 400 km), with no other records for the Pyrenees. Warncke et al. (1974: carte 140) gave Bordeaux as the south-western limit for A. simillima (specimen leg. Pérez, coll. Warncke OÖLM). I consider the dot on the maps of Gusenleitner and Schwarz (2002) to be of doubtful providence; I have examined no material from Iberia of this species, and I could not locate a Spanish specimen in the Warncke collection. No other authors have ever recorded A. simillima from the Pyrenees, though A. (Cnemidandrena) nigriceps (Kirby, 1802) is common in this region. Males of A. simillima and A. nigriceps are extremely difficult to separate, and it is possible that this dot represents a misidentification of A. nigriceps. Moreover, this record is ecologically questionable, since A. simillima is predominantly a northern species (sensu lato, since cryptic taxa may be present) found in temperate habitats. In the absence of available specimens, the isolated nature of this record, and the identification difficulties associated with this group, I remove A. simillima from the Iberian fauna.

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 (Gusenleitner and Schwarz 2002), there are dots for A. chrysopyga from south-western France, but none in Iberia proper. Andrena chrysopyga is a species of dry grassland and steppe, extending east across the Great Eurasian Steppe to Kazakhstan. Given the absence or scarcity of steppe habitats in France, its presence in much of the country seems ecologically unlikely. Given the overly generous distribution of A. chrysopyga given by Warncke (e.g. the species is known only from the extreme east of Belgium yet Warncke’s map indicates the presence of this taxon throughout Belgium, see Wood 2023a), the lack of available specimens, and the identification mistakes known to occur between A. chrysopyga and A. gravida, A. chrysopyga is not considered to be part of the Iberian fauna until such a time as validated specimens can be found or the species can be newly captured. Its historical and contemporary distribution in France must also be clarified, as I believe that most records are misidentifications of A. gravida.

Species added to baseline list

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 (Wood et al. 2021); A. (Plastandrena) nigrospina Thomson, 1872 (Ortiz-Sánchez et al. 2022); A. (Taeniandrena) afzeliella (Kirby, 1802) and A. (Taeniandrena) ovata Schenck, 1853 (Praz et al. 2022); A. (Avandrena) erodiorum Wood & Ortiz-Sánchez, 2022, A. (Avandrena) melacana Warncke, 1967, A. (Taeniandrena) lusitania Wood & Ortiz-Sánchez, 2022, and A. (Suandrena) gades Wood & Ortiz-Sánchez, 2022 (Wood & Ortiz-Sánchez, 2022); A. (incertae sedis) ramosa Wood, 2022 (Wood et al. 2022a); A. (Taeniandrena) contracta Wood, 2022 (Wood 2022); and A. (Andrena) clarkella (Kirby, 1802) (Álvarez Fidalgo and Aguado Martín 2022).

Therefore, relative to the baseline of Ortiz-Sánchez (2020), 12 species are removed, and 28 species are added (including Andrena aff mica). In sum, the taxonomic changes and faunal updates presented here and in the referenced papers brings the Iberian Andrena fauna to 228 species, a substantial increase on the 175 species recorded by Warncke (1975a), with 228 species known from mainland Spain and 128 species from mainland Portugal. A full checklist can be found in Suppl. material 1.

Dietary niche of Iberian Andrena species

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. Westrich 1989) when these same species do not differ in their pollen foraging behaviour in Iberia. Pollen was analysed and removed from a total of 1,127 specimens from 76 species from 12 countries. Findings are summarised in Table 1. Andrena species are grouped by subgenus to highlight their often conserved patterns of pollen collection.

Demonstration of oligolecty in understudied species

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. 61A). However, two species deviate from this pattern, A. bellidis Pérez, 1895 and A. leucophaea Lepeletier, 1841. Andrena bellidis appears to be polylectic, whereas A. leucophaea was associated with Bellis (Asteraceae; Asteroideae) and may be oligolectic on this subfamily. Both species fly early in the year (typically February-April) and are uncommon in collections; more study is required, but neither species is likely to be specialised on Brassicaceae.

Figure 61. 

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. 61B). Based on direct observations, all five Iberian members of the relata-group (including A. macroptera and A. murana) appear to be narrowly oligolectic on Reseda, but additional pollen analysis is necessary for confirmation; this being limited by the scarcity of these species in collections.

All studied members of the subgenus Chlorandrena are specialists of Asteraceae. However, the only published associations relate to the subfamily Cichorioideae (e.g. Westrich 1989; Schwenninger 2015). However, three Iberian Chlorandrena collect exclusively from Asteraceae subfamily Asteroideae: A. abrupta, A. elata (Fig. 61C), and A. leucolippa. These three species form a clade within the Chlorandrena along with A. boyerella which is also specialised on Asteraceae subfamily Asteroideae (data to be published in the North African revision). This situation resembles the situation within the genus Panurgus, members of which are specialised on Asteraceae but for which members of different lineages within Panurgus utilise either Asteroideae or Cichorioideae (Wood et al. 2022b).

Within the newly erected subgenus Ovandrena, no clear pattern was seen, as A. farinosa is a specialist of small-flowered Fabaceae (Fig. 61D) and A. oviventris is a specialist of Reseda (Fig. 61E). At the moment, A. farinosa must be considered a broad oligolege, as the identity of pollen grains removed from museum specimens cannot be identified to the genus level. In Iberia, all my observations of this species come from Lotus dorycnium Linnaeus (=Dorycnium pentaphyllum Scopoli), and so the species may be more narrowly specialised on the genus Lotus, but this requires additional study.

Within the subgenus Simandrena, central and northern European species are well-known to be polylectic (Westrich 1989). In contrast, three West Mediterranean species appear to be oligolectic, A. vetula (Fig. 61F) and A. antigana Pérez, 1895 on Brassicaceae and potentially A. rhypara Pérez, 1895 on Reseda. All three species belong to the group of Simandrena with a strongly shagreened and almost impunctate scutum that is distributed predominantly across the Mediterranean basin, with the highest diversity in North Africa and the Levant. Further study is required to establish the dietary niche of A. cilissaeformis which may be polylectic.

Most studied members of the subgenus Truncandrena are specialised on Brassicaceae (Westrich 1989), though this subgenus is species-poor in Central Europe and most taxa are Mediterranean and understudied. In Iberia, A. doursana, A. ferrugineicrus, and A. nigropilosa are also specialised on Brassicaceae. However, A. villipes was strongly associated with Cistus (Cistaceae) based on pollen analysis and direct observations, a behaviour that has not previously been demonstrated for this subgenus. As discussed above, this is likely to be the dietary niche for A. ghisbaini, the two species together appear basally in the COI phylogeny (Fig. 29). Additional study is required to establish whether this basal position is supported by more powerful genetic analyses.

The subgenus Micrandrena contains mixture of oligolectic and polylectic species in central and northern Europe, though polylectic species predominate (Westrich 1989). In Iberia, numerous species display oligolectic behaviour, specifically A. ampla (Apiaceae), A. djelfensis and A. fabrella Pérez, 1903 (Cistaceae), A. longibarbis, A. nitidula, A. orana, and A. tenuistriata (Brassicaceae), and A. omnilaevis (Crassulaceae). The dietary status of A. spreta is somewhat unclear, as specimens from both the spring and summer generations analysed here collected predominantly Brassicaceae pollen (93.1%). There is a bias towards spring generation specimens which are typically more abundant, and which fly at a time of year with high Brassicaceae availability. Additional samples from the summer generation are required, but the species can provisionally be considered to be polylectic with a strong preference for Brassicaceae.

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 Müller and Kuhlmann (2008), this species would not necessarily be classified as oligolectic on Geraniaceae. However, Geraniaceae pollen grains are very large, usually in excess of 50 μm, and it is possible for contaminant pollen to be present between these large grains in a way that is much less common when dealing with smaller grains which pack into scopal hairs more tightly. Behavioural observations of A. panurgina, including those of males which patrol around Erodium (Geraniaceae) plants in March and early April, strongly suggests oligolecty on Geraniaceae. All behavioural pollen-foraging observations of A. panurgina, along with other western Avandrena Warncke, 1968 species A. avara, A. erodiorum, and A. melacana Warncke, 1967 have been made at Erodium (Wood and Ortiz-Sánchez 2022; pers. obs., Álvarez Fidalgo in litt.). Consequently, the presence of Asteraceae and Brassicaceae pollen in the quantitative analysis is considered likely to be contamination, and A. panurgina is considered to be an oligolege of Geraniaceae along with the other western Avandrena species (see Pisanty et al. 2022a).

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 (Wood 2023b) which requires genetic confirmation of its placement. The two West Mediterranean species are instantly recognisable in the female sex because of their extraordinarily long and incredibly plumose hairs of the tibial and femoral scopae and both flocculi (see Fig. 64H). Baldock et al. (2018) identified the use of Plantago (Plantaginaceae) pollen by A. orbitalis, but did not come to a firm conclusion as to the overall dietary niche of this species. Analysis of scopal pollen loads and behavioural observations (A. orbitalis in the Algarve, Portugal and A. rufiventris in the Middle Atlas, Morocco) show that both A. orbitalis and A. rufiventris are specialists of Plantago. In the spring, usually in the months of March and April, both species can be seen visiting low-growing Plantago species. Upon arriving (Fig. 62C) at a small flower head, they grab onto its side and present the underside of their body so that falling grains will land on their venter and scopae (Fig. 62A). They will then manipulate the anthers using their fore legs and mandibles (Fig. 62D), often directly scraping pollen from the anthers using their mandibles (Fig. 62E). When the flower head is long, they will sometimes work from bottom to top, often vigorously dislodging pollen grains that can form a small but distinctly visible cloud (Fig. 62B). Pollen landing on the body is then groomed into the scopae.

Figure 62. 

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. 62F). Pollen is packed into the scopae dry, and inspection of freshly caught and pinned specimens shows that pollen rapidly falls out of the scopae with gentle manipulation, for example with an entomological pin. This probably explains both why the scopal hairs are so densely and finely plumose in order to retain these small pollen grains (typically <20 μm in diameter), and also why very few specimens in museum collections have scopae that retain pollen, as dry grains are easily dislodged after specimen collection. Use of pollen from wind-pollinated plants is well-known in bees, predominantly by social polylectic species (Saunders 2018), but also by solitary species, including species belonging to genera such as Lipotriches Gerstaecker, 1858 (Halictidae) that can be oligolectic upon members of the Poaceae (Immelman and Eardley 2000). Whilst some Andrena species are known to collect pollen from wind-pollinated plants (Wood and Roberts 2017; Saunders 2018), to my knowledge this is the first time that any Andrena species has been demonstrated to be specialised for pollen collection on a wind-pollinated plant.

Finally, a note on the pollen collection preferences of A. afzeliella and A. ovatula is beneficial. Praz et al. (2022) demonstrated that A. afzeliella is polylectic with a strong preference for Fabaceae, and A. ovatula is oligolectic on Fabaceae. Additional detail and context can be provided based on observations in Iberia which help to explain their differing distributions and ecologies. Andrena afzeliella typically visits herbaceous Fabaceae, such as Trifolium