identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
3FDD9A403F8E5D2E83F67CA614B784DD.text	3FDD9A403F8E5D2E83F67CA614B784DD.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Aciandrena) chelma Warncke 1975	<div><p>Andrena (Aciandrena) chelma Warncke, 1975</p><p>Andrena (Aciandrena) chelma Warncke, 1975: 66, ♀ [Greece, OÖLM, examined]</p><p>Material examined.</p><p>GREECE • 2 ♀; Chelmos, Peloponnese; 4 Jun. 1962; M. Schwarz leg.; OÖLM (holotype and paratype) • 1 ♀; Chelmos, Peloponnese; 4 Jun. 1962; M. Schwarz leg.; OÖLM (paratype of Andrena pulicaria Warncke, 1975) • 1 ♀; Chelmos, Peloponnes; 1900 m a. s. l.; 4 Jun. 1962; M. Schwarz leg.; OÖLM (not part of type series; originally determined as Andrena tenuis Morawitz, 1877 by Warncke) • 1 ♂, 1 ♀; Chelmos [ Aroania]; 1700 m a. s. l.; 27–29 May 1987; H. Teunissen leg.; RMNH • 1 ♀; Melano Mts ( Arkadia); 1600 m a. s. l.; 27 May 1986; H. Teunissen leg.; RMNH • 2 ♀; Pelop. Tripolis 30 km Sparta; 870 m a. s. l.; 23 Apr. 2014; K. Deneš leg.; OÖLM • 4 ♂, 1 ♀; Peloponnese, Mount Maenalon, 4.5 km NW Kardaras; 1600 m a. s. l.; 4 Apr. 2024; T. J. Wood leg.; TJWC • 19 ♂, 2 ♀; Central Greece, Mt. Parnassos, 4.4 km NNW of Kalyvia Livadiou Arachovas; 1250 m a. s. l.; 9 Apr. 2024; T. J. Wood leg.; TJWC .</p><p>Notes.</p><p>Warncke (1975) introduced a complicated situation when describing A. chelma . On page 66 he described A. chelma in the female sex only (male newly diagnosed below) with the holotype coming from Chelmos in the Peloponnese (Mount Aroania). Then, on page 68 he described A. pulicaria in both the female and male sexes, with a female holotype coming from Ürgüp in central Anatolia in Turkey. Additional paratypes came from Yeşilhisar and Konya (Warncke 1974 a: 100). However, in Warncke (1975: 69) he writes “ außerdem Griechenland: Chelmos ”. Indeed, in the Warncke collection there is a female paratype of A. pulicaria from the exact same collecting event (4 Jun. 1962) as the type series of A. chelma . The two species are very similar in the female sex due to the elongate head with the process of the labrum extremely small, narrowly triangular and pointed (see also Andrena afghana Warncke, 1974).</p><p>Furthermore, a specimen in Warncke’s boxes of “ in progress ” material (i. e. outside of the main catalogued Warncke collection) contained another specimen from the same collecting event which was determined by Warncke as A. tenuis, a species that does not occur in Greece. As Warncke never dated his determination labels (e. g. see Ebmer 1999: 183), it is not possible to reconstruct a timeline as to how his species concepts developed. Examination of the specimens from Chelmos (determined by Warncke as A. chelma, “ A. pulicaria ”, and “ A. tenuis ”), as well as additional specimens collected by Teunissen and contemporary collections suggests that there is just a single taxon in Greece that can be referred to as A. chelma . Morphological differentiation in female Aciandrena species is extremely challenging, but the newly recognised males can be diagnosed with reference to their genital capsule. New sequences of A. chelma have been generated from Chelmos and the Melano mountains (Fig. 1); additional sequencing from Turkey is required to understand genetic differentiation from A. pulicaria .</p><p>Diagnosis.</p><p>The male of A. chelma can be recognised as an Aciandrena due to its small body size (Fig. 2 A), overall dark integument with the exception of a partially yellow-marked clypeus (Fig. 2 B), finely granulose propodeal triangle (Fig. 2 C; lacking lateral or internal rugae), and “ typical ” genital capsule which has gonocoxae teeth and elongate gonostyli (Fig. 2 D; not as in Graecandrena Warncke, 1968 with the gonostyli short and stubby and without gonocoxal teeth). The genital capsule is elongate, which allows separation from Andrena aciculata Morawitz, 1886 and Andrena lamiana Warncke, 1965 (see photographs in the identification key below), placing A. chelma close to A. pulicaria . The two species can be separated due to the genital capsule. In A. chelma, the inner margins of the gonostyli are raised but smooth, without a kink (in A. pulicaria with the inner margins of the gonostyli raised and showing a distinct kink medially), and the penis valves at their base have the outer margins slightly raised, penis valves broadest slightly before the gonocoxal teeth (in A. pulicaria with the penis valves laterally lacking raised margins, penis valves continuing to broaden as they meet the gonocoxal teeth). Overall, these characters are slight, but are typical for species-level differences in Aciandrena . Given the small nature of the differences, a formal description of the male of A. chelma is not considered to be necessary, especially as it is integrated into the identification key below. In this context, A. chelma is best considered to be an isolated species restricted to southern Greece and geographically separated from populations of A. pulicaria in Anatolian Turkey.</p><p>Distribution.</p><p>Greece (Peloponnese and south-central Greece on Mount Parnassus).</p></div>	https://treatment.plazi.org/id/3FDD9A403F8E5D2E83F67CA614B784DD	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
E4F4E997187A52439D9FA0305580F5D8.text	E4F4E997187A52439D9FA0305580F5D8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Aciandrena) Warncke 1968	<div><p>Subgenus Aciandrena Warncke, 1968</p><p>Genetics.</p><p>The Subgenus Aciandrena is taxonomically complex, and difficult to characterise both morphologically and genetically using barcodes. Some taxa placed in the subgenus such as Andrena janthina Warncke, 1975 (see Gusenleitner and Schwarz 2002) actually represent undescribed subgenera (Pisanty et al. 2022 a, 2022 b), and are not considered here, as all constituent species are Levantine (S. Bossert, G. Pisanty, T. J. Wood, unpublished data). Even in a narrow sense, Aciandrena s. str. seems particularly prone to the generation of NUMTs (Nuclear Mitochondrial DNA sequences) based on results from nanopore sequencing which can produce multiple different sequences from the same individual (unpublished data). As such, individual unusual sequences must be treated with great care to avoid the overdescription of taxa, particularly since females are often impossible to separate with confidence morphologically.</p><p>A particularly troublesome species pair are Andrena aciculata Morawitz, 1886 and Andrena lamiana Warncke, 1965 . Males are separable after dissection and examination of the apex of sternum 8 (truncate in A. aciculata, narrowly pointed in A. lamiana, see male identification key), but females are much more challenging if not impossible to separate morphologically. Genetically, sequences of A. aciculata were available from Czechia, North Macedonia, and Greece (Samos, Rhodes), and sequences of A. lamiana were available from Greece (East Macedonia; Fig. 1). Andrena aciculata showed moderate intraspecific variation of 3.37 % (range 0.45–6.28 %) but formed a ladder of sequences towards A. lamiana . The two A. lamiana sequences were identical, and were separated from A. aciculata by an average genetic distance of 3.76 % (range 0.86–5.05 %). Resolution of these two taxa remains obscure, and it is very possible that many specimens are misidentified in the female sex. Identification continues to rely on dissection of males. Additional focused study of this subgenus using additional molecular tools including nuclear DNA is necessary to resolve species boundaries across the East Mediterranean and Middle East.</p></div>	https://treatment.plazi.org/id/E4F4E997187A52439D9FA0305580F5D8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
2536BCE6250E513EAE02ACC0AF7E9966.text	2536BCE6250E513EAE02ACC0AF7E9966.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Campylogaster) erberi Morawitz 1871	<div><p>Andrena (Campylogaster) erberi Morawitz, 1871</p><p>Andrena erberi Morawitz, 1871: 222, ♀ ♂ [Greece, ZISP, examined by photograph]</p><p>Andrena kozarowi Nedialkov, 1914: 189, ♀ [Bulgaria, type lost] syn. nov.</p><p>Material examined.</p><p>ALBANIA • 1 ♂; Xhafaj; 15 Jul. 2017; K. Janssen leg.; KJCB ; BULGARIA • 1 ♂; Blagoevgrad, Dolno Spanchevo, 500 m S, sandy / calcareous grassland; 150 m a. s. l.; 16 May 2024; T. J. Wood leg.; TJWC • 2 ♀; Kresna [Кресна]; 9 Jul. 1990; B. &amp; O. Tkalců leg.; OÖLM • 2 ♂; Kresna [Кресна]; 12 Jul. 1990; B. &amp; O. Tkalců leg.; OÖLM • 1 ♀; S Sinitovo vill; 310 m a. s. l.; 8 May 2023; T. Trifonov leg.; IBER • 1 ♀; Struma, Valkovo, Sandanski; 3 Jul. 2019; M. Halada leg.; OÖLM • 2 ♀; údolí řeky [river valley] Lebnice; 1 Jul. 2000; J. Straka leg.; JSPC</p><p>Notes.</p><p>Nedialkov (1914) described Andrena kozarowi based on an unknown number of female specimens from Stara Zagora collected in July (Mense Iulio). The name was not included in the catalogue of Warncke (1967 b), but it was indicated as a possible synonym of A. erberi by Gusenleitner and Schwarz (2002: 252). Given this regional revision, it is appropriate to deal with it more definitively here.</p><p>The species name is written using the Latin alphabet, and the original description is written in Latin and then duplicated and translated into Bulgarian; the name is consequently available, as it fulfils ICZN (1999) requirements. It is diagnosed as “ Andrena erberi Mor similis ”, indicating close proximity to Morawitz’s species. The description presents a bee that conforms to the key aspects of A. erberi, including the scutum, scutellum, and metapostnotum densely covered with ferruginous squamous hairs (thorax superne (etiam scutellum et postscutellum) densissime ferrugineo squamosus, lateribus albido villosus), metasoma elongate, convex, and dull, subtly punctate and with T 1–2 laterally weakly reddish (Abdomen elongatum, convexiusculum, opacum, subtilissime punctatum; segmenta 1 et 2 lateribus subtus rufescentia), the tergal margins with ochraceous squamous hair bands (segmenta 1–4 apice densissime ochraceo squamosa (late fasciata)), and body length of 13 mm.</p><p>The overall bee is described as black (nigra), and A. erberi can present dark specimens such as the type (Astafurova et al. 2021) and contemporary specimens caught in Bulgaria and northern Greece (Fig. 3 A, B), as well as specimens extensively lightened red (Fig. 3 C, D) as in Andrena amasia Radoszkowski, 1876 (a junior synonym described from Amasya in northern Turkey; the specimen illustrated is the possible holotype in the ZMHB collection). Concerning A. kozarowi, searches in the NZMS unfortunately could not locate Nedialkov’s material (T. Ljubomirov, pers. comm.), but it is not considered necessary to create a neotype, as the identity of the taxon appears clear based on the description. It is formally synonymised under A. erberi syn. nov.; the species continues to be present in southern Bulgaria.</p><p>Distribution.</p><p>Albania *, North Macedonia, Greece, Bulgaria, Romania, Ukraine, Crimea, Turkey, Syria,? Egypt [unclear; based on the known distribution, A. erberi is a species of warm and humid East Mediterranean habitats, but not of dry desert habitats], Georgia, Armenia, Azerbaijan, Iran (Fig. 4; Gusenleitner and Schwarz 2002; Osytshnjuk et al. 2005; Astafurova et al. 2021).</p></div>	https://treatment.plazi.org/id/2536BCE6250E513EAE02ACC0AF7E9966	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
5A748AF7ECB652EB9F044750A9885628.text	5A748AF7ECB652EB9F044750A9885628.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Cryptandrena) Pittioni 1948	<div><p>Subgenus Cryptandrena Pittioni, 1948</p><p>Genetics.</p><p>Sequences of Andrena ventricosa Dours, 1873 were available from Portugal, Spain, Italy, Romania, and Lebanon, and sequences of Andrena brumanensis Friese, 1899 were available from Greece and Israel (Fig. 1). A single sequence from Crete that was tentatively identified as A. brumanensis was separated from A. ventricosa by an average genetic distance of 10.29 % (range 9.76–10.56 %) and from A. brumanensis by an average genetic distance of 7.07 % (range 6.89–7.20 %). Within A. ventricosa, two clades were seen, a western clade (Portugal, Spain, Italy) and an eastern clade (Romania and Lebanon). Overall intraspecific variation was 1.80 % (range 0.00–3.34 %) with the greatest separation (3.34 %) seen between the Romanian sequences and three identical sequences from Portugal, Spain, and Italy. Variation within the western clade was 0.21 % (range 0.00–0.47 %) and within the eastern clade was 0.40 % (0.00–0.76 %). Although two clades are formed, the lack of morphological differentiation suggests that this is simply isolation by distance, and the current broad concept of A. ventricosa is maintained. In A. brumanensis, intraspecific variation was 1.43 % (range 0.00–3.50 %). Due to the strong genetic isolation combined with morphological differences, the Cretan taxon is therefore newly described below as Andrena testudo sp. nov.</p><p>More broadly, revision of Cryptandrena material from the West Palaearctic (see Suppl. material 2) revealed an interesting pattern by the four members of this subgenus (Fig. 5). In all cases, no more than two Cryptandrena species ever seem to be found in sympatry, these being Andrena rotundata Pérez, 1895 and A. ventricosa in Portugal (see Gaspar et al. 2025), north-west Africa, and Sicily, A. brumanensis and A. ventricosa from south-eastern France to Iran, and A. testudo and A. ventricosa on Crete. Andrena ventricosa therefore appears to be able to co-exist with the other members of the subgenus, but the others do not. This is reminiscent of the situation in the proxima - group (McLaughlin et al. 2022) where Andrena proxima (Kirby, 1802) and Andrena ampla Warncke, 1967 cannot exist in sympatry, but the slightly later flying Andrena alutacea Stöckhert, 1942 can be found at the same sites as A. proxima . McLaughlin et al. (2022) concluded that phenological and Wolbachia - induced mating incompatibilities may maintain species boundaries in this species group.</p><p>Phenologically, A. brumanensis and A. ventricosa are only weakly differentiated (Fig. 6), with A. brumanensis peaking between 16–31 May, and A. ventricosa between 1–15 June. However, both have overall long activity periods from March to the end of July or early August; there may be partial or facultative bivoltinism in parts of the range. Restricting the analysis to just the southern Balkans (815 specimens compared to 1,543 specimens in the full analysis), A. brumanensis peaks at the same period of 16–31 May and A. ventricosa between 1–15 June, suggesting no difference from the overall pattern or any clear evidence of consistent bivoltinisim in this region. The small number of available records for A. rotundata make comparisons difficult; the greatest number of records (nine) occur in the 16–30 June period, but records occur between the beginning of March and the end of July. Overall, some factor is maintaining species boundaries between A. brumanensis and A. ventricosa, and interactions between brumanensis - rotundata - testudo require further study. The status of A. rotundata ssp. diomedia Warncke, 1975 (endemic to Sardinia) also requires further study. It may well deserve species-status.</p></div>	https://treatment.plazi.org/id/5A748AF7ECB652EB9F044750A9885628	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
1EFE51B476275C6B8C125E74B87501D3.text	1EFE51B476275C6B8C125E74B87501D3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Cryptandrena) testudo Wood 2026	<div><p>Andrena (Cryptandrena) testudo sp. nov.</p><p>Figs 66 A – C, 66 E, 66 G, 67 A – E</p><p>Type material.</p><p>Holotype. GREECE • 1 ♀; Crete, Plateau Katharo [<a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=25.5598&amp;materialsCitation.latitude=35.1473" title="Search Plazi for locations around (long 25.5598/lat 35.1473)">W of Katharon</a>]; 35.1473°N, 25.5598°E; 1160 m a. s. l.; 1 Jun. 2023; R. Le Divelec leg.; RMNH, RMNH.INS.1716253; [BOLD accession number ORBIT 190-23].</p><p>Paratypes. GREECE • 1 ♂; Crete, Plateau Katharo [<a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=25.5598&amp;materialsCitation.latitude=35.1473" title="Search Plazi for locations around (long 25.5598/lat 35.1473)">W of Katharon</a>]; 35.1473°N, 25.5598°E; 1160 m a. s. l.; 1 Jun. 2023; R. Le Divelec leg.; RMNH, RMNH.INS.1716254 • 1 ♀; Crete, Plateau Katharo [<a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=25.5598&amp;materialsCitation.latitude=35.1473" title="Search Plazi for locations around (long 25.5598/lat 35.1473)">W of Katharon</a>]; 35.1473°N, 25.5598°E; 1160 m a. s. l.; 1 Jun. 2023; R. Le Divelec leg.; TJWC • 2 ♂; Crete, Dept. Chania, Mescla [Meskla]; 3 May 1973; v. Ooststroom leg.; RMNH, RMNH.INS.1264915 • 1 ♂, 1 ♀; Crete, Lasithi, 15 km NW Males; 850 m a. s. l.; 14 May 2003; O. Sauša leg.; OÖLM • 1 ♂; Crete, SE, Kato Simi; 1000 m a. s. l.; 6 May 2003; O. Sauša leg.; OÖLM • 1 ♂; Kreta sept., Bali; 6–11 May 2003; B. &amp; O. Tkalců leg.; OÖLM • 1 ♀; Kreta, Chania, Lefka Ori, 5 km SW Omalos; 1200–1250 m a. s. l.; 4 Jun. 1993; H. &amp; R. Rausch leg.; OÖLM • 1 ♂; Kreta, Rethimnon [Rethimno]; 19–30 Apr. 1998; H. Wolf leg.; OÖLM .</p><p>Description.</p><p>Female. Body length: 6–6.5 mm (Fig. 66 A). Head: Dark, 1.2 times wider than long (Fig. 66 C). Clypeus domed, medially slightly flattened, surface polished and shining, densely punctate, punctures separated by 0.5–1 puncture diameters. Process of labrum rounded rectangular, 3 times broader than long, surface polished. Gena slightly exceeding width of compound eye; ocelloccipital distance subequal to diameter of lateral ocellus. Foveae dorsally occupying slightly&gt; 1 / 2 space between compound eye and lateral ocellus, poorly defined, ventrally slightly narrowing, still exceeding diameter of scape; facial foveae filled with pure white hairs. Head covered with short whitish hairs, none equalling length of scape. Antennae dark, A 5–12 ventrally lightened by presence of orange-brown scales; A 3 equalling A 4 + 5.</p><p>Mesosoma: Scutum and scutellum polished and shining, slightly irregularly punctate, puncture separated by 0.5–1.5 puncture diameters, scutum posteromedially with punctures separated by almost 2 puncture diameters (Fig. 66 E). Pronotum rounded. Mesepisternum and dorsolateral parts of propodeum irregularly roughened with network of rugosity, surface weakly shining. Propodeal triangle laterally weakly indicated by presence of carina, internal surface covered with raised longitudinal ridges with short lateral carinae (Fig. 66 B). Mesepisternum with finely plumose whitish hairs, becoming shorter and light brown on scutum and scutellum. Propodeal corbicula lacking anterior fringe, dorsal fringe composed of shortly plumose buffish hairs, internal surface with long buffish simple hairs. Legs dark, pubescence light brown, hind tarsal claws with inner subapical tooth. Posterior face of hind femur with row of posteriorly-projecting 6 short tooth-like pegs. Flocculus complete, composed of white plumose hairs; femoral and tibial scopae composed of white predominantly simple hairs, tibial scopae at most with some lateral hairs plumose. Wings hyaline, stigma and venation dark brown, nervulus interstitial.</p><p>Metasoma: Terga dark, apical rims narrowly lightened hyaline brown (Fig. 66 G). Tergal discs polished and shining, densely punctate, punctures separated by 0.5–1 puncture diameters; punctures extending onto tergal margins. Tergal discs with very short and obscure scattered hairs, T 2–4 laterally with dense white apical hairbands, interrupted medially, obscuring underlying surface. T 5 and hairs flanking pygidial plate orange to orange-brown; pygidial plate rounded triangular, surface dull.</p><p>Male. Body length: 5–6 mm (Fig. 67 A). Head: Dark, 1.4 times wider than long (Fig. 67 B). Clypeus dark, structurally as in female. Process of labrum rounded trapezoidal, 2 times wider than long. Gena equalling width of compound eye; ocelloccipital distance slightly shorter than diameter of lateral ocellus. Head covered with moderately long whitish hairs, none equalling length of scape. Antennae dark, A 4–13 ventrally lightened by presence of brownish scales; A 3 exceeding length of A 4, slightly shorter than A 4 + 5 (Fig. 67 E).</p><p>Mesosoma: Scutum and scutellum polished and shining, moderately densely punctate, punctures separated by 1–2 puncture diameters (Fig. 67 C). Pronotum rounded. Mesepisternum and propodeum as in female. Mesosoma covered in long white hairs. Legs dark, apical tarsal segments lightened orange-brown, pubescence whitish to light brownish, hind tarsal claws with small subapical tooth. Wings hyaline, stigma and venation brown, nervulus interstitial.</p><p>Metasoma: Terga dark, apical margins lightened brown to hyaline brown. Tergal discs finely microreticulate, weakly shining, moderately punctate, punctures separated by 1–2 puncture diameters, extending onto marginal areas. Tergal discs covered with short whitish hairs, not forming distinct apical hairbands. S 8 narrow, apex rounded, ventral surface covered with short fringe of light brown hairs. Genital capsule slightly elongate, more-or-less parallel sided, gonocoxal teeth very slightly produced into narrowly rounded points (Fig. 67 D). Gonostyli broadly flattened-triangular apically, surface slightly concave, internal angle raised and rounded. Penis valves long, occupying 1 / 3 space between gonostyli, gradually tapering, acutely pointed apically.</p><p>Diagnosis.</p><p>Andrena testudo can swiftly be recognised as a Cryptandrena due to the presence of a row of short tooth-like spines on the posterior face of the hind femur (females only) combined with the small body size (&lt;7 mm), short and broad process of the labrum (around 3 times broader than long), simple scopal hairs (not plumose), and densely punctate metasomal terga (Fig. 66 G; punctures separated by &lt;1 puncture diameter), as well as by the typical male genital capsule which is relatively long and narrow, with the apices of the gonostyli flattened and weakly triangular with the inner angle upturned, and with long narrow penis valves (Fig. 67 D).</p><p>After excluding Andrena monacha Warncke, 1965 and Andrena aruana Warncke, 1969 which lack spines on the posterior face of the female hind femur and belong in an undescribed subgenus (Pisanty et al. 2022 a; S. Bossert, G. Pisanty, T. J. Wood, unpublished data), only three other species are found in this subgenus: A. brumanensis, A. rotundata, and A. ventricosa . The intriguing geographic distributions are discussed in the section on Subgenus Cryptandrena Pittioni, 1948 (Fig. 5). Andrena testudo can be separated from the most widespread member of the group A. ventricosa due to the male A 3 which is much longer than A 4, and almost as long as A 4 + 5 (Fig. 67 E; in A. ventricosa with the male A 3 essentially equal to the length of A 4, Fig. 67 G) and the female scutum which is relatively sparsely punctate, with punctures separated by 0.5–1.5 puncture diameters (Fig. 66 E; in A. ventricosa with the female scutal punctures separated by around 0.5 puncture diameters, therefore almost confluent).</p><p>The difference from A. brumanensis is subtle, but in side-by-side comparison, the differences can be clearly seen. In both sexes, the clypeus is distinctly longitudinally domed in its median half (Fig. 66 C; in A. brumanensis with the clypeus almost flattened, Fig. 66 D), the male clypeus is entirely dark (in A. brumanensis males with the clypeus showing a yellow median marking of variable size), the female scutum has relatively spaced punctures separated by 0.5–1.5 puncture diameters and approaching 2 puncture diameters posteromedially (Fig. 66 E; in A. brumanensis with the scutum showing punctures separated by 0.5–1 puncture diameters at most, Fig. 66 F), the male scutum has punctures separated by 1–2 puncture diameters and with the interspaces shining (Fig. 67 C; male A. brumanensis have the scutum with punctures separated by &lt;1 puncture diameters, scutum appearing relatively dull), and in the female with the tergal punctation separated by 0.5–1 puncture diameters, interspaces shining, metasoma therefore appearing relatively shiny (Fig. 66 G; in A. brumanensis females with the metasomal punctation denser, separated by &lt;0.5–0.5 puncture diameters, overall metasoma therefore appearing more dull, Fig. 66 H). Andrena brumanensis is also a slightly but distinctly larger bee, with females around 8 mm in length and in side by side comparison appearing broader.</p><p>These same differences allow for separation from A. rotundata, which have the clypeus as flattened as in A. brumanensis, the male A 3 is only slightly longer than A 4 (Fig. 67 H; not almost as long as A 4 + 5 in A. testudo, Fig. 67 E), and the female scutum with punctures separated by at most 1 puncture diameter.</p><p>Etymology.</p><p>From the Latin testudo meaning tortoise in reference to the (relatively) domed clypeus, and also the dense integumental punctation typical of Cryptandrena species which poetically resembles a tight-knit defensive formation, like the testudo formation used by Roman soldiers in the capture of (amongst other cities) Heracleium in Macedon in Ancient Greece in 169 B. C. (described by Livy and Polybius). It is a noun in apposition.</p><p>Distribution.</p><p>Greece (Crete) (Fig. 5).</p></div>	https://treatment.plazi.org/id/1EFE51B476275C6B8C125E74B87501D3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
55EEC9331178586FB911D71C0CFBAB85.text	55EEC9331178586FB911D71C0CFBAB85.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Euandrena) amieti Praz, Muller & Genoud 2019	<div><p>Andrena (Euandrena) amieti Praz, Müller &amp; Genoud, 2019</p><p>Andrena (Euandrena) amieti Praz, Müller &amp; Genoud, 2019: 20, ♀ ♂ [Switzerland, MHNN, not examined]</p><p>Material examined.</p><p>GREECE • 1 ♀; [Peloponnese] top ski center Kalavrita, Ahaia [Achaea]; 20 Jun. 2022; K. Janssen leg.; TJWC • 2 ♀; Chelmos [ Aroania]; 1700 m a. s. l.; 27 May 1987; H. Teunissen leg.; RMNH • 2 ♀; Chelmos, Peloponnes; 2250 m a. s. l.; 25 Aug. 1987; K. Warncke leg.; OÖLM • 74 ♀; Peloponnese, Melano Mountains; 1500 m a. s. l.; 12 May 1987; H. Teunissen leg.; RMNH • 2 ♀; Mt. Olymp. Ost-flanke; 2200–2500 m a. s. l.; 19 Sep. 1989; D. Lange &amp; T. Osten leg.; OÖLM • 3 ♀; Mt. Tymfi / Pindos; 18–2000 m a. s. l.; K. Warncke leg.; OÖLM • 1 ♂; Olymp, Agios Antonis; 2500 m a. s. l.; 2 Aug. 1985; L. Blank leg.; OÖLM • 1 ♀; Olymp, Balkoni N; 1900 m a. s. l.; 31 Jul. 1985; L. Blank leg.; OÖLM • 1 ♀; Olymp.; 2000 m a. s. l.; 24 Aug. 1981; K. Warncke leg.; OÖLM • 2 ♀; Olymp.; 2000–2200 m a. s. l.; 18 Aug. 1978; K. Warncke leg.; OÖLM • 3 ♀; Peloponn., Menalo Mts., Kardaras NW; 1450 m a. s. l.; 20 Apr. 1999; Blank &amp; Kutzscher leg.; DEI • 7 ♂, 1 ♀; Western Macedonia, Florina, 3 km N of Vigla; 1800–1900 m a. s. l.; 13 Jun. 2024; T. J. Wood leg.; TJWC • 1 ♂; Western Macedonia, Florina, immediately north of Vigla; 1500 m a. s. l.; 9 Jun. 2024; T. J. Wood leg.; TJWC ; NORTH MACEDONIA • 1 ♀; Galicia Mt.; 29 Jun. 1985; J. Hladil leg.; OÖLM • 1 ♀; Ochrid, Calicica-Geb, 0.6 km E Livada-Pass; 1595–1650 m a. s. l.; 8 Jul. 2023; E. &amp;. J. Hüttinger leg.; LBCM .</p><p>Notes.</p><p>The first confirmed records from the southern Balkans. As in the Alps, where altitudinal data are available, the species is found at high altitude above 1400 m up to 2500 m. Most records are from the second summer generation, but some specimens from April and May have also been collected.</p></div>	https://treatment.plazi.org/id/55EEC9331178586FB911D71C0CFBAB85	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
DF640D9784C15E2FA30AE78A5F1A42E8.text	DF640D9784C15E2FA30AE78A5F1A42E8.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Euandrena) bulgarica Wood 2026	<div><p>Andrena (Euandrena) bulgarica sp. nov.</p><p>Fig. 68</p><p>Type material.</p><p>Holotype. BULGARIA • 1 ♀; Blagoevgrad, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=23.3106&amp;materialsCitation.latitude=41.4563" title="Search Plazi for locations around (long 23.3106/lat 41.4563)">General Todorov, 2 km E</a>; 41.4563°N, 23.3106°E; 120 m a. s. l.; 19 May 2024; T. J. Wood leg.; RMNH, RMNH.INS.1152454 [BOLD accession number NATBE 081-25].</p><p>Paratypes. BULGARIA • 1 ♀; Blagoevgrad, Sandanski, 3.5 km W, Struma river; 110 m a. s. l.; 23 May 2024; T. J. Wood leg.; RMNH, RMNH.INS.1152503 [BOLD accession number NATBE 112-25] • 1 ♀; Petrich env., steppes near Struma river; 100 m a. s. l.; 10 May 2018; D. Benda leg.; NMPC [BOLD accession number WPATW 994-23] • 3 ♀; Slanĉev Brjag [Sunny Beach]; 10 Jun. 1972; M. Kocourek leg.; OÖLM • 1 ♀; Slanĉev Brjag [Sunny Beach]; 1–30 Jun. 1972; M. Kocourek leg.; OÖLM ; GREECE • 1 ♀; Western Macedonia, Filotas, 3 km N of Antigonos; 600–700 m a. s. l.; 11 Jun. 2024; T. J. Wood leg.; RMNH, RMNH.INS.1152611 [BOLD accession number NATBE 173-25] .</p><p>Description.</p><p>Female. Body length: 10–11 mm (Fig. 68 A). Head: Dark, 1.2 times wider than long (Fig. 68 B). Clypeus weakly domed, medially somewhat flattened, surface polished and shining; densely punctate, punctures separated by &lt;0.5–1 puncture diameters, punctures denser laterally, becoming slightly sparser apically. Process of labrum weakly trapezoidal, 2.5 times wider than long, apical margin broadly and shallowly emarginate. Gena slightly exceeding width of compound eye; ocelloccipital distance 1.5 times diameter of lateral ocellus. Foveae dorsally occupying 1 / 3 space between compound eye and lateral ocellus, narrowing ventrally to just below lower margin of antennal insertions; facial foveae filled with dark brown hairs. Head covered with brown hairs, light brown on face and lower part of gena, becoming golden brown on vertex; occasional black hairs on upper frons laterally; no hairs equalling length of scape. Antennae dark, A 5–12 ventrally slightly lightened by presence of greyish scales; A 3 equalling A 4 + 5.</p><p>Mesosoma: Scutum anteriorly and laterally shagreened, weakly shining, becoming polished and shining in posteromedial 1 / 2 (Fig. 68 C); surface densely punctate, punctures separated by 0.5–1 puncture diameters over majority of surface, becoming slightly sparser medially. Scutellum dull posteriorly and laterally, medially broadly shining, irregularly punctate, punctures separated by 0.5–2 puncture diameters. Pronotum rounded. Mesepisternum and dorsolateral parts of propodeum finely microreticulate, covered with shallow irregular punctures, separated by 0.5–2 puncture diameters on mesepisternum, density increasing to 0.5–1 puncture diameters on propodeum; surface dull. Propodeal triangle laterally not delineated by carinae, internal surface impunctate, covered with fine granular shagreen, basomedially with network of fine wrinkles. Mesepisternum with brown hairs, pale whitish brown ventrally, becoming golden brown dorsally (Fig. 68 D); hairs on scutum and scutellum shorter, densely plumose but not subsquamous, golden brown (Fig. 68 E). Propodeal corbicula lacking anterior fringe, dorsal fringe composed of long golden brown plumose hairs, internal surface with long partially-plumose golden hairs. Legs dark, pubescence dark brown, hind tarsal claws with inner subapical tooth. Flocculus complete, composed of buff plumose hairs; femoral and tibial scopae composed of simple orange hairs. Wings hyaline, stigma and venation dark orange-brown, nervulus interstitial to weakly antefurcal.</p><p>Metasoma: Terga dark, apical rims narrowly lightened hyaline brown (Fig. 68 F). Tergal discs predominantly polished and shining, with obscure weak sculpturing most visible at base of discs. Tergal discs irregularly punctate, punctures separated by 0.5–2 puncture diameters, extending onto tergal margins, becoming slightly sparser and less distinct. Tergal discs with short hairs, pale on T 1–3, black on T 4–5, not obscuring underlying surface. Marginal area of T 1 with loose light brown hairs, marginal areas of T 2–4 with apical hairbands of pale hairs, complete in fresh specimens, obscuring underlying surface. T 5 and hairs flanking pygidial plate dark brown; pygidial plate rounded triangular, surface dull and featureless.</p><p>Male. Unknown.</p><p>Diagnosis.</p><p>Andrena bulgarica can be recognised as a Euandrena due to the moderate body size, facial foveae which dorsally occupy 1 / 3 of the space between the compound eye and a lateral ocellus and which further narrow ventrally (comma or drop-shaped), A 3 = A 4 + 5, and lack of any other distinctive characters (simple tibial scopae, simple propodeal triangle, posterior face of hind femur rounded, hind tibial spurs unmodified). Recognition is challenging, and requires experience within the group; use of the identification key is recommended to learn the major divisions within the subgenus in the southern Balkans.</p><p>Due to the pubescence which is brown to golden brown on the mesosomal dorsum (Fig. 68 D) and the tibial scopae which are orange (Fig. 68 A), it can be placed in the group of species around A. bicolor . The clypeus is uniformly punctate and lacks a weak median longitudinal impunctate line medially and the pronotum is rounded, excluding A. glidia . The most similar species are consequently A. aff. limosa and A. bicolor . Andrena bulgarica can be separated due to the facial pubescence which is almost entirely light brown with at most a few black hairs along the inner margins of the compound eyes (Fig. 68 B; variable, either extensively black-haired in A. bicolor or partially black-haired in A. aff. limosa, but in all cases with clear black hairs on the clypeus), clypeus densely punctate but with the underlying surface and hence interspaces strongly polished and shining (Fig. 68 B; in A. bicolor and A. aff. limosa with the clypeus densely punctate with interspaced shagreened and shagreened and weakly shining to dull), mesepisternum with pubescence light brown to golden brown, without black hairs (Fig. 68 D; highly variable, but usually in A. bicolor with black hairs on the mesepisternum, in A. aff. limosa usually with the mesepisternum with entirely brown hairs), and scutum widely polished and shining over most of its posterior half (Fig. 68 C; in A. aff. limosa with the scutum also extensively polished medially, in A. bicolor the condition is quite variable, sometimes with small shining area medially, sometimes almost entirely dull; this character is variable between Levantine populations which always have the scutum dull to European populations which are most extensively polished). A final character is body size, with A. bulgarica reasonably large for a Euandrena at 10–11 mm in length, whereas A. bicolor and A. aff. limosa tend to average a little smaller at 8–10 mm on average. This character can only be appreciated with series assembled together, at which point it becomes easier to observe. Through this combination of characters, it is possible to diagnose A. bulgarica, but consultation of barcoded or reliably determined specimens is preferable until the concept has been appreciated.</p><p>Notes.</p><p>All records to date come from low to moderate altitude sites (100–700 m a. s. l.); based on current evidence, A. bulgarica does not seem to be a species of mountains.</p><p>Etymology.</p><p>Named after the country of Bulgaria, which currently lacks an eponymous species given the synonymy of Andrena bulgariensis Warncke, 1965 with Andrena inconstans Morawitz, 1877 (Wood 2023 a). It is an adjective in the feminine singular nominative form.</p><p>Distribution.</p><p>Bulgaria (Blagoevgrad, Burgas) and Greece (Western Macedonia).</p></div>	https://treatment.plazi.org/id/DF640D9784C15E2FA30AE78A5F1A42E8	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
343CC20491E15D388F86E2DAD9D466A2.text	343CC20491E15D388F86E2DAD9D466A2.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Euandrena) glabriventris Alfken 1935	<div><p>Andrena (Euandrena) glabriventris Alfken, 1935</p><p>Andrena glabriventris Alfken, 1935: 158, ♀ [Turkey, SMFD, examined]</p><p>Material examined.</p><p>GREECE • 1 ♀; Western Macedonia, Anatoliko, 2.8 km E, fields; 750 m a. s. l.; 15 Jun. 2024; T. J. Wood leg.; RMNH, RMNH.INS.1152469 ; TURKEY • 4 ♀; 10 May 1934; A. Seitz leg.; SMFD (holotype and paratypes) • 1 ♀; 20 km E Karakurt / Arastal; 12 Jun. 1977; K. Warncke leg.; OÖLM • 1 ♀; Acigel [Acıgöl]; 19 May 1999; K. Deneš leg.; TJWC • 1 ♀; Kars, 15 km E Karakurt; 1460 m a. s. l.; 2 Jun. 1988; K. Warncke leg.; OÖLM • 1 ♀; Kulu / Konya; 1000 m a. s. l.; 3 Jun. 1978; M. Kraus leg.; OÖLM • 3 ♂, 4 ♀; N Güzelsu / Van; 9 Jun. 1977; K. Warncke leg.; OÖLM • 1 ♀; Şereflikoçhisar; 24 May 1972; K. Warncke leg.; K. Warncke det.; OÖLM .</p><p>Notes.</p><p>Andrena glabriventris strongly resembles A. bicolor in its colour pattern with orange-brown hairs on the mesosoma dorsally (Fig. 13 A), becoming black on the mesepisternum (Fig. 13 C), dark facial hairs (Fig. 13 B), and with orange tibial scopae. However, it can be recognised due to the irregular tergal punctation, with punctures separated by 1–3 puncture diameters and with polished and shining interspaces (Fig. 13 D). In A. bicolor, the tergal punctation is much more regular, with interspaces typically not exceeding the diameter of a puncture on average. The sculpture of the interspaces is variable, but polished shining interspaces are typically not visible.</p><p>Distribution.</p><p>Greece * and Turkey (Alfken 1935; Warncke 1974 a; Pisanty et al. 2022 b).</p></div>	https://treatment.plazi.org/id/343CC20491E15D388F86E2DAD9D466A2	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
244775BA77955F2E8B9AC1E35F4E4B44.text	244775BA77955F2E8B9AC1E35F4E4B44.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Euandrena) glidia Warncke 1965	<div><p>Andrena (Euandrena) glidia Warncke, 1965</p><p>Andrena glidia Warncke, 1965 a: 49, ♀ ♂ [Greece, OÖLM, examined] (Fig. 14)</p><p>Material examined.</p><p>GREECE • 1 ♀; Graecia [no further information]; OÖLM (holotype) . See Suppl. material 2 for additional examined material.</p><p>Notes.</p><p>Warncke unhelpfully selected the oldest possible specimen to serve as holotype, as was sometimes his wont. Andrena glidia has traditionally been considered a species of south-eastern Europe and Turkey, but the available DNA barcodes and examined specimens support an extension of its range south into the Levant (Fig. 15). Warncke (1968: 87) placed A. glidia in the Subgenus Ptilandrena Robertson, 1902 (currently restricted to North American species only, see Pisanty et al. 2022 a) due to the presence of a pronotal angle (in both sexes) and a broadened gena that exceeded the width of the compound eye (males only). However, some specimens in the original type series have weak to absent pronotal angles, and specimens from the Levant are much more challenging and ambiguous. Specimens from Israel lack pronotal angles, and display a dull scutum compared to specimens from the southern Balkans (G. Pisanty, pers. comm.). At the present time, a full understanding of this morphological variation is not possible, even though the species appears to be very well characterised genetically. Whilst the classical morphological characters as used by Warncke are here included in the identification key, this substantial caveat is here emphasised and users of the key are urged to treat carefully in this challenging group.</p><p>This caveat notwithstanding, A. glidia has an unusual pattern of activity; it appears to be bivoltine, with a generation in the spring and another generation in the autumn to early winter. Warncke (1974 a: 113) gives activity in April, with additional November activity based on a single female specimen. Based on specimens examined here, the activity period can be described as March (Cyprus, Greece; n specimens = 3), April (Cyprus, Greece, Greece: Crete, Israel, Jordan, Lebanon, Turkey; n = 25), May (Greece, Greece: Crete, Israel, Turkey; n = 18), June (Greece, Greece: Crete, Turkey; n = 6), October (Greece: Crete; n = 17), and December (Cyprus, Israel; n = 11). This would appear to support a broad spring generation centred on April and a poorly characterised autumn-early winter generation.</p><p>Distribution.</p><p>? Croatia, Greece, Turkey, Cyprus, Lebanon *, Israel *, Jordan * (Warncke 1965 a, 1974 a; Gusenleitner and Schwarz 2002; Fig. 15). The Croatia listing comes from a single dot on the Warncke distribution maps presented by Gusenleitner and Schwarz (2002: 1052). It has not been possible to examine any specimens from this country; it it not impossible that it actually refers to a specimen of A. aff. limosa (see below).</p></div>	https://treatment.plazi.org/id/244775BA77955F2E8B9AC1E35F4E4B44	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
CBD8EAEC96CE5EB191C0929CAD33921F.text	CBD8EAEC96CE5EB191C0929CAD33921F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Euandrena) Hedicke 1933	<div><p>Subgenus Euandrena Hedicke, 1933</p><p>Genetics.</p><p>The Subgenus Euandrena is very young by Andrena standards (Pisanty et al. 2022 a), and contains a swarm of morphologically variable and challenging to characterise taxa. The situation is particularly acute in the East Mediterranean, where multiple faunas meet. There are also many mountainous areas; these landscape features create many areas with phenologically distinct microclimates, and seem to support high Euandrena species richness when considered at a global scale (Osytshnjuk et al. 2008; Praz et al. 2019; Wood et al. 2021; Wood 2021, 2023 b, 2023 c, 2024 a).</p><p>Since the work of Praz et al. (2019) which resolved taxon delineation issues in Central European Euandrena, a large number of sequences have now becomes available from the East Mediterranean. This is pertinent, as Praz et al. left a number of questions open; whilst elevating Andrena pileata Warncke, 1975 and treating it distinct from Andrena allosa Warncke, 1975 (Spain, France, Switzerland). Andrena pileata was described from Chelmos mountain in the Peloponnese, and Praz et al. considered the described paratype male not to be conspecific with the holotype male. They also identified three uncertain taxa supported by DNA barcodes which could not at the time be named; sp. 1, sp. 2, and sp. 3. Analysis of DNA barcodes, including the most pertinent barcodes of Praz et al., provides resolution to these open questions (Fig. 7). To begin with, sp. 2 sensu Praz et al. (HYMAA 038-18) comes from the island of Lesbos, and almost perfectly matches sequences of Andrena robusta Warncke, 1975 from Lesbos and Samos (separated by 0.26 %); the species was originally described from the island of Rhodes. This question at least is straightforward to answer.</p><p>More challenging is sp. 1 sensu Praz et al.; sequences from Chelmos and Mount Olympus (HYMAA 037-18; HYMAA 043-18) placed adjacent to a long series of sequences of A. pileata from Mount Maenalon (Peloponnese), Mount Parnassus (south Central Greece), and the Pirin Mountains (south-western Bulgaria) (Fig. 8), separated by an average genetic distance of 2.02 % (range 1.67–2.44 %). As in the analysis of Praz et al., sp. 1 was placed close to A. allosa, but the addition of new Balkan sequences has resulted in a topology where clades of A. pileata and A. allosa are reciprocally monophyletic, although bootstrap support is relatively low, potentially indicating an unstable topology. Taking A. pileata (inclusive of sp. 1), it was separated from A. allosa by an average genetic distance of 1.46 % (range 0.95–2.28 %); excluding sp. 1, A. pileata was separated by an average genetic distance of 1.35 % (range 0.95–1.67 %). sp. 1 alone was separated from A. allosa by an average genetic distance of 1.89 % (range 1.55–2.28 %).</p><p>Adjacent to A. allosa are two sequences, NATBE 120-25 which is a female Euandrena from Petrich (Bulgaria) on low altitude grassland at the Rupite mineral baths (20 May 2024) and WPATW 1645-25 which is a male Euandrena from Armenia. They were separated from A. allosa by an average genetic distance of 2.22 % (range 1.27–3.19 %). The male specimen is badly abraded and essentially uncharacterisable given the challenges associated with identification of isolated male Euandrena specimens. The female specimen shows the densely plumose scutal hairs typical of allosa + pileata, but the clypeus is comparatively slightly shorter and the terga are shinier than in A. pileata . Moreover, it was collected at an altitude of 90 m a. s. l., when the one available sequenced specimen of A. pileata from Bulgaria was collected at an altitude of approximately 2,000 m a. s. l. (3.5 km NNE of Popina Luka).</p><p>Given what is known about Euandrena and their ecological and altitudinal niches, and the likely break in gene flow between low and high altitude in this group, it is difficult to imagine these being conspecific. In this context, it is best to currently consider A. allosa and A. pileata (inclusive of sp. 1) as distinct following Praz et al. (2019), and to consider the Bulgarian + Armenian sequences of uncertain identity at the present time until more data become available. Praz et al. (2019) also considered that Warncke’s paratype male of A. allosa ssp. pileata was not conspecific with the holotype (even though they were collected in sympatry, just two days apart), but based on the examination of a barcoded male of A. pileata (NATBE 056-25; Fig. 9), it seems to match Warncke’s specimen in all aspects including the shape of the genital capsule, and so Warncke’s existing male paratype is considered to have been correctly associated.</p><p>Before moving to sp. 3 sensu Praz et al., it is also necessary to report Andrena amieti Praz, Müller &amp; Genoud, 2019 as new for the southern Balkans. Four sequences were available from northern Greece collected at an altitude of 1800–1900 m a. s. l. which were separated from A. amieti group 1 (Switzerland, including the holotype; bootstrap support of 86) sequences by an average genetic distance of 0.96 % (range 0.91–1.07 %) and from A. amieti group 2 (Switzerland and Italy; bootstrap support of 95) by an average genetic distance of 2.45 % (range 2.31–2.58 %). Compared to the analysis of Praz et al. (2019: 16), all Andrena amieti sequences formed a monophyletic clade with bootstrap support of 86, rather than fall into two distinct clades which were interrupted by the clade of A. allosa . The Greek clade of A. amieti received bootstrap support of 89, and overall seems to have have helped resolve the topology, grouping all three clades together with bootstrap support of 86. Although there is slight population structuring, the morphology of A. amieti remains recognisable over this range. Examination of 111 A. amieti specimens from Greece and North Macedonia (see Suppl. material 2) shows activity between 12 May and 25 August, supporting the bivoltine behaviour reported in Switzerland by Praz et al. (2019). In this context, A. amieti can be considered a species found in high mountains in the Alps, southern Apennines, and southern Balkans (Fig. 8). Collectively, covering all three clades, a low average intraspecific genetic distance of 1.04 % (range 0.00–2.74 %) was observed. The distribution is not unlike that of Andrena montana Warncke, 1973; the one available sequence from northern Greece (Mount Kajmakčalan) clustered with sequences from Switzerland, showing minimal average genetic differentiation of 0.25 % (range 0.15–0.46 %).</p><p>For sp. 3 sensu Praz et al. (HYMAA 036-18), the sequence clustered with a long series of sequences from Bulgaria, Greece, Lebanon, and Jordan. This group of sequences requires discussion, and an analysis of what Warncke referred to as Andrena rufitibialis Friese, 1899 in a broad sense. Friese (1899: 342) described A. rufitibialis from two female specimens from 13 Apr. 1899 collected at Jericho by Francis Morice. Gusenleitner and Schwarz (2002: 647) give ZMHB or OUMNH as the possible type repositories. Examination of these collections produced three female specimens (Figs 10, 11), but only the specimen in the ZMHB was actually labelled by Friese with one of his red type labels. Friese’s type material is often difficult to interpret (see Rasmussen and Ascher 2008); although the OUMNH material may well be syntypic, it is best to treat as only possibly syntypic, and the female in the ZMHB collection illustrated here is designated as the lectotype. It was later redetermined as Andrena biguttata Friese, 1923 by Stöckhert for inexplicable reasons, given that this name is younger, regardless of specimen morphology.</p><p>Warncke (1969 a) listed A. rufitibialis, and then described Andrena rufitibialis ssp. limosa Warncke, 1969 from Turkey (locus typicus), Syria, and Israel which is darker, with the metasoma entirely dark, but still with partial reddish colouration of the hind tibiae (Fig. 12). This reddish colouration of the hind tibiae is absent on many “ dark ” specimens that can be collected in the southern Balkans and the Levant. Rasmont et al. (2017: 18) elevated Andrena rufitibialis ssp. limosa to species status based on personal communication from Erwin Scheuchl and reported it as new for Greece based on specimens from the East Aegean Islands. No species concept was discussed, or global range indicated, for example whether Turkish material is conspecific with dark material from the southern Balkans and Levant. However, this elevation is justified due to their differing morphologies combined with the ecological affinities of the taxa; A. rufitibialis s. str. is a very rarely collected species of arid habitats (see Material examined below), and is completely absent from the Mediterranean zone in the Levant whereas darker specimens (lacking extensive red tergal markings, A. limosa sensu lato) are common throughout the Mediterranean zone (G. Pisanty, pers. comm.).</p><p>In the current analysis, it has not been possible to generate a barcode from either i) a specimen with orange hind tibiae and abundant red markings on the terga ( A. rufitibialis s. str.) or ii) a generally dark specimen still displaying partially orange hind tibiae ( A. limosa), but many sequences are available from entirely dark specimens which form a broad clade from southern Bulgaria to Lebanon and north-western Jordan (Fig. 7). It is necessary to highlight the sequence NATBE 129-25 which was generated from a specimen that was part of a long series of dark specimens collected from Campanula ( Campanulaceae) flowers near Gorno Spanchevo in south-western Bulgaria. It shows no morphological differentiation from other specimens from the same collecting event (e. g. NATBE 128-25, NATBE 131-25), but shows a level of genetic differentiation from all remaining dark specimen sequences of 3.38 % (range 2.93–3.67 %). It is unclear if this is variation or if the sequence has problems, for example, if it was generated from nuclear mitochondrial DNA (a NUMT), sequences that are notoriously difficult to differentiate from genuine mitochondrial fragments (e. g. Williams et al. 2020, 2024). If it is excluded, the clade receives bootstrap support of 98 and the average intraspecific variation is 0.82 % (range 0.00–2.69 %), and if included then average intraspecific variation is 1.08 % (range 0.00–3.67 %). In any case, inclusion or exclusion does not affect the overall topology of the clade. Based on the morphology of these specimens, including their dark legs, it is not clear if they actually correspond to A. limosa . Until it is possible to generate a sequence from a specimen displaying at least partially orange legs (ideally from Turkey), to be as cautious as possible these sequences are referred to as A. aff. limosa (Fig. 7). Based on the available sequences, A. aff. limosa is still well-differentiated from its nearest neighbour Andrena glidia Warncke, 1965 by an average genetic distance of 5.69 % (range 4.84–6.23 %). The two species are in principle easy to separate in the male sex (at least in the southern Balkans) due to the male head morphology, but it is unclear if these characters are stable (see comments in the Andrena glidia section below) and females can be challenging; differences are given in the identification key below.</p><p>The very widespread and common species Andrena bicolor Fabricius, 1775 is one of the most challenging Andrena species in the West Palaearctic due to its abundance, widespread distribution, and variable morphology, particularly pubescence colouration. It has been described many times from across its range (Gusenleitner and Schwarz 2002), and has acted as something of a waste bin for difficult to characterise or place Euandrena taxa. Warncke lumped many names under a broad A. bicolor concept, and the process of revalidating valid species or confirming existing synonymies from this “ waste bin ” concept is ongoing (Praz et al. 2019; Wood 2023 b, 2023 c). Generation of A. bicolor sequences from the East Mediterranean confirms the presence of a broad A. bicolor taxon across the southern Balkans and into the Levant (Fig. 7). No comparative pattern of two moderately separated clades as seen in the analysis of Praz et al. (2019) was found, with complex structuring observed, though a focused analysis including all available sequences would be necessary to investigate this possible pattern further. One sequence of Andrena bicolor ssp. apricaria Warncke, 1975 from Cyprus was placed as the base of the bicolor - clade, but was grouped with a sequence (NATBE 253-25) of a morphologically ambiguous specimen from Belgium. The status of ssp. apricaria must wait for a future analysis. Taking a broad concept of A. bicolor inclusive of the Cypriot and Belgian sequences show an average intraspecific genetic distance of 3.37 % (range 0.00–6.01 %). The greatest distance of 6.01 % was seen between the specimen from Cyprus and a specimen from the Sierra Nevada mountain in southern Spain (WPATW 290-21).</p><p>What can be resolved here is that sequences were available from a Euandrena taxon from southern Bulgaria and northern Greece which is noticeably larger and shinier than typical A. bicolor specimens in the southern Balkans. Four sequences were generated which were consistently placed at the base of a broad bicolor - clade with bootstrap support of 100. They were separated from the bicolor - clade by an average genetic distance of 5.99 % (range 5.22–6.91 %). Based on this consistent genetic differentiation and observed differences in size, integument sculpture, and existence in sympatry with A. bicolor s. str., they are described below as Andrena bulgarica sp. nov.</p><p>Andrena fulvata Stöckhert, 1930 sequences from Greece and Bulgaria currently do not have strong genetic support, being more closely placed next to Andrena angustior (Kirby, 1802) sequences than A. fulvata sequences from Germany and Slovenia (Fig. 7). Based on their observed morphology and knowledge of the western- and eastern-biased distribution of the two taxa, respectively (Wood et al. 2021), assignment to A. fulvata is made without doubts. Greek and Bulgarian sequences were separated from German and Slovenian sequences by an average genetic distance of 3.45 % (range 3.16–3.99 %), and from A. angustior by an average genetic distance of 3.17 % (range 2.80–3.53 %). Alignment with either clades is therefore somewhat arbitrary purely based on genetic distance and without consideration of specimen morphology. It may well be the case that additional sampling in countries such as Serbia and Romania produces intermediate sequences and results in a change in the topology of the tree, and a monophyletic A. fulvata is formed. Currently, bootstrap support for Greek and Bulgarian A. fulvata + A. angustior is moderate at 77. Additional study is required, and the current morphological concept of A. fulvata is maintained (Schmid-Egger and Scheuchl 1997; Wood et al. 2021).</p><p>Finally, specimen NATBE 091-25 is a female Euandrena from near to Anatoliko in West Macedonia (Greece) collected in fields at an altitude of 750 m a. s. l. (15 Jun. 2024). It was strongly separated from all other Euandrena taxa studied here, and matched a sequence of Andrena glabriventris Alfken, 1935 from Turkey (G. Pisanty, unpublished data). It morphologically corresponds to that species, and is therefore newly reported from Greece and Europe; it is treated below.</p></div>	https://treatment.plazi.org/id/CBD8EAEC96CE5EB191C0929CAD33921F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
60909A0CFA885D73A153153C24FEC5F4.text	60909A0CFA885D73A153153C24FEC5F4.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Euandrena) limosa Warncke 1969	<div><p>Andrena (Euandrena) limosa Warncke, 1969</p><p>Andrena (Euandrena) rufotibialis [sic] ssp. limosa Warncke, 1969 a: 408, ♀ ♂ [Turkey, OÖLM, examined] (Fig. 12)</p><p>Material examined ( Andrena limosa).</p><p>TURKEY • 1 ♀; Konya; 15 Jun. 1965; M. Schwarz leg.; OÖLM (holotype); SYRIA • 1 ♂; N. Syria, Ladikye [Latakia]; 1–31 May 1885; E. Leuthner leg.; OÖLM (paratype) .</p><p>( Andrena aff. limosa, indicative). ALBANIA • 1 ♀; Gjirokastër [ SW of Këlcyrë]; 1 Jun. 2024; W. Vertommen leg.; WVC ; BULGARIA • 1 ♀; 30 km SW Svilengrad; 550 m a. s. l.; 21 Jun. 2008; M. &amp; Z. Halada leg.; OÖLM • 1 ♀; S Petrich vill. [near Sofia]; 593 m a. s. l.; 20 May 2016; T. Ljubomirov leg.; IBER ; CROATIA • 2 ♀; Split, Gornji Muc, MT [malaise trap]; 500 m a. s. l.; 14 May – 8 Jun. 2014; B. Kokan leg.; OÖLM ; JORDAN • 3 ♀; 15 km W Jerash, Dibbin; 2 May 2006; K. Deneš leg.; TJWC • 1 ♀; Aljoun; 28 Apr. 2012; M. Kafka leg.; TJWC • 1 ♀; Irbid, Saham vill; 25 Apr. 2003; I. Pljushtch leg.; TJWC ; LEBANON • 2 ♀; Bcharre d., Aabdine; 950 m a. s. l.; 12 May 2003; V. Soon leg.; TUZ, TUZ 346124 • 1 ♀; Mount Lebanon Governorate, Jabal Moussa Biosphere Res.; 1400 m a. s. l.; 14 May 2023; V. Soon leg.; TUZ, TUZ 341652 ; ROMANIA • 1 ♀; SW, Resita, Anina; 4 Jun. 2000; M. Snížek leg.; OÖLM . See Suppl. material 2 for additional examined specimens.</p><p>Notes.</p><p>Following the discussion above, A. limosa is treated as a species distinct from A. rufitibialis, but doubts are expressed as to whether the commonly encountered species with dark legs that is found in the southern Balkans and represented here in the genetic sampling is actually conspecific with it. These records are presented as A. aff. limosa, and are subject to change following future study of populations in Turkey and the Levant.</p><p>Distribution.</p><p>Croatia *, Romania *, Albania *, Bulgaria *, Greece, Turkey, Syria, Lebanon *, Israel &amp; West Bank, Jordan * (Fig. 15; Warncke 1969 a, 1974 a; Gusenleitner and Schwarz 2002; Rasmont et al. 2017, inclusive of A. aff. limosa).</p></div>	https://treatment.plazi.org/id/60909A0CFA885D73A153153C24FEC5F4	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
09217BB91307552B9B5EA5165FE414DB.text	09217BB91307552B9B5EA5165FE414DB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Euandrena) obscuricrus Wood 2026	<div><p>Andrena (Euandrena) obscuricrus sp. nov.</p><p>Fig. 69</p><p>Type material.</p><p>Holotype. GREECE • 1 ♀; Creta, Lassithi Ebene [Plateau]; 24 Apr. 1973; H. Teunissen leg.; RMNH, RMNH.INS.1716258.</p><p>Paratypes. GREECE • 3 ♀; Lasithi Vlakte [Plateau]; 17 Apr. 1982; H. Teunissen leg.; RMNH, RMNH.INS.1716259 • 1 ♀; Creta, Omalos; 2 Jun. 1987; Kudrna leg.; OÖLM • 1 ♀; W. Kreta, Rhodhákinon [Ano Rodakino] nahe S. Küste; 20 Mar. 1925; A. Schulz leg.; OÖLM • 1 ♀; Kreta, Mesa Potami; 28 Apr. 2004; R. &amp; W. Zarre leg.; OÖLM • 1 ♀; Kreta, Spili; 4 Jun. 1980; M. Kraus leg.; OÖLM .</p><p>Description.</p><p>Female. Body length: 8.5–10 mm (Fig. 69 A). Head: Dark, 1.2 times wider than long (Fig. 69 B). Clypeus weakly domed, surface covered with fine granular microreticulation, basally dull, becoming weakly shining apically; surface with large and distinctly separated punctures separated by 0.5–3 puncture diameters, typically by 1–2 puncture diameters. Process of labrum rounded trapezoidal, 2 times wider than long, apical margin straight. Gena slightly exceeding width of compound eye; ocelloccipital distance slightly exceeding diameter of lateral ocellus. Foveae dorsally occupying 1 / 3 space between compound eye and lateral ocellus, narrowing ventrally to just below lower margin of antennal insertions; facial foveae filled with brown hairs. Face with whitish hairs on clypeus and around antennal insertions, with black hairs along inner margins of compound eyes and frons. Gena ventrally with whitish hairs, becoming weakly golden on vertex; no hairs equalling length of scape. Antennae dark, A 5–12 ventrally slightly lightened by presence of greyish scales; A 3 slightly exceeding A 4 + 5, shorter than A 4 + 5 + 6.</p><p>Mesosoma: Scutum and scutellum finely microreticulate, dull, surface with irregular scattered shallow punctures separated by 0.5–3 puncture diameters (Fig. 69 C). Pronotum rounded. Mesepisternum with fine granular microreticulation, dull. Dorsolateral parts of propodeum with fine granular microreticulation, microreticulation slightly raised, forming network of weak pseudopunctures, surface dull. Propodeal triangle narrow, weakly indicated, internal surface with dense fine granular microreticulation, with raised rugae in basal 1 / 3–1 / 2 (Fig. 69 D). Mesepisternum with intermixed light brown and black hairs, scutum and scutellum with shorter light brown hairs intermixed with shorter dark hairs. Propodeal corbicula lacking anterior fringe, dorsal fringe composed of mixture of light brown and black plumose hairs, internal surface with weakly plumose light brown hairs. Legs dark, apical tarsal segments brownish, pubescence brown, hind tarsal claws with inner subapical tooth. Flocculus complete, composed of pale plumose hairs; femoral and tibial scopae composed of pale golden to orange hairs. Tibial scopae composed of mixture of simple and obscurely plumose hairs (Fig. 69 E). Wings hyaline, stigma and venation dark orange-brown, nervulus interstitial.</p><p>Metasoma: Terga dark, marginal areas obscurely lightened brownish, apical rims narrowly lightened hyaline-whitish (Fig. 69 F). Tergal discs with fine granular microreticulation, weakly shining; surface obscurely punctate, with small scattered very fine punctures, appearing broadly impunctate. Tergal discs with scattered short hairs, marginal areas of T 2–4 with weak apical hairbands laterally, widely interrupted, not obscuring underlying surface. T 5 and hairs flanking pygidial plate dark brown; pygidial plate rounded triangular, lateral margins impunctate, remaining surface densely punctate, punctures almost confluent; surface dull.</p><p>Male. Unknown.</p><p>Diagnosis.</p><p>Andrena obscuricrus can be recognised as a Euandrena due to the moderate body size, facial foveae which dorsally occupy 1 / 3 of the space between the compound eye and a lateral ocellus and which further narrow ventrally (comma or drop-shaped), A 3 slightly exceeding A 4 + 5, and lack of any other distinctive characters (simple propodeal triangle, posterior face of hind femur rounded, hind tibial spurs unmodified). It should be noted that, unusually for a West Palaearctic Euandrena, the tibial scopae of A. obscuricrus is composed of partially plumose hairs (Fig. 69 E). It is difficult to neatly categorise this as either simple or plumose.</p><p>Warncke (1965 a: 50) listed Andrena (Euandrena) ruficrus Nylander, 1848 from “ Westkreta ” based on a specimen collected by Schulz on 20 Mar. 1925. It has been possible to examine a specimen with this collecting information, which now forms part of the type series. The two are genuinely morphologically similar due to the largely impunctate terga, dull and shallowly punctate scutum, and predominantly pale facial vestiture with the exception of black hairs along the inner margins of the compound eyes. However, apart from the strong geographic separation (the nearest confirmed records of A. ruficrus come from around Croatia and Hungary, with two additional specimens from Mount Olympus in Greece; Gusenleitner and Schwarz 2002, see below), A. obscuricrus can be separated due to its clypeus which has large and distinctly separated punctures which are separated by 0.5–3 puncture diameters but typically by 1–2 puncture diameters (Fig. 69 B; in A. ruficrus with the clypeus covered in small regular punctures separated by 0.5–1 puncture diameters), the dark legs (Fig. 69 E; hind tibiae and tarsi lightened orange in A. ruficrus), and the partially plumose tibial scopae (tibial scopae with simple hairs in A. ruficrus).</p><p>Andrena obscuricrus is more likely to be confused with A. glidia which occurs on Crete (Fig. 15). However, A. obscuricrus can be rapidly recognised due to the rounded pronotum ( A. glidia with the pronotum laterally displaying a subtle but distinct humeral angle), entirely dull scutum (Fig. 69 C; A. glidia with the scutum extensively polished and shining, at least medially), sparsely punctate clypeus ( A. glidia with the clypeus covered with dense punctures separated by 0.5–1 puncture diameters), pale facial vestiture with black hairs along the inner margins of the compound eyes ( A. glidia with uniformly light brown facial vestiture), and tergal discs almost impunctate (Fig. 69 F; tergal discs of A. glidia with weak but distinct punctures separated by around 1–2 puncture diameters, most visible laterally).</p><p>Etymology.</p><p>Since ruficrus (rufus + crus) means red-legged in Latin, and the Cretan specimens have dark legs, they are named obscuricrus from obscurus + crus. It is a compound noun.</p><p>Distribution.</p><p>Greece (Crete) (Warncke 1965 a, as A. ruficrus misidentification).</p></div>	https://treatment.plazi.org/id/09217BB91307552B9B5EA5165FE414DB	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
A410AAE3125D55EBA244BA1E0462D304.text	A410AAE3125D55EBA244BA1E0462D304.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Euandrena) pileata Warncke 1975	<div><p>Andrena (Euandrena) pileata Warncke, 1975</p><p>Andrena (Euandrena) allosa ssp. pileata Warncke, 1975: 85, ♀ ♂ [Greece, OÖLM, examined]</p><p>Andrena (Euandrena) allosa ssp. canigica Warncke, 1975: 86, ♀ [Turkey, OÖLM, examined] (see Wood 2024 a)</p><p>Material examined.</p><p>BULGARIA • 1 ♀; Blagoevgrad, Popina Luka, 3.5 km NNE, path to Spano Pole hut; 1800–2000 m a. s. l.; 18 May 2024; T. J. Wood leg.; RMNH, RMNH.INS.1152448 • 1 ♀; Pirin, Kozia dolina [valley]; 2000 m a. s. l.; 30 May 1999; J. Straka leg.; JSPC • 1 ♀; Pirin, Popina Luka [Попина Лъка]; 1800 m a. s. l.; 14 Jul. 1966; M. Kocourek leg.; OÖLM ; GREECE • 1 ♀; Chelmos, Peloponnes; 1900 m a. s. l.; 2 Jun. 1962; M. Schwarz leg.; OÖLM (holotype of Andrena allosa ssp. pileata) • 1 ♂; Chelmos, Peloponnes; 1900 m a. s. l.; 1 Jun. 1962; M. Schwarz leg.; OÖLM (paratype of Andrena allosa ssp. pileata) • 1 ♀; Chelmos, Peloponnes; 1900 m a. s. l.; 1 Jun. 1962; M. Schwarz leg.; TJWC • 6 ♀; Central Greece, Mt. Parnassos, immediately NW of ski centre; 1750 m a. s. l.; 9 Apr. 2024; T. J. Wood leg.; RMNH / TJWC • 2 ♀; Chelmos [Aroania]; 1700 m a. s. l.; 27 May 1987; H. Teunissen leg.; RMNH, RMNH.INS.1264657 • 1 ♂, 4 ♀; Peloponnese, Mount Maenalon, 4.5 km NW Kardaras; 1600 m a. s. l.; 4 Apr. 2024; T. J. Wood leg.; RMNH, RMNH.INS.1152409 . See Wood (2024: 95) for examined material from Turkey.</p><p>Notes.</p><p>The dot maps of Warncke (Gusenleitner and Schwarz 2002) indicate the presence of A. pileata (as Andrena allosa ssp. pileata) in southern Bulgaria. This can be confirmed with modern samples from the Pirin Mountains (Fig. 8).</p><p>Distribution.</p><p>Bulgaria, Greece, Turkey (Warncke 1974 a, 1975; Praz et al. 2019; Wood 2024 a).</p></div>	https://treatment.plazi.org/id/A410AAE3125D55EBA244BA1E0462D304	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
27B0377866765C41B18F78C0B5696A13.text	27B0377866765C41B18F78C0B5696A13.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Euandrena) rufitibialis Friese 1899	<div><p>Andrena (Euandrena) rufitibialis Friese, 1899</p><p>Andrena rufitibialis Friese, 1899: 342, ♀ [West Bank, ZMHB, lectotype by present designation] (Fig. 10)</p><p>Material examined.</p><p>ISRAEL • 1 ♀; Kinereth [Sea of Galilee]; 13 Apr. 1959; SMNHTAU, SMNH-TAUI 331515 ; SYRIA • 1 ♂; Damascus; [probably 1899]; O. Schmiedeknecht leg.; K. Warncke det.; OÖLM; WEST BANK • 1 ♀; Jericho; 13 Apr. 1899; F. Morice leg.; ZMHB (lectotype by present designation) • 1 ♀; Jericho; 13 Apr. 1899; F. Morice leg.; OUMNH; ENT-HYME 294-01 (possible syntype) • 1 ♀; Jericho; 13 Apr. 1899; F. Morice leg.; OUMNH; ENT-HYME 294-02 (possible syntype) .</p><p>Notes.</p><p>Following the discussion above, A. rufitibialis is considered in a narrow sense (only specimens matching the lectotype), giving it a small distribution in drier parts of the Levant.</p><p>Distribution.</p><p>Syria, Israel &amp; West Bank (Fig. 15; Friese 1899; Warncke 1969 a).</p></div>	https://treatment.plazi.org/id/27B0377866765C41B18F78C0B5696A13	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
67F99BF7D52D58DAA7B649236895E783.text	67F99BF7D52D58DAA7B649236895E783.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Graecandrena) cretica Wood 2026	<div><p>Andrena (Graecandrena) cretica sp. nov.</p><p>Fig. 70 A – C, E, F</p><p>Type material.</p><p>Holotype. GREECE • 1 ♀; Crete, Plateau Katharo [<a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=25.5598&amp;materialsCitation.latitude=35.1473" title="Search Plazi for locations around (long 25.5598/lat 35.1473)">W of Katharon</a>]; 35.1473°N, 25.5598°E; 1160 m a. s. l.; 1 Jun. 2023; R. Le Divelec leg.; RMNH, RMNH.INS.1716255; [BOLD accession number ORBIT 180-23].</p><p>Paratypes. GREECE • 1 ♀; Crete, Plateau Katharo [ W of Katharon]; 1160 m a. s. l.; 1 Jun. 2023; R. Le Divelec leg.; RMNH, RMNH.INS.1716256 • 5 ♀; Crete, Foinikilas [ E of Agios Ioannis]; 450 m a. s. l.; 6 Jun. 2023; R. Le Divelec leg.; RLDC • 1 ♀; Crete, Mousoures [ W of Prases]; 500 m a. s. l.; 9 Jun. 2023; R. Le Divelec leg.; RLDC • 1 ♀; Crete, Dept. Chania, Mescla [Meskla]; 3 May 1973; v. Ooststroom leg.; RMNH, RMNH.INS.1264834 • 1 ♀; Kriti, Spilia – Drakona, 25 km W of Chania; 18 May 1982; J. P. Duffels leg.; RMNH; ZMA.INS.5103925 .</p><p>Description.</p><p>Female. Body length: 5 mm (Fig. 70 A). Head: Dark, 1.25 times wider than long (Fig. 70 C). Clypeus weakly domed, finely shagreened and weakly shining, moderately punctate, punctures separated by 1–3 puncture diameters. Process of labrum small, narrowly trapezoidal, slightly wider than long. Gena slightly exceeding width of compound eye; ocelloccipital distance slightly &lt;1 / 2 diameter of lateral ocellus. Foveae occupying 1 / 3 space between pound eye and lateral ocellus, equalling width of flagellum, moderately impressed, ventrally extending below lower level of antennal insertions; foveae filled with light brown hairs. Head covered with pale to light brown hairs, none equalling length of scape. Antennae dark, A 5–12 ventrally lightened by presence of greyish scales; A 3 equalling A 4 + 5.</p><p>Mesosoma: Scutum and scutellum very weakly sculptured, almost polished and shining, weakly punctate, punctures separated by 1–3 puncture diameters (Fig. 70 E). Pronotum rounded. Mesepisternum and dorsolateral parts of propodeum with fine granular microreticulation, dull to weakly shining. Propodeal triangle broad, laterally not differentiated from dorsolateral parts of propodeum, basally with greatly reduced rugae, present only immediately adjacent to the junction between the propodeum and metanotum, not exceeding the width of a flagellum in length (Fig. 70 B). Mesepisternum with whitish densely plumose hairs, becoming shorter and sparser on scutum and scutellum. Propodeal corbicula lacking anterior fringe, dorsal fringe composed of long plumose whitish hairs, internal surface with long scattered simple pale hairs. Legs dark, apical tarsal segments lightened brownish, pubescence light brown. Flocculus incomplete, composed of long scarcely curving white plumose hairs, femoral and tibial scopae composed of white simple hairs. Inner tarsal claws with inner subapical tooth. Wings hyaline, stigma and venation brownish orange, nervulus interstitial.</p><p>Metasoma: Terga dark, apical rims narrowly lightened hyaline brown (Fig. 70 G). Tergal discs with fine granular microsculpture, completely impunctate, weakly shining. Tergal discs with scattered short hairs, not forming apical hairbands. T 5 and hairs flanking pygidial plate orange-brown; pygidial plate narrowly rounded triangular, surface dull.</p><p>Male. Unknown.</p><p>Diagnosis.</p><p>Andrena cretica can be recognised as a Graecandrena due to the small body size, hind tarsal claws with a small inner subapical tooth, and propodeal triangle which is predominantly covered with granular shagreen with the exception of a narrow section at its base with very short rugae (Fig. 70 B). Due to the short ocelloccipital distance (less than the diameter of a lateral ocellus), it is closest to A. hyemala (mainland southern Balkans) and A. repressa (the island of Rhodes only). It can be separated from both comparison species (treated together) by the clypeus which is finely shagreened and weakly shining (Fig. 70 C; in comparison species with the clypeus microreticulate and dull, Fig. 70 D), propodeal triangle with basal rugae greatly reduced, present only immediately adjacent to the junction between the propodeum and metanotum, not exceeding the width of a flagellum in length (Fig. 70 B; in comparison species with the propodeal triangle more extensively wrinkled basally, rugae extending to the midpoint of the propodeal triangle, exceeding the width of a flagellum in length), scutum very weakly sculptured, almost polished and shining (Fig. 70 E; in comparison species with the scutum shagreened, at most weakly shining, Fig. 70 F), and frons with weak and superficial sculpturing, with only hints of longitudinal striations (Fig. 70 G; in comparison species with the frons dull, showing raised longitudinal striations, Fig. 70 H). Andrena cretica is also very slightly smaller at 5 mm in length compared to 6 mm in the comparison species, but a series of specimens is required to appreciate this difference. As demonstrated above, the biogeography of A. cretica suggests a non-overlapping distribution, with neither A. hyemala or A. repressa present on Crete.</p><p>Etymology.</p><p>Named after the island of Crete. It is an adjective in the feminine singular nominative form.</p><p>Distribution.</p><p>Greece (Crete) (Fig. 23).</p></div>	https://treatment.plazi.org/id/67F99BF7D52D58DAA7B649236895E783	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
47FEEF5D04635E129B8D8C331A553781.text	47FEEF5D04635E129B8D8C331A553781.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Graecandrena) repressa (Warncke 1975) Warncke 1975	<div><p>Andrena (Graecandrena) repressa Warncke, 1975 stat. nov.</p><p>Andrena hyemala ssp. repressa Warncke, 1975: 62, ♀ ♂ [Turkey, OÖLM, examined] Figs 21 B, 21 D, 21 F.</p><p>Material examined (illustrative).</p><p>CYPRUS • 1 ♀; Paphos, Polis, Chrysochous; 10 m a. s. l.; 1 Jun. 2023; P. Rosa leg.; UMONS ; GREECE • 1 ♂, 4 ♀; Rhodos; 23 Apr. 1976; K. Vegeter leg.; RMNH • 1 ♂; 23 Apr. 1970; v. Oostroom leg.; RMNH; ISRAEL • 2 ♀; Mas’ada [Masada]; 19 Apr. 1988; M. Kraus leg.; OÖLM • 6 ♂, 1 ♀; Mt. Carmel, En Hod; 4 Apr. 1999; S. P. M. Roberts leg.; SPMRC • 10 ♂, 3 ♀; Hasharon, Zikhron Ya’aqov, Ramas Hanadiv; 25 Mar. 1990; R. Leys leg.; RMNH ; JORDAN • 1 ♀; 20 km N Amman; 620 m a. s. l.; 23 Apr. 2006; K. Deneš leg.; OÖLM • 3 ♀; 30 km NW Ajlun [Ajloun]; 600 m a. s. l.; 29 Apr. 2006; K. Deneš leg.; OÖLM / TJWC • 4 ♀; Ajlun [Ajloun] env; 840 m a. s. l.; 1 May 2006; K. Deneš leg.; OÖLM / TJWC LEBANON • 3 ♀; Beqaa, Beqaa valley, Mansourah, Aammiq wetland preserve; 850 m a. s. l.; T. J. Wood leg.; TJWC • 2 ♂; Beqaa, Beqaa valley, Mansourah, Kafraiya village; 1000 m a. s. l.; T. J. Wood leg.; TJWC • 5 ♀; Beqaa, Beqaa valley, Qaraoun, 3.5 km W of Madjal Balhis; 900 m a. s. l.; T. J. Wood leg.; TJWC • 2 ♂; Mount Lebanon, Hazerta; c. 1400 m a. s. l.; 6 May 1996; C. G. Roche leg.; OUMNH ; SYRIA • 1 ♀; Jisr al-Shughur [written as Jisr-esh-Shughur], Syria; 18 Apr. 1992; K. Warncke leg.; OÖLM • 1 ♀; occ. An Nasrah env [Al-Nasrah]; 8–13 Apr. 2005; J. Saki leg.; OÖLM • 1 ♂, 11 ♀; Orontes, 30 km SW Homs; 500 m a. s. l.; 15 Apr. 1992; K. Warncke leg.; OÖLM • 1 ♀; Stausee [Dam], 500 m, 10 km SW Homs; 15 Apr. 1992; K. Warncke leg.; OÖLM TURKEY • 1 ♂, 1 ♀; Adana; 6 Apr. 1971; K. Warncke leg.; OÖLM (male holotype, female paratype) .</p><p>Notes.</p><p>Andrena repressa was originally described from Turkey (Adana), with additional material from Greece (Rhodes), and Israel (Ramla). A selection of records is presented above; full details can be found in the Suppl. material 2.</p><p>Distribution.</p><p>Greece (Rhodes), Turkey, Cyprus *, Syria *, Lebanon *, Israel, Jordan * (Fig. 23; Warncke 1975, as A. hyemala ssp. repressa).</p></div>	https://treatment.plazi.org/id/47FEEF5D04635E129B8D8C331A553781	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
B2516C398EDC5E26B3AC93BEB0306AB3.text	B2516C398EDC5E26B3AC93BEB0306AB3.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Graecandrena) Warncke 1968	<div><p>Subgenus Graecandrena Warncke, 1968</p><p>Genetics.</p><p>Sequences of Andrena hyemala Warncke, 1973 were available from continental Greece (Peloponnese) and Crete, and sequences of A. hyemala ssp. repressa Warncke, 1975 were available from Rhodes, Lebanon, and Israel (Fig. 20). However, the sequence from Crete was surprisingly strongly separated from that of continental Greece. Whilst A. hyemala s. str. from continental Greece was separated from A. h. repressa by an average genetic distance of 8.09 % (range 7.75–8.36 %), it was also separated from the Cretan sequence by 8.89 %. The separation between the Cretan “ A. hyemala ” and A. h. repressa was also high at an average genetic distance of 7.08 % (range 6.95–7.27 %). These strong genetic distances combined with morphological differences mean that it is appropriate for A. repressa stat. nov. to be treated at a specific level, and for Cretan specimens to be described as Andrena cretica sp. nov. Morphologically, A. repressa can most easily be recognised in the male sex through comparison of the genital capsule and specifically the width of the penis valves (Fig. 21), broader in A. hyemala (Fig. 21 E) and narrower in A. repressa (Fig. 21 F); full diagnosis is made through the identification key below which includes A. hyemala, A. repressa, and A. cretica, as well as the diagnosis section in the description of A. cretica . Formal elevation is carried out below.</p><p>Additionally, comment is necessary on the placement of Andrena helenica Warncke, 1965 . Warncke (1968: 58) placed A. helenica in the Subgenus Graecandrena, but A. helenica does not show the typical genital capsule form seen in most members of the group, and the female has very narrow facial foveae which could be indicative of the Distandrena Warncke, 1968 group (= Micrandrena Ashmead, 1899). Doubts also existed about Andrena verticalis Pérez, 1895 which was only tentatively placed in the Graecandrena until Pisanty et al. (2022 a) confirmed this placement. Andrena helenica sequences placed most closely to A. verticalis (Fig. 20), and whilst this is weak evidence, it is enough to maintain its current subgeneric placement. Andrena helenica remains endemic to Greece, extending north to Siatista and Trikala (Warncke 1965 a; Fig. 22).</p></div>	https://treatment.plazi.org/id/B2516C398EDC5E26B3AC93BEB0306AB3	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
0C2C585EB273523BB93282A54ED129B4.text	0C2C585EB273523BB93282A54ED129B4.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Holandrena) decipiens Schenck 1861	<div><p>Andrena (Holandrena) decipiens Schenck, 1861</p><p>? Apis albifrons Christ, 1791: 191 [Germany, type lost] nec. Forster, 1771 and Rossi, 1790.</p><p>Andrena decipiens Schenck, 1861: 250, ♀ ♂ [Germany, SMFD, examined]</p><p>Andrena fonscolombii Dours, 1872: 357, ♀ ♂ [Greece: Corfu, MSNG, examined]</p><p>? Andrena insolita Dours, 1872: 422, ♀ ♂ [Spain, Algeria, possibly MNHN, not examined]</p><p>? Andrena strigosa Dours, 1872: 428, ♀ ♂ [France, types lost]</p><p>Andrena flavilabris Schenck, 1874: 170 [Germany, SMFD, examined] syn. nov.</p><p>Andrena deceptoria Schmiedeknecht, 1884: 745, ♀ ♂ [France, types lost]</p><p>Andrena fimbriata var. paganettii Alfken, 1904: 294, ♀ ♂ [Greece: Corfu, ZMHB, not examined]</p><p>Andrena decipiens ssp. capta Warncke, 1974 b: 46, ♀ ♂ [Morocco, OÖLM, examined]</p><p>Material examined.</p><p>ALBANIA • 1 ♀; Fier region, Lushnje env., pastures and steppes; 177 m a. s. l.; 29 Jun. 2018; D. Benda leg.; JSPC ; GERMANY • 1 ♀; Wiesbaden; Schenck collection; SMFD (syntype of Andrena decipiens) • 1 ♀; Bamburg; Schenck collection; SMFD (syntype of Andrena flavilabris) ; GREECE • 1 ♂, 1 ♀; Corfu; Gribodo collection; MSNG (male lectotype and female paralectotype of Andrena fonscolombii) ; LEBANON • 1 ♂; Kherbet Kanafar [Kherbet Qanafar], Bekaa; 16 Jun. 1953; G. Schmidt leg.; OÖLM ; MOROCCO • 1 ♀; El Hajeb; 22 Jul. 1932; Ad. Nadig leg.; OÖLM (holotype of Andrena decipiens ssp. capta) • 1 ♂; Ifrane; 30 May 1973; H. Teunissen leg.; OÖLM (paratype of Andrena decipiens ssp. capta) . See Suppl. material 2 for additional examined material.</p><p>Notes.</p><p>Wood and Bossert (2025: 287) located type material of A. fonscolombii, which was actually described from Corfu and was not a junior synonym of A. variabilis as inferred by Warncke (1967 b: 272).</p><p>Distribution.</p><p>Morocco, Algeria, Spain, Andorra, France, Switzerland, Germany, Austria, Czechia, Slovakia, Hungary, Italy, Malta, Slovenia, Croatia, Bosnia &amp; Herzegovina, Serbia, Romania, Poland, Belarus, Ukraine, Russia (European part), Montenegro, Albania *, Greece, Bulgaria, Turkey, Lebanon *, Georgia, Armenia, Azerbaijan, Iran (Dours 1872, as A. fonscolombii; Warncke 1965 a, 1974 a, 1974 b; Gusenleitner and Schwarz 2002; Osytshnjuk et al. 2008; Standfuss et al. 2011; Wood 2021, 2023 c, as A. flavilabris; Wood and Bossert 2025). Records from Tajikistan and Uzbekistan presented by Wood and Monfared (2022: 74) are here considered to be misidentifications of A. labialis, rendering A. decipiens a strictly West Palaearctic species.</p></div>	https://treatment.plazi.org/id/0C2C585EB273523BB93282A54ED129B4	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
82E259E9B10B524AB4C5221CCD13683E.text	82E259E9B10B524AB4C5221CCD13683E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Holandrena) forsterella Osytshnjuk 1978	<div><p>Andrena (Holandrena) forsterella Osytshnjuk, 1978</p><p>Andrena fimbriata Brullé, 1833 sensu auctorum (Warncke 1965 a: 57, ♀ ♂)</p><p>Andrena forsterella Warncke, 1967 b: 205–206 nom. nov. pro. Andrena fimbriata sensu auctorum (nomen nudum)</p><p>Andrena forsterella Osytshnjuk, 1978: 318, 321, 337, 350, ♀ ♂ [Azerbaijan, lectotype designated by Schönitzer et al. 1995: 829, SIZK, not examined]</p><p>Andrena wilhelmi Schuberth, 1995: 808, ♀ ♂ [Greece, ZSM, partially examined] syn. nov.</p><p>Material examined.</p><p>ALBANIA • 1 ♂; Butrint NP; 2 Jul. 2021; M. Jacobs leg.; KJCB • 2 ♂, 1 ♀; Buz [Buzë]; 16 Jul. 2017; K. Janssen leg.; KJCB • 1 ♀; Novaj [3 km N Therepelë]; 18 Jul. 2017; K. Janssen leg.; KJCB • 1 ♀; rivier Gjerbës; 21 Jul. 2017; K. Janssen leg.; KJCB • 3 ♂; Xhafaj; 15 Jul. 2017; K. Janssen leg.; KJCB ; AZERBAIJAN • 1 ♂; Sabirabad; 5 May 1969; ZSM (labelled as paratype of A. wilhelmi) ; GREECE • 1 ♂; Olympia; 24 Apr. 1964; W. Grünwaldt leg.; ZSM (labelled as “ n. sp. ”, probable paratype of A. wilhelmi) • 1 ♂; Olympia; 24 Apr. 1964; W. Grünwaldt leg.; ZSM (labelled as “ n. sp. ”, probable paratype of A. wilhelmi) • 1 ♂; Olympia; 24 Apr. 1964; W. Grünwaldt leg.; ZSM (labelled as paratype of A. wilhelmi) . See Suppl. material 2 for additional examined material.</p><p>Notes.</p><p>Concerning the type material of A. wilhelmi, the holotype male from Olympia (24 Apr. 1964) is supposed to be deposited in the ZSM, but examination of this collection could only produce three males from this collecting event (Fig. 27), only one of which was labelled as paratype. Three paratype males from this collection event are listed by Schuberth (1995: 816), and so they are likely simply all paratypes, but the precise location of the holotype is currently unclear. Regardless, they are representative of the taxon and there is no doubt as to what the name applies to.</p><p>Distribution.</p><p>Italy, Croatia, Albania *, North Macedonia, Bulgaria, Greece, Turkey, Ukraine, Crimea, Cyprus, Syria, Lebanon, Israel &amp; West Bank, Georgia, Armenia, Azerbaijan, Iran (Fig. 24; Schuberth 1995; Schönitzer et al. 1995; Osytshnjuk et al. 2008).</p></div>	https://treatment.plazi.org/id/82E259E9B10B524AB4C5221CCD13683E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
DA3CBBA9958F517ABD42BE287AE88EAA.text	DA3CBBA9958F517ABD42BE287AE88EAA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Holandrena) Perez 1890	<div><p>Subgenus Holandrena Pérez, 1890</p><p>Genetics.</p><p>There are two major issues within this subgenus which must be resolved, specifically the statuses of the two pairs of nominally univoltine taxa which fly in the spring and summer, respectively, these being Andrena flavilabris Schenck, 1874 – Andrena decipiens Schenck, 1861 and Andrena wilhelmi Schubert, 1995 – Andrena forsterella Osytshnjuk, 1978 . Whilst Warncke (1967 b, 1974 a, 1974 b) treated both “ pairs ” as bivoltine species, but variable treatments have been produced since them which are dissected below.</p><p>Andrena forsterella is discussed first, as the creation of this name has been complex, requiring an understanding of changing concepts presented by Warncke (1965 a, 1967 b). A summary was presented by Schönitzer et al. (1995), but in short, Andrena forsterella Warncke, 1967 is an invalid name as it does not meet ICZN (1964, 1985, 1999) standards. Under Article 13.1. 3, a name created after 1930 must be be proposed expressly as a new replacement name (nomen novum) for an available name – Andrena forsterella Warncke, 1967 was proposed as a replacement name for a sensu auctorum, i. e. not an available name. The correct authority is Andrena forsterella Osytshnjuk, 1978 .</p><p>This background is important to note, as A. forsterella is a problematic taxon; originally considered to be bivoltine, the “ first ” generation was described as a new species by Schuberth (1995) as Andrena wilhelmi Schubert, 1995 . The separation of A. wilhelmi from A. forsterella is morphologically challenging in females if the collecting date information has been removed. Schuberth cites primarily the slightly smaller size, darker colouration, longer hairs, less strongly developed tergal hairbands, and yellow facial markings in the male restricted to the clypeus (and not extending onto the lower paraocular areas). However, these are characters which are subject to intergenerational variation due to developing and flying in different seasons; the different generations of Andrena variabilis Smith, 1853 also display these types of colouration and pubescence differences. For structural differences, Schuberth cites slightly sparser tergal punctation, the slightly more strongly upturned fore margin of the male clypeus, and more strongly striate gena. However, slight differences in sculpture are also subject to intergenerational variation, as can be seen in Andrena (Micrandrena) Ashmead, 1899 such as in the widely accepted to be bivoltine Andrena minutula (Kirby, 1802) and Andrena minutuloides Perkins, 1914 . Schuberth asserts that the characters are clear, and that they cannot be subject to seasonal variation, but evidence to support this claim is required given the pattern of morphological intergenerational variation often seen in bivoltine Andrena .</p><p>Schuberth (1995: 818) noted that the two taxa seemed to avoid occupying the same sites, at least at a small geographic scale (Der Vergleich der Verbreitungskarten der beiden Arten erweckt vielmehr den Anschein, als würden sich die beiden Arten zumindest kleinflächig meiden). However, examination of 123 specimens of A. wilhelmi and 127 specimens of A. forsterella from across their known ranges (see Suppl. material 2 for full details) make it difficult to believe this idea (Fig. 24). Whilst only a small quantity of summer-flying material from the Caucasus has been examined and none from Crimea, Schönitzer et al. (1995) presented material from Azerbaijan (including the lectotype designation of A. forsterella) and northern Iran along the Caspian Sea, and Osytshnjuk (1978) cited the presence of A. forsterella in Crimea. Therefore, combining the specimen records presented here and by Schuberth (1995) and Schönitzer et al. (1995), A. wilhelmi and A. forsterella are considered to have completely overlapping ranges. The consistent northern boundary of both taxa can be seen, for example, across Bulgaria, and both taxa extend consistently into the Levant (Fig. 24).</p><p>Genetic evidence provides a level of clarity (Fig. 25). Spring-flying sequences were available from Israel and Lebanon (n = 2), and summer-flying sequences were available from Italy and Israel (n = 3). All sequences were grouped together with bootstrap support of 100. If treating the taxa as distinct, average intraspecific distance for A. wilhelmi was 0.30 %, and average intraspecific distance for A. forsterella was 0.41 % (range 0.30–0.46 %). Average “ interspecific ” genetic distance was 0.41 % (range 0.15–0.61 %). Treating them as a single species showed average intraspecific variation of 0.40 % (range 0.15–0.61 %). Unsurprisingly, these very low genetic distances strongly support just a single species that is bivoltine. When compared to other Holandrena species, high interspecific distances are seen, such as an average of 9.12 % (range 8.03–9.58 %) between Andrena labialis (Kirby, 1802) and Andrena variabilis . Consequently, A. wilhelmi syn. nov. is synonymised under A. forsterella; this change is formalised below, and A. forsterella is integrated into the identification key as a bivoltine species.</p><p>Finally, it is appropriate to discuss the status of Andrena flavilabris Schenck, 1874 as distinct from Andrena decipiens Schenck, 1861 . Long considered (with a degree of uncertainty) to be just the spring generation of A. decipiens (e. g. Schönitzer et al. 1995; Schuberth 1995), Mandery et al. (2008) returned A. flavilabris to species status based on morphological and 16 S ribosomal DNA genetic data, specifically divergence of 3 % between A. decipiens and A. flavilabris . Using the different COI marker, two clades were detected in a contemporary analysis. One contained four sequences identified as A. flavilabris which showed average intraspecific variation of 0.08 % (range 0.00–0.16 %). The second clade was composed of eight sequences showing intraspecific variation of 0.00 % (range 0.00–0.00 %). The two clades were separated by an average genetic distance of 1.99 % (range 1.11–2.51 %), and consequently appeared visually well-separated, although the clade with four sequences only received bootstrap support of 56 compared to 96 for the clade with eight sequences (Fig. 25).</p><p>However, the clade of eight sequences contained six specimens identified as A. decipiens and two sequences (CODAB 349-21 and WPATW 1776-25) identified as A. flavilabris . The specimen CODAB 349-21 was caught on 16 Apr. 2012 in France, and WPATW 1776-25 was caught on 8 May 2025 in central Spain. The issue is illustrated by specimens from central Spain. WPATW 1776-25 (a female specimen) and WPATW 1777-25 (a male specimen) were identified as A. flavilabris and were caught a few days apart in May 2025 in the provinces of Segovia and Guadalajara, but showed a genetic difference of 1.11 %. They consequently fell into the two observed clades. Specimen WPATW 1776-25 was caught just 6 kilometers east of sequence WPATW 327-21 that was caught on 9 Jul. 2021, and the two sequences were identical.</p><p>There is therefore a lack of congruence between the morphology, the observed flight date, and the genetic analysis. Looking at other lines of evidence, in Germany, both A. decipiens and A. flavilabris have reappeared simultaneously at several sites in recent years after decades without evidence of their presence, and A. flavilabris in the spring flies to the same nesting sites as A. decipiens in the summer (E. Scheuchl, pers. comm.). This strongly suggests that they are the same bivoltine species. Inspection of museum material illustrates that A. decipiens and A. flavilabris have completely overlapping ranges (Fig. 26), as is the case in the bivoltine A. forsterella . The two taxa share the same northern and western limit, and the south-eastern limit is a little blurry but is limited by the availability of museum specimens representing spring sampling in Lebanon and Iran. For the barcoding results, as the mitochondrial genome is inherited matrilineally, there is no recombination and these mixed results could be as a result of incomplete lineage sorting. The fact that specimens flying at the same time in close geographical proximity (central Spain, May 2025) showed barcodes from the two clades, and that specimens flying in both spring and summer in near sympatry (central Spain, July 2021 and May 2025) showed identical barcodes strongly suggests that the observed genetic distance is not biologically meaningful. Consequently, the broad concept of a bivoltine A. decipiens (as used by Schönitzer et al. 1995; Schuberth 1995; Osytshnjuk et al. 2008) is returned to, and the change is formalised below.</p></div>	https://treatment.plazi.org/id/DA3CBBA9958F517ABD42BE287AE88EAA	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
0FD5B70DF0D85B13A16F22924C19E59E.text	0FD5B70DF0D85B13A16F22924C19E59E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Lepidandrena) caprimulga Warncke 1975	<div><p>Andrena (Lepidandrena) caprimulga Warncke, 1975</p><p>Andrena (Lepidandrena) caprimulga Warncke, 1975: 94, ♀ ♂ [Turkey, OÖLM, examined] (Fig. 17)</p><p>Andrena (Lepidandrena) statusa Gusenleitner, 1998: 119, ♀ ♂ [Turkey, OÖLM, examined] syn. nov.</p><p>Material examined.</p><p>LEBANON • 1 ♂; Beqaa, Beqaa valley, Qaraoun dam; 850 m a. s. l.; 4 Apr. 2023; T. J. Wood leg.; TJWC • 1 ♂; Tannourine el Tahta, Wadi Ain al Raha; 936 m a. s. l.; 11 Apr. 2017; M. Boustani leg.; TJWC ; ISRAEL • 1 ♂; 850 m S Bet Oren; 256 m a. s. l.; 31 Mar. 2012; A. Dorchin leg.; ADC • 1 ♂, 1 ♀; Galilea, 10 km E of Nahariyya; 9 Apr. 1988; R. Leys leg.; RMNH • 5 ♂, 16 ♀; Hasharon, Zikhron Ya’aqov, Ramas Hanadiv; 12 Mar. 1990; R. Leys leg.; RMNH / TJWC • 1 ♂, 3 ♀; Hasharon, Zikhron Ya’aqov, Ramas Hanadiv; 25 Mar. 1990; R. Leys leg.; RMNH • 1 ♂; Jerusalem env; 7 Apr. 1946; Bytinski-Salz leg.; OÖLM (paratype of Andrena statusa) • 3 ♀; Mt Carmel, Denia; 8 Apr. 1999; S. P. M. Roberts leg.; SPMRC • 3 ♀; Mt. Gilboa, Merav [Meirav], 0.33 km W; 353 m a. s. l.; 20 Mar. 2021; A. Dorchin leg.; ADC • 1 ♂; Tiberias; 26 Mar. 1996; H. Wolf leg.; OÖLM ; TURKEY • 2 ♀; Horasan, Arastal; 14 Jun. 1973; K. Warncke leg.; OÖLM (holotype and stylopised paratype of Andrena caprimulga) • 4 ♂, 3 ♀; Abanoz (Rte Anamur Kazenci), Taurus; 1200 m a. s. l.; 10 May 1991; H. Teunissen leg.; RMNH • 3 ♀; Tanin-Tanin-Pass; 1700 m a. s. l.; 12 Jun. 1984; K. Warncke leg.; OÖLM / TJWC • 14 ♀; E of Şırnak / Siirt; 4 Jun. 1977; K. Warncke leg.; OÖLM (paratypes of Andrena statusa) • 1 ♀; Uludere, Hakkari; 5 Jun. 1977; K. Warncke leg.; OÖLM (holotype of Andrena statusa) ; WEST BANK • 1 ♂; Judea, Har Gilo; 6 Apr. 1988; R. Leys leg.; RMNH .</p><p>Distribution.</p><p>Crimea, Turkey, Lebanon, Israel &amp; West Bank (Fig. 18; Warncke 1974 a, 1975; Osytshnjuk et al. 2008; Pisanty et al. 2018, as A. statusa; Wood et al. 2020, as A. statusa). Although the key presented by Osytshnjuk et al. (2008: 142) includes only the female sex and not the distintive male, the listing from Crimea can be confirmed through personal correspondence from Erwin Scheuchl:</p><p>“ I own one female and three genitalized males from Crimea:</p><p>1 ♂, 18. 06. 1997 \ Crimea, Balaklava District \ W. Cholodorov leg.</p><p>1 ♂, 18. 06. 1997 \ Crimea, Balaklava Area, Chernorech \ W. Cholodorov leg.</p><p>1 ♀, 09. 06. 1997 \ Crimea, Balaklava Area, Chernorech \ W. Cholodorov leg.</p><p>1 ♂, 06. 06. 1997 \ Ukraine, Crimea, Area Tylovoje Village \ Ivanov leg.</p><p>[Chernorechye = Chernorechye, 44.55°N, 33.68°E, Tylovoje Village = Tylove Village.</p><p>Osytshnjuk actually first encountered the male of A. caprimulga in 1997 during her stay in Munich at the Grünwaldt Collection. The female from Alushta, which Osytshnjuk used to include it in the Central and Eastern Palaearctic Key, was confirmed by Grünwaldt as A. caprimulga ”.</p></div>	https://treatment.plazi.org/id/0FD5B70DF0D85B13A16F22924C19E59E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
638E34D3F5725BC6AA7D8747F6070449.text	638E34D3F5725BC6AA7D8747F6070449.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Lepidandrena) haladai Wood 2026	<div><p>Andrena (Lepidandrena) haladai sp. nov.</p><p>Figs 71, 72 A, B</p><p>Type material.</p><p>Holotype. GREECE • 1 ♂; Central Greece, Erateini, 3 km N of <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=22.2553&amp;materialsCitation.latitude=38.3998" title="Search Plazi for locations around (long 22.2553/lat 38.3998)">Panormos</a>; 38.3998°N, 22.2553°E; 600 m a. s. l.; 10 Apr. 2024; T. J. Wood leg.; RMNH, RMNH.INS.1148903 [BOLD accession number NATBE 001-25].</p><p>Paratypes. GREECE • 1 ♂; Central Greece, Erateini, 3 km N of Panormos; 600 m a. s. l.; 10 Apr. 2024; T. J. Wood leg.; RMNH, RMNH.INS.1148904 • 1 ♂; Peloponnes / Elis, Lambia-Geb, 5 km SW Lambia; 21 May 1992; H. Rausch leg.; OÖLM • 1 ♂; Peloponnese, 12 km NWW Sparti; 26 May 2005; M. Kadlecová leg.; OÖLM • 4 ♂; Peloponnese, 35 km NW Tripoli, Langadia SW [Lagkadia]; 3 May 2005; M. Kadlecová &amp; J. Halada leg.; OÖLM .</p><p>Description.</p><p>Female. Unknown.</p><p>Male. Body length: 8–9 mm (Fig. 71 A). Head: Dark, 1.2 times wider than long (Fig. 71 B). Clypeus weakly domed, basally and laterally shagreened and dull, medially becoming polished and shining; surface punctate, punctures separated by 0.5–2 puncture diameters. Process of labrum rounded trapezoidal, apical margin emarginate, slightly thickened and upturned. Surface of galea polished and shining over majority of surface, becoming shagreened only apically. Gena slightly exceeding width of compound eye; ocelloccipital distance slightly exceeding diameter of lateral ocellus. Head covered with long whitish hairs, some black hairs laterally on paraocular areas; longest hairs not equalling length of scape. Antennae basally dark, A 4–13 ventrally lightened by presence of greyish scales; A 3 exceeding length of A 4, shorter than A 4 + 5.</p><p>Mesosoma: Scutum and scutellum densely but somewhat irregularly punctate, punctures separated by &lt;0.5–1 puncture diameters, underlying surface shagreened, weakly shining (Fig. 71 C). Pronotum rounded. Mesepisternum irregularly roughened with network of rugosity, forming weak pseudopunctures, surface weakly shining. Propodeum with posterior and lateral faces covered in dense network of rugosity, surface dull. Propodeal triangle laterally indicated by fine carinae, internal surface with raised areolate rugae, interspaces shining. Mesosoma covered with long whitish hairs, none equalling length of scape. Legs dark, pubescence light brown, hind tarsal claws with small subapical tooth. Wings hyaline, stigma and venation orange-brown, nervulus postfurcal.</p><p>Metasoma: Terga dark, apical rims narrowly lightened hyaline brown (Fig. 71 D). Terga densely and uniformly punctate, punctures separated by 0.5–1 puncture diameters, punctures extending onto tergal margins; underlying surface polished and shining. Tergal discs with short scattered pale hairs, T 2–5 laterally with long loose hair fringes of pale hairs, not obscuring underlying surface. S 8 compact, robust, stem parallel-sided, disc rounded, ventral surface covered with dense fringe of light brown hairs (Fig. 72 A). Genital capsule compact, gonocoxal teeth short but strongly produced, gonostyli short, narrow, spatulate and parallel-sided, only slightly exceeding apical breadth of penis valves (Fig. 72 B). Penis valves narrow, medially occupying 1 / 2 space between gonostyli, slightly narrowing apically.</p><p>Diagnosis.</p><p>Andrena haladai can be recognised as a Lepidandrena due to the densely and uniformly punctate terga (Fig. 71 D), moderately long A 3 (Fig. 71 B), narrow and moderately indicated propodeal triangle, S 8 with produced disc ventrally covered with dense hairs (Fig. 72 A), and genital capsule compact with moderately produced gonocoxal teeth (Fig. 72 B). These are not particularly strong characters in isolation, but they are typical for Lepidandrena males, and the placement of A. haladai is supported by its DNA barcode (Fig. 16).</p><p>Andrena haladai has the external faces of the galea polished and shining, separating it from species around A. mocsaryi . Determination can be made by the combination of S 8 and the genital capsule. The disc of S 8 is short and thumb-shaped and is not noticeably wider or narrower than the stem (Fig. 72 A), whereas in A. paucisquama the stem of S 8 is very narrow before broadening into the disc which has its lateral margins concave and apical margin deeply emarginate (Fig. 72 C). The structure of S 8 is much more like A. pandellei (Fig. 72 E) and A. curvungula, but in these two species the gonostyli are apically broadened, shaped like the apex of an ice hockey stick and even slightly broadening up to their apices (Fig. 72 F). In contrast, A. haladai has the gonostyli apically narrow, only slightly broader than the breadth of the penis valves, definitively not broadening towards their apices (Fig. 72 B). This genital capsule form is much more similar to that seen in A. paucisquama (Fig. 72 D). In short, A. haladai displays a genital capsule similar to that of A. paucisquama, but a S 8 similar to that of A. pandellei; this combination of characters allows immediate recognition.</p><p>The female of A. haladai is currently unknown. There is a distinct possibility that it is very similar to the female of A. pandellei or A. paucisquama and is currently being misidentified as that. DNA barcoding of Lepidandrena females from sites with A. haladai present is necessary to convincingly locate the unknown female and study its morphology.</p><p>Remarks.</p><p>The males from Panormos were collected flying around a Campanula species ( Campanulaceae). Given the known specialisation of A. curvungula, A. pandellei, and A. paucisquama on Campanula (Westrich 1989; Schmid-Egger and Scheuchl 1997), it seems highly likely that A. haladai is also narrowly oligolectic on this plant genus.</p><p>Etymology.</p><p>Named after the Czech collector Jiří Halada who for many years has travelled the world collecting insects, many of which are deposited in the OÖLM collection and have served as the basis in whole or in part for (probably) hundreds of new species descriptions. His collections, along with those of other members of his family, have immeasurably increased our understanding of insect biodiversity. The name is in the genitive form.</p><p>Distribution.</p><p>Greece (Peloponnese, Central Greece).</p></div>	https://treatment.plazi.org/id/638E34D3F5725BC6AA7D8747F6070449	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
ACAD96DC757958C3B22CE9103C2B1468.text	ACAD96DC757958C3B22CE9103C2B1468.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Lepidandrena) Hedicke 1933	<div><p>Subgenus Lepidandrena Hedicke, 1933</p><p>Genetics.</p><p>Many sequences were available covering all species of the southern Balkans. Most species were well-resolved with high bootstrap support of&gt; 95 for species such as Andrena rufizona Imhoff, 1834, Andrena dorsalis Brullé, 1833, Andrena florivaga Eversmann, 1852, Andrena curvungula Thomson, 1870, and Andrena paucisquama Noskiewicz, 1924 . However, other species require discussion (Fig. 16).</p><p>Andrena pandellei Pérez, 1895 formed a monophyletic clade with bootstrap support of 88, but showed moderate average intraspecific genetic variation of 4.80 % (range 0.15–8.05 %). Three sequences from Bulgaria clustered together with bootstrap support of 100, with three sequences from Portugal, Germany, and Slovenia grouped together with bootstrap support of 90. Andrena pandellei was originally described from Algeria (lectotype: Theniet el Had) from a cedar forest in the Tell Atlas mountain range. The species is very rare in North Africa, and Warncke (1967 b: 215) described Andrena pandellei ssp. europaea Warncke, 1967 for the much more abundant, widespread, and commonly encountered European population. No authors have recognised population structuring within Europe, and hence a broad concept of A. pandellei is maintained for the time being.</p><p>Warncke adopted a broad species concept for Andrena gamskrucki Warncke, 1965 (locus typicus: Greece, Larisa), later describing Andrena gamskrucki ssp. eburnea Warncke, 1975 (locus typicus: Turkey, Ayvalik) and Andrena gamskrucki ssp. impasta Warncke, 1975 (locus typicus: Turkey, Birecik). Subsequent authors have treated these as distinct taxa (Dubitzky et al. 2010), and DNA barcodes support this distinction, at least for the two sampled taxa. Sequences of A. gamskrucki from mainland Greece were separated from sequences of A. impasta from Lebanon by an average genetic distance of 7.52 % (range 7.29–7.75 %). Unfortunately no barcodes are available for A. eburnea, but based on the morphological differences (see identification key), the conclusions of Dubitzky et al. (2010) can be followed. Andrena gamskrucki is treated here in a narrow sense as a species endemic to the southern Balkans. See species entry in the Updated Faunal List for discussion of A. eburnea on the East Aegean Islands. One additional morphologically-unrelated Lepidandrena sequence from southern Greece, NATBE 001-25, was placed as sister to A. gamskrucki and A. impasta, and was strongly isolated from all other Lepidandrena sequences by an average genetic distance of 12.18 % (range 10.97–13.23 %). It shows a genital and sternal morphology with aspects of A. pandellei and A. paucisquama, and is newly described below as Andrena haladai sp. nov.</p><p>The greatest problems come in the mocsaryi - group of species. These seem to be taxa strongly associated with Ornithogalum ( Asparagaceae) (e. g. Westrich 2010). It contains Andrena mocsaryi Schmiedeknecht, 1884 (types lost, but described from Hungary), Andrena baetica Wood, 2020 (locus typicus: Portugal, Portalegre), Andrena caprimulga Warncke, 1975 (locus typicus: Turkey, Horasan), and probably Andrena elisaria Gusenleitner, 1998 (locus typicus: Turkey, Hakkâri: Uludere) and Andrena statusa Gusenleitner, 1998 (locus typicus: Turkey, Hakkâri: Uludere). This group can be recognised due to the combination of the outer surface of the female galea which is strongly roughened and dull and the male sternum 8 which is flat and not divided into horizontal and vertical parts (as is the case in the florivaga - group sensu Schmid-Egger and Scheuchl 1997).</p><p>Five clades were observed, all with bootstrap support&gt; 93 (Fig. 16). One contained A. baetica (Spain and Portugal), one contained A. caprimulga from Turkey plus A. statusa from Lebanon, one contained A. mocsaryi from Austria and Greece (Peloponnese) ( mocsaryi # 1), one contained A. mocsaryi from Italy ( mocsaryi # 2; see also Mendicino et al. 2026), and one contained A. mocsaryi from the Peloponnese only ( mocsaryi # 3). The three clades of “ A. mocsaryi ” were all strongly separated. mocsaryi # 1 was separated from mocsaryi # 2 by an average genetic distance of 5.86 % (range 4.88–6.33 %) and from mocsaryi # 3 by an average genetic distance of 9.87 % (range 8.61–10.70 %). mocsaryi # 2 was separated from mocsaryi # 3 by an average genetic distance of 8.83 % (range 8.51–9.17 %). The situation is particularly confusing because the two mocsaryi lineages that can be found in sympatry in the Peloponnese (nos. # 1 and # 3) show the greatest observed separation of 10.70 % (sequences NATBE 030-25 and BGENL 1535-24). This genetic distance is substantial, and may indicate the presence of a cryptic species. Unfortunately, all of the barcoded Greek specimens of “ A. mocsaryi ” are female (all selected males failed), and in general it has only been possible to examine two male specimens from Greece compared to 36 female specimens. It seems that most collectors travel to Greece slightly too late to encounter males. If a cryptic species truly is present, then it is mocsaryi # 1 that represents the true A. mocsaryi since this clade contains a specimen from Austria, close to the terra typica of Hungary. More male material is required to understand what is happening in these different mitochondrial lineages of A. mocsaryi .</p><p>More broadly, the three mocsaryi lineages do not form a monophyletic clade. Andrena baetica was nested within, but was strongly separated from mocsaryi # 1 (5.66 %, range 4.88–6.43 %), mocsaryi # 2 (5.75 %, range 4.95–6.55 %), and mocsaryi # 3 (9.71 %, range 9.58–9.88 %). Morphologically, in principle there is no strong barrier to including A. baetica within a broad concept of A. mocsaryi as a geographically isolated population (differences are mostly related to female scutal hair length and structure and male clypeus colour), but the topology of the tree does not allow this given the strong genetic separation of A. caprimulga which is also nested within the broad A. mocsaryi sensu lato clade, in addition to the differences in the structure of the male S 8.</p><p>The inclusion of sequences of A. caprimulga is important to exclude the possibility that one of the “ lineages ” of A. mocsaryi in the Peloponnese is actually A. caprimulga . However, A. caprimulga sequences were strongly separated from the lineages present in the Peloponnese, namely mocsaryi # 1 (7.47 %, range 6.43–8.16 %) and mocsaryi # 3 (9.74 %, range 8.87–10.35 %). Morphologically, A. caprimulga is differentiated from A. mocsaryi by a small but distinct difference; it was not well-expressed by Warncke in his writing, but he did illustrate the key difference which is that the 8 th sternum is produced into three distinct points in A. caprimulga, and is rounded with a tiny apical point in A. mocsaryi (illustrated Warncke 1975: 95; see also Schmid-Egger and Scheuchl 1997: 156 for the character state of A. mocsaryi). None of the examined males from the southern Balkans have shown this distinctive A. caprimulga structure of the 8 th sternum. This difference has been slightly obscured because in the OÖLM collection, although Blank and Kraus (1994) indicated two female (holotype and paratype) and one male (paratype) specimens in the collection of Warncke during its accession, only the two female specimens are currently present. Moreover, the female paratype is stylopised and displays intersex features (e. g. a partially yellow-marked clypeus; Fig. 17).</p><p>The sequence of A. statusa from Lebanon is separated from the Turkish sequences by an average of 1.01 % (range 0.91–1.07 %), but the topology indicates that it is nested between them. Examination of a paratype male from Israel (directly; photographs also available on zobodat. at) indicate that the 8 th sternum shows the typical A. caprimulga shape, with three distinct points. This is consistent between Levantine specimens, newly examined and sequenced specimens from the Taurus mountains (RMNH), and Warncke’s original type series from Horasan (as illustrated by Warncke 1975). Consequently, A. statusa is formally synonymised under A. caprimulga below. Finally, Pisanty et al. (2018) already expressed doubts about the nominal differences described between A. elisaria and A. statusa based on co-occurrence of the taxa in Israel, conclusing that they may simply be dark and red-marked specimens of A. statusa (= A. caprimulga). Some male Lepidandrena specimens from south-eastern Turkey (OÖLM / TJWC) with extensive red maculations show a S 8 that is laterally rounded – these may represent A. elisaria, but the description combined with the holotype designated in the female sex is currently limiting. No taxonomic action for A. elisaria is taken at the present time, awaiting the generation of additional genetic sequences from Turkey.</p></div>	https://treatment.plazi.org/id/ACAD96DC757958C3B22CE9103C2B1468	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
2AA2EFF75FA75C10B3D8823452D4B83C.text	2AA2EFF75FA75C10B3D8823452D4B83C.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Leucandrena) Hedicke 1933	<div><p>Subgenus Leucandrena Hedicke, 1933</p><p>Notes.</p><p>Hedicke (1933: 215) described the Subgenus Leucandrena with the type species of Apis sericea Christ, 1791 nec. Forster, 1771 = Andrena barbilabris (Kirby, 1802) . This name has been used widely at a subgeneric level in the classification of Andrena since its original description. However, it has been discovered (Doug Yanega, pers. comm.) that Haeckel (1872: Band II: 172) created the name Leucandrena Haeckel, 1872 “ in cohors ” (sub-subgenus) within the genus Leucandra Haeckel, 1872 Subgenus Leucogypsa Haeckel, 1872 . This name is considered to be available under Article 10.4 (ICZN 1999) as even though it was created as a tertiary division of a genus, it was accompanied by the term “ cohors ” which can be considered equivalent to a term such as “ section ” or “ division ”.</p><p>There is no evidence that Leucandrena Haeckel has ever been used in any subsequent publications, and its recognition would result in subgeneric instability within Andrena, a challenging enough issue given the difficulties associated with applying and delineating Leucandrena Hedicke (see Pisanty et al. 2022 a). For this reason, the application of the Principle of Priority should be set aside and I here invoke reversal of precedence in accordance with Article 23.9. 1 of the Code (ICZN 1999). The senior homonym Leucandrena Haeckel has not been used as a valid name after 1899, and the junior homonym Leucandrena Hedicke has been used in at least 25 works published by at least 10 different authors in the last 50 years and encompassing a span of not less than 10 years (LaBerge 1987; Schmid-Egger and Scheuchl 1997; Xu and Tadauchi 1997, 2009; Gusenleitner and Schwarz 2002; Gusenleitner et al. 2005; Grünwaldt et al. 2005; Dubitzky 2006; Michener 2007; Osytshnjuk et al. 2008; Dubitzky et al. 2010; Ortiz-Sánchez 2011, 2022; Hazir et al. 2014; Scheuchl and Willner 2016; Álvarez Fidalgo et al. 2021; Astafurova et al. 2021; Bossert et al. 2022; Pisanty et al. 2022 a; Wood and Monfared 2022; Wood 2021, 2023 b, 2024 a, 2025 a; Ghisbain et al. 2023; Gautam et al. 2024).</p><p>Therefore, I establish the reversal of precedence according to Article 23.9. 1 giving the senior homonym Leucandrena Haeckel, 1872 the status of nomen oblitum and the junior homonym Leucandrena Hedicke, 1933 the status of nomen protectum.</p></div>	https://treatment.plazi.org/id/2AA2EFF75FA75C10B3D8823452D4B83C	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
516F6FC6FD6E5665A94DC00B6FA80C08.text	516F6FC6FD6E5665A94DC00B6FA80C08.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Limbandrena) toelgiana Friese 1921	<div><p>Andrena (Limbandrena) toelgiana Friese, 1921 sp. resurr.</p><p>Andrena tölgiana Friese in Fahringer &amp; Friese, 1921: 171, ♀ [Turkey, DEI, examined] (Fig. 19)</p><p>Material examined.</p><p>GREECE • 3 ♀; Lesbos, Agiassos; 658 m a. s. l.; 5 Jun. 2009; M. Hull leg.; WML ; TURKEY • 1 ♀; Amanusgeb [Amanus mountains], Jaribaschi; DEI (probable holotype) • 1 ♀; Akyaka, Yeşilova; 20 Jun. 2016; M. Kasparek leg.; OÖLM • 1 ♀; Ayvalik; 13 Apr. 1965; K. Warncke leg.; K. Warncke det.; OÖLM • 1 ♀; Diyarbakır; 15 Apr. 1972; Bytinski-Salz leg.; K. Warncke det.; OÖLM • 1 ♀; Side; 23 Apr. 1973; K. Warncke leg.; K. Warncke det.; OÖLM • 4 ♀; Side; 10 Apr. 1974; K. Warncke leg.; K. Warncke det.; OÖLM • 2 ♀; Antalya, Gençler; 341 m a. s. l.; 23 Apr. 2025; Ghisbain, Michez, Santerre leg.; TJWC / UMONS • 2 ♀; Çanakkale; 2024; L. Efil leg.; IUZC .</p><p>Notes.</p><p>A single specimen of A. toelgiana is preserved in the DEI collection; it is almost certainly the holotype, as Friese simply writes “ ♀ von Jaribaschi im Amanusgebirge, 7. Mai ”, whereas for other species he specifies the number of specimens, suggesting that only a single specimen was considered.</p><p>Wood (2023 b: 325) synonymised A. toelgiana under the widespread A. limbata Eversmann, 1852 due to the inconsistent and variable nature of the “ key ” discriminative character provided in the literature, namely the yellow-marked clypeus (Gusenleitner and Schwarz 2002). However, what was not clear from the literature is that this character is not at all important for identification; what is instead key is the short ocelloccipital distance which only slightly exceeds the diameter of a lateral ocellus (Fig. 19 D; 1–1.5 times), whereas in A. limbata this distance equals three times the diameter of a lateral ocellus. That the clypeus is yellow marked is incidental given the partial expression of this character in some specimens of A. limbata (see photographs in Wood 2023 b). In this context, A. toelgiana sp. resurr. is restored to species status, as the original synonymy was unjustified without the inspection of the original type material. The male remains unknown and undescribed.</p><p>In the southern Balkans, it is known only from the island of Lesbos (Grace 2010); this can be confirmed through new specimen examination. It has also been possible to examine specimens from Ayvalık and Çanakkale from the adjacent parts of nearby Turkey.</p><p>Distribution.</p><p>Greece (Lesbos) and Turkey descending into the Levant (Syria and northern Israel; Gusenleitner and Schwarz 2002) but ideally Levantine material should be located and validated.</p></div>	https://treatment.plazi.org/id/516F6FC6FD6E5665A94DC00B6FA80C08	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
654E5F4CE90E5604A9D7D0578E4A7894.text	654E5F4CE90E5604A9D7D0578E4A7894.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Melandrena) creticola (Strand 1915) Strand 1915	<div><p>Andrena (Melandrena) creticola Strand, 1915 stat. rev.</p><p>Andrena creticola Strand, 1915: 150, ♀ ♂ [Greece: Crete, DEI, examined]</p><p>Andrena creticola var. antennalis Strand, 1915: 151, ♂ [Greece: Crete, DEI, examined]</p><p>Andrena strigifrontalis Strand, 1921: 281, ♂ [Greece: Crete, DEI, examined]</p><p>Andrena pectoralis var. cretensis Friese, 1922: 211, ♀ [Greece: Crete, ZMHB, not examined]</p><p>Material examined.</p><p>GREECE • 1 ♂, 4 ♀; Canea [Chania] (Creta); 1 Mar. – 30 Jun. 1914; Paganetti leg.; DEI (syntypes of Andrena creticola) • 1 ♂; Canea [Chania] (Creta); 1 Mar. – 30 Jun. 1914; Paganetti leg.; DEI (holotype of Andrena creticola var. antennalis) • 1 ♂; Canea [Chania] (Creta); 1 Mar. – 30 Jun. 1914; Paganetti leg.; DEI (holotype of Andrena strigifrontalis) . See Suppl. material 2 for additional examined material.</p><p>Notes.</p><p>Warncke (1967 b: 251) placed A. creticola in combination with A. vachali Pérez, 1895 as A. vachali ssp. creticola . Wood (2023 b: 267) placed A. vachali in combination with A. discors, a taxon ranging from the Canary Islands into Iberia and across North Africa to the southern Levant. From the island of Crete, it was possible to examine 110 specimens (64 female, 46 male; see Suppl. material 2) collected between 1914–2023; the species appears common. The genetic results (Fig. 28) strongly support the recognition of A. creticola as distinct from A. discors, and more closely related to Andrena korleviciana Friese, 1887 (Adriatic region), Andrena vulcana Dours, 1873 (Iberia, north-west Africa, and the Canaries in a broad sense), and Andrena gravida (Europe and Turkey).</p><p>Morphologically, there are few differences from A. discors, leading to Warncke’s original combination (as A. vachali). In the female sex the tibial scopae are black (orange-red in A. discors), the scutal pubescence is entirely brown (variable, from foxy red to orange brown in A. discors but always with some short black hairs intermixed medially), and the pattern of punctures on the clypeus only weakly suggests the presence of longitudinal wrinkles (in A. discors, with a comparatively stronger pattern of longitudinal wrinkles on the clypeus, in North African specimens with almost shining interspaces). Males are structurally identical. Despite these very slight morphological differences, based on the known Andrena fauna of Crete, it does not have North African Andrena elements. With the revisions presented here, all of the species present are either very widely distributed pan-Mediterranean, European, or Cretan endemic species with the closest relative present in mainland Greece (e. g. Andrena pelagica Warncke, 1975 – Andrena rugothorace Warncke, 1965; Andrena incustodita Wood, 2024 – Andrena abbreviata Dours, 1873; Andrena lediveleci Wood, 2024 – Andrena sandanskia Warncke, 1973; Andrena testudo – Andrena brumanensis; Andrena cretica – Andrena hyemala). Based on this pattern, and the pattern suggested by the DNA barcodes, A. creticola stat. rev. is considered to be an isolated lineage (Fig. 29) descended from a European ancestor which due to colouration and the typical minimal morphological differentiation observed within members of the flavipes - group (here defined as corresponding to the former Subgenus Zonandrena Hedicke, 1933, see Pisanty et al. 2022 a) simply resembles A. discors . A similar pattern can be found in the Subgenus Taeniandrena Hedicke, 1933 (Praz et al. 2022) where structurally similar but differently coloured and partially geographically isolated species are strongly differentiated by DNA barcodes ( Andrena russula Lepeletier, 1841; Andrena croceiventris Morawitz, 1871; Andrena antonellae Praz &amp; Genoud, 2022).</p><p>Distribution.</p><p>Greece (Crete) (Fig. 29).</p></div>	https://treatment.plazi.org/id/654E5F4CE90E5604A9D7D0578E4A7894	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
8E5A4CDCC32F5A7081198FF01562D2C2.text	8E5A4CDCC32F5A7081198FF01562D2C2.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Melandrena) Perez 1890	<div><p>Subgenus Melandrena Pérez, 1890</p><p>Genetics.</p><p>The Subgenus Melandrena remains challenging to resolve using barcodes (e. g. nitida - limata-thoracica complex), and the widespread species Andrena morio Brullé, 1833 (originally described from Greece) is currently incompletely characterised with seemingly multiple mitochondrial lineages (Wood 2023 b). These two groups are not presented in the current phylogenetic analysis, as the subgenus would benefit from a revision utilising nuclear markers. Their statuses remain open questions, and for now the typical Warncke concepts (as modified by Wood 2023 b) are used here. One further reason for this is that the sequences of Andrena hungarica Friese, 1887 and Andrena quadrimaculata Friese, 1921 presented here are strongly intermixed with sequences of the nitida-limata-thoracica complex. Those of A. hungarica fall into BIN BOLD: AGU 5376 which contains sequences of A. nitida (Müller, 1776) and A. limata Smith, 1853, and those of A. quadrimaculata fall into BIN BOLD: AAE 1815 which contains sequences of A. nitida, A. limata, and A. thoracica (Kirby, 1802), as well as a likely mis-identification of A. assimilis Radoszkowski, 1876 . This group of species requires molecular revision using nuclear DNA to understand current levels of gene flow; their exclusion from the current barcode tree represents the challenging use of this marker in the Subgenus Melandrena, and a desire for higher-quality data, rather than an attempt to avoid dealing with this complex issue.</p><p>Concerning the taxa that are included here, many East Mediterranean taxa are represented by DNA barcodes for the first time (Fig. 28). Members of the subgenus showed sometimes strange patterns of very low genetic divergence, with species like Andrena flavipes Panzer, 1799 (Portugal to India; 3.85 %), Andrena albopunctata (Rossi, 1792) (Italy to Kyrgyzstan; 0.30 %), A. assimilis (Morocco to Kyrgyzstan; 0.95 %) separated by low percentages considering the geographic distances involved. Equally, some geographically close sequences showed moderate divergence, specifically between Levantine and Cypriot sequences of Andrena pyrozonata Friese, 1921 (average 3.16 %; range 2.74–3.93 %) and Andrena elmaria Gusenleitner, 1998 (average 4.46 %; range 4.41–4.56 %). Surprisingly, sequences of Andrena discors Erichson, 1841 from Portugal and Israel were very strongly separated from sequences from Crete ( ssp. creticola Strand, 1915) by an average genetic distance of 8.85 % (range 8.81–8.97 %). This resulted in a major topographical separation, placing creticola much closer to Andrena gravida Imhoff, 1832, Andrena korleviciana Friese, 1887, and Andrena vulcana Dours, 1873 . This case is discussed here.</p></div>	https://treatment.plazi.org/id/8E5A4CDCC32F5A7081198FF01562D2C2	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
75D741FDB4E9575B9B23C1EB8D152830.text	75D741FDB4E9575B9B23C1EB8D152830.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Melandrena) pyropygia Kriechbaumer 1873	<div><p>Andrena (Melandrena) pyropygia Kriechbaumer, 1873</p><p>Andrena pyropygia Kriechbaumer, 1873: 52, ♀ [“ Palestine ”, ZSM, lectotype by present designation] (Fig. 30)</p><p>Material examined.</p><p>ISRAEL • 1 ♀; “ Palästina ” [actually Jerusalem]; Dr. Roth leg.; ZSM (lectotype by present designation) .</p><p>Notes.</p><p>The locations of types described by Kriechbaumer are not always clear, so it is beneficial to designate the one remaining specimen in the ZSM collection as the lectotype (Fig. 30). The species was originally described from two females from “ Palästina ”, with Kriechbaumer noting in the publication that specimens were collected in Jerusalem. Friese labelled the specimen as “ morio v. pyropygia ”, indicating a level of scepticism as to the status of A. pyropygia, and indeed there are minimal structural differences between the two taxa. Warncke (1965 a, 1967 b) treated A. pyropygia as a valid species, and in the southern Balkans it can only be found in Greece. For now, A. pyropygia is maintained as a species in the sense of Warncke.</p><p>Distribution.</p><p>Greece, Crimea, Russia (European part), Turkey, Georgia, Armenia, Cyprus, Syria, Lebanon, Israel &amp; West Bank (Warncke 1974 a; Gusenleitner and Schwarz 2002; Osytshnjuk et al. 2008).</p></div>	https://treatment.plazi.org/id/75D741FDB4E9575B9B23C1EB8D152830	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
607B6FA37B2A5EBEB9F30199D2C813A5.text	607B6FA37B2A5EBEB9F30199D2C813A5.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Micrandrena) Ashmead 1899	<div><p>Subgenus Micrandrena Ashmead, 1899</p><p>Genetics.</p><p>Before discussing the genetics, it is necessary to highlight two taxonomic actions. The first is that Andrena corpana Warncke, 1965 (described from Larisa in Greece) is currently treated as a junior synonym of Andrena simontornyella Noskiewicz, 1939 (Gusenleitner and Schwarz 2002: 701). The second is that Pisanty et al. (2025) synonymised Andrena dargia Warncke, 1965 (described from Delphi in Greece) with Andrena minutula (Kirby, 1802); barcodes of “ Andrena dargia ” from the Peloponnese are presented here; these were also used in the analysis of Pisanty et al. (2025; Fig. 31). New barcodes of eastern Micrandrena are presented here, and the presence of Andrena oedicnema Warncke, 1975 in Greece is here discussed. There was some complexity seen with Andrena rugulosa Stöckhert, 1935 and Andrena illyrica Warncke, 1975, with A. illyrica intermixed between sequences of A. rugulosa . The rugulosa - group of species is taxonomically challenging with cryptic and only recently described Levantine species of high mountains (Pisanty et al. 2025), and the topology of this group is unstable; detailed analysis with additional samples is needed for clarity, and no action is taken here. Morphologically, the two taxa are separable, as indicated in the identification key.</p></div>	https://treatment.plazi.org/id/607B6FA37B2A5EBEB9F30199D2C813A5	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
DDEB963FDACD5F6EB7F72BB0DDAF4A0B.text	DDEB963FDACD5F6EB7F72BB0DDAF4A0B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Micrandrena) cedricola Wood 2020	<div><p>Andrena (Micrandrena) cedricola Wood, 2020</p><p>Andrena (incertae sedis) cedricola Wood in Wood et al. 2020: 10, ♀ ♂ [Lebanon, RBINS, examined]</p><p>Material examined.</p><p>GREECE • 1 ♀; Ad Klitoria [Ano Klitoria] - Kalavrita (Pelop.); 21 May 1987; H. Teunissen leg.; RMNH • 5 ♀; Erimanthos Mts (Peloponn); 28 May 1987; H. Teunissen leg.; RMNH • 2 ♀; Erimanthos Mts (Peloponn); 2 Jun. 1987; H. Teunissen leg.; RMNH • 2 ♀; Kalambaka; 17 May 1978; H. Teunissen leg.; RMNH ; TURKEY • 1 ♀; Abanoz (Rte Anamur), Kazanci; 11–1200 m a. s. l.; 11 May 1991; H. Teunissen leg.; RMNH • 1 ♀; Akseki; 1000 m a. s. l.; 5 May 1991; H. Teunissen leg.; RMNH • 6 ♀; Uzuncaburç, 30 km N of Silifke; 28 May 1996; Mi. Halada leg.; OÖLM / TJWC .</p><p>Notes.</p><p>Rasmont et al. (2017) reported Andrena oedicnema Warncke, 1975 (described from Turkey) for Europe based on the dot maps of Warncke (Gusenleitner and Schwarz 2002) which show a point in central Greece. No precise details for this specimen or specimens have ever been located. Examination of Teunissen material from the RMNH collection unexpectedly produced several specimens of Andrena cedricola, a species that is closely morphologically related to A. oedicnema . This identification is also supported by DNA barcode data (Fig. 31). The sequences from Greece were clustered with a sequence from northern Israel (Mount Hermon), separated by an average genetic distance of 1.32 % (range 0.91–2.13 %). Collectively, the Greek and Israeli sequences of A. cedricola were separated from four Israeli sequences of A. oedicnema by an average genetic distance of 10.37 % (range 8.42–14.42 %). Consequently, due to this high interspecific genetic distance, A. cedricola received bootstrap support of 98 and A. oedicnema received bootstrap support of 99.</p><p>After its original description from high-altitude sites in Lebanon and Syria, A. cedricola was recorded from Israel, Turkey, and northern Iraq (see Pisanty et al. 2022 b; Wood et al. 2024 for specimen details). Since then, additional Turkish specimens from central Taurus mountains have been discovered. The records from Greece come from the northern Peloponnese and central Greece, mountainous areas with a faunal affinity with the Taurus mountains (Fig. 32). In this context, the Warncke dot of “ Andrena oedicnema ” from central Greece is considered to belong to A. cedricola which replaces it on the Greek and European list. The precise specimen examined by Warncke may have been one of Teunissen’s specimens from Kalambaka, but this cannot currently be traced or demonstrated.</p><p>Distribution.</p><p>Greece *, Turkey, Lebanon, Syria, Israel, Iraq (Fig. 32; Wood et al. 2020, 2024; Pisanty et al. 2022 b).</p></div>	https://treatment.plazi.org/id/DDEB963FDACD5F6EB7F72BB0DDAF4A0B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
6E7425DF7A795F7D97C21B06A9D122CF.text	6E7425DF7A795F7D97C21B06A9D122CF.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Notandrena) albipila (Warncke 1967) Warncke 1967	<div><p>Andrena (Notandrena) albipila Warncke, 1967 stat. nov.</p><p>Andrena langadensis ssp. albipila Warncke, 1967 b: 237, ♀ ♂ [Spain, OÖLM, examined] (Figs 36, 37)</p><p>Material examined.</p><p>MOROCCO • 1 ♀; Agadir; 2–8 Apr. 1999; M. Hrodsky leg.; TJWC • 2 ♀; High Atlas, Aoulouz; 28 May 1995; J. Macek leg.; NMPC ; SPAIN • 1 ♀; Cádiz, Finca WWF (Trebujera); 19 Feb. 2022; Á. P. Gómez leg.; Á. P. Gómez Colln • 1 ♂; Caracoles (P. N. Doñana); 7 Apr. 2018; C. Molina leg.; EBDC • 1 ♀; Caracoles (P. N. Doñana); 19 Apr. 2021; C. Molina leg.; EBDC • 1 ♀; Convento de la Luz (P. N. Doñana); 8 Mar. 2017; C. Molina leg.; EBDC • 1 ♀; Granada, Cerro del Chupa, 1200 m [10 km NNW Otivar]; 4 Jun. 1986; W. Perraudin leg.; OÖLM • 1 ♀; Huelva, Isla Mayor; 17 Mar. 2012; I. Cross leg.; I. Cross Colln . • 1 ♂; Pozuelo de Ca. [Pozuelo de Calatrava]; La Fuente leg.; OÖLM (paratype of Andrena langadensis ssp. albipila) • 1 ♀; Villaverde; 6 Jun. 1909; J. M. Dusmet y Alonso leg.; OÖLM (holotype of Andrena langadensis ssp. albipila) .</p></div>	https://treatment.plazi.org/id/6E7425DF7A795F7D97C21B06A9D122CF	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
52925514608150E581CC41BE5C74D493.text	52925514608150E581CC41BE5C74D493.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Notandrena) clanga (Warncke 1965) Warncke 1965	<div><p>Andrena (Notandrena) clanga Warncke, 1965 stat. nov.</p><p>Andrena langadensis ssp. clanga Warncke, 1965 b: 259, ♀ ♂ [Turkey, OÖLM, examined] (Figs 38, 39)</p><p>Material examined.</p><p>CYPRUS • 1 ♀; North Cyprus, 15 km E Diakarpaz, Golden Sands; 10 Apr. 2007; C. Schmid-Egger leg.; CSE • 2 ♀; Sov. Base Area, Akrotiri, marsh; 1 m a. s. l.; 4 Apr. 2024; R. Santerre leg.; TJWC / UMONS ; GREECE • 1 ♀; Lesbos, Parakila; 14–17 May 1980; H. Teunissen leg.; RMNH, RMNH.INS.1716257 ; ISRAEL • 1 ♂; Akko [Acre], swamp; 1 Apr. 2012; A. Freidberg leg.; SMNHTAU ; JORDAN • 1 ♀; Irbid, Saham vill; 19 Apr. 2003; I. Pljushtch leg.; OÖLM ; SYRIA • 1 ♂; Ras al Basit; 19 May. 1995; K. Deneš leg.; OÖLM ; TURKEY • 5 ♂, 1 ♀; Adana; 25 Mar. 1967; K. Warncke leg.; OÖLM • 1 ♀; Ceyhan / Adana; 4 May 1964; K. Warncke leg.; OÖLM (paratypes of Andrena langadensis ssp. clanga) • 1 ♂; Dalyan, salty meadow [2 km S Dalyan]; 30 Apr. 2016; Barták &amp; Kubik leg.; OÖLM • 1 ♀; Hakkari, 5 km N Oramar [river]; 1450 m a. s. l.; 11 Jun. 1981; M. Kraus leg.; OÖLM • 1 ♀; Kadirli / Adana; 5 May 1964; K. Warncke leg.; OÖLM (holotype of Andrena langadensis ssp. clanga) • 8 ♀; Karatas [Karataş] / Adana; 29 Apr. 1964; K. Warncke leg.; OÖLM (paratypes of Andrena langadensis ssp. clanga) • 1 ♂; Milet / Aydin [ Milet Antik Kenti]; 22 Apr. 1981; K. Warncke leg.; OÖLM • 1 ♀; Tanin-Tanin-Pass; 2300–2600 m a. s. l.; 3 Jun. 1980; M. Schwarz leg.; OÖLM .</p></div>	https://treatment.plazi.org/id/52925514608150E581CC41BE5C74D493	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
1B6C93EB81165997A6AE3B206364EF96.text	1B6C93EB81165997A6AE3B206364EF96.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Notandrena) meridionalis Dalla Torre & Friese 1895	<div><p>Andrena (Notandrena) meridionalis Dalla Torre &amp; Friese, 1895 sp. resurr.</p><p>Andrena gravida Dours, 1872: 429, ♀ ♂ [Greece, MSGN, lectotype by present designation] nec. Imhoff, 1832 and Eversmann 1852 (Fig. 33)</p><p>Andrena meridionalis Dalla Torre &amp; Friese, 1895: 46, nom. nov. pro Andrena gravida</p><p>Andrena commutata Schulz, 1906: 241, unnecessary replacement name.</p><p>Andrena langadensis Warncke, 1965 a: 51, ♀ ♂ [Greece, OÖLM, examined] syn. nov. (Figs 34, 35)</p><p>Material examined.</p><p>ALBANIA • 1 ♀; Ksamil bij Butrinti; 24 May 2017; M. Jacobs leg.; KJCB ; GREECE • 1 ♀; “ Al. ” [= Algeria, considered mislabelled]; MSNG (lectotype) • 1 ♀; Corfu, Acharavi; 26 Apr. 1994; M. Koponen leg.; MZH • 1 ♂, 3 ♀; Corfu, Agios Spiridon; 7–17 May 1978; M. Viitasaari leg.; MZH • 1 ♂; Corfu, Sidari; 5 May 1978; H. Teunissen leg.; TJWC • 1 ♂; Crete, Kato Zakros; 11 Apr. 1988; M. Archer leg.; M. Archer Colln . • 1 ♂, 1 ♀; Crete: Lassithi, Goudouras; 11 Apr. 1992; S. P. M. Roberts leg.; SPMRC • 5 ♀; Hellas, Evros, Evros Deltagebied; 4 May 2004; J. v. d. Nieuwegiessen leg.; RMNH • 1 ♀; Lamia; 15 Apr. 1963; K. Warncke leg.; OÖLM (holotype of Andrena langadensis) • 1 ♂, 1 ♀; Langada b. Saloniki; 26 May 1945; Bodenstein leg.; OÖLM (paratypes of Andrena langadensis) • 3 ♀; Messinia, Agios Nikolaos; 26 May 1998; H. &amp; J. E. Wiering leg.; RMNH / TJWC • 2 ♀; Messinia, Gialova; 16 May 1998; H. &amp; J. E. Wiering leg.; RMNH • 1 ♀; Messinia, Petrochori; 17 May 1998; H. &amp; J. E. Wiering leg.; RMNH • 1 ♀; O. Creta, Gournia; 21 Apr. 1973; H. Teunissen leg.; OÖLM • 1 ♀; Peloponnese NE, N of Astros, Vernenon beach; 24 Mar. 2016; M. Snížek leg.; OÖLM • 1 ♀; Saloniki, Thessalien; 1–30 May 1942; Babiy leg.; OÖLM (paratype of Andrena langadensis) • 1 ♀; Syra [Syros]; OÖLM • 26 ♀; W, N of Amfilochia, Bukka [Mpoukka]; 1 Apr. 2016; M. Snížek leg.; OÖLM / TJWC .</p></div>	https://treatment.plazi.org/id/1B6C93EB81165997A6AE3B206364EF96	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
D7B8E22A917A5622BE24EF2BD8181F6B.text	D7B8E22A917A5622BE24EF2BD8181F6B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Notandrena) oblonga (Warncke 1967) Warncke 1967	<div><p>Andrena (Notandrena) oblonga Warncke, 1967 stat. nov.</p><p>Andrena langadensis ssp. oblonga Warncke, 1967 b: 236, ♀ ♂ [Libya, OÖLM, examined] (Figs 40, 41)</p><p>Material examined.</p><p>EGYPT • 1 ♀; Dekhela [Ad Dakhilah]; 20 Feb. 1917; Storey leg.; OÖLM (paratype of Andrena langadensis ssp. oblonga) ; LIBYA • 2 ♀; Cirenaica, Ras El Hilal [Ra’s Al Hilal]; 26 Apr. 1967; G. Dellacasa &amp; P. Maifredi leg.; MSNG / TJWC • 1 ♀; Cyrenaica, Bengasi; 14 Mar. 1922; G. C. Krüger leg.; OÖLM (holotype of Andrena langadensis ssp. oblonga) • 1 ♀; Cyrenaica, R. U. Agrario; 1 Jun. 1925; G. C. Krüger leg.; OÖLM (paratype of Andrena langadensis ssp. oblonga) • 1 ♀; Cyrenaica, R. U. Agrario; 17 Mar. 1929; G. C. Krüger leg.; OÖLM (paratype of Andrena langadensis ssp. oblonga) • 1 ♀; Cyrenaica, R. U. Agrario, Bu Teija; G. C. Krüger leg.; MSNG • 1 ♂, 1 ♀; Cyrenaika, Mars el Brega; 8–11 Feb. 1942; Kirchberg leg.; OÖLM (paratype of Andrena langadensis ssp. oblonga) • 1 ♀; Libia occ., u. Ramla; 1–31 Mar. 1938; G. C. Krüger leg.; MSNG .</p><p>Notes.</p><p>This group of species was treated by Warncke (1965 a, 1965 b, 1967 b) as a pan-Mediterranean taxon under the name A. langadensis . However, Warncke (1967 b) made several incorrect treatments and assessments of Notandrena species, leading to confusion and requiring taxonomic action to stabilise names and concepts (Gusenleitner and Schwarz 2002; Wood and Monfared 2022; Wood 2023 b). One stabilised species is Andrena (Notandrena) griseobalteata Dours, 1872 . Although the modern concept of this species is clear, there is one remaining issue which is that Warncke (1967 b: 288) treated A. meridionalis as a junior synonym (as A. erythrocnemis auctorum), and this remains the current placement of the taxon (Gusenleitner and Schwarz 2002: 323); this treatment requires dissection.</p><p>Andrena meridionalis is a replacement name for Andrena gravida Dours which is a junior primary homonym of A. gravida Imhoff, 1832 (and also A. gravida Eversmann, 1852). Dours described the species from “ Iles de l’archipel grec. Algérie ”. Whilst A. griseobalteata is common in Greece, it is not known from North Africa (Wood 2023 b); it is not clear if the original type series contained multiple taxa, or has been misinterpreted. A neotype was designated from southern France, and A. griseobalteata was considered to be a North Mediterranean species distributed from Spain to Turkey (Wood 2023 b). The identity of A. griseobalteata and A. meridionalis has always been unclear and confused because the collection of Dours was destroyed in the bombing of Amiens in the First World War (e. g. see Wood 2023 b); this is what necessitated the neotype designation for A. griseobalteata . However, what is less known is that some of Dours’ type material survived as specimens were traded with other hymenopterists such as Giovanni Gribodo (1846 – † 1924), and a number of syntypes for taxa described by Dours have been located in the MSNG collection (see Wood and Bossert 2025).</p><p>Examination of the Gribodo collection produced a female specimen labelled as “ Andrena gravida Typus Dours ” and nominally coming from Algeria (Fig. 33). Due to the short and broad head, pronotum with a humeral angle, and densely punctate terga, it can be placed into the Subgenus Notandrena, as is expected. However, the punctation of the scutum is dense, but not as dense as in A. griseobalteata (punctures in this species almost confluent), and instead conform to those seen in A. langadensis sensu lato . It is beneficial to reproduce the female description here:</p><p>“ ♀. Noire; antennes noires; mandibules ferrugineuses au bout. Tête arrondie, ponctuée; les poils cendré roux. Corselet assez fortement ponctué, les poils en dessus roux, plus pâles sur les côtés et en dessous. Abdomen elliptique très-élargi au niveau des 2 e et 3 e segments, nu, assez fortement ponctué. Bord inférueur des 2 e, 3 e, 4 e segments cilié de poils couchés, courtes, d’un blanc de neige. Les poils du 5 e segment et de l’anus sont roux doré. En dessous, le bord des segments est cilié de longs poils cendrés. Pattes noires. Tibias intermédiaires et postérieurs ferrugineux hérissées de poils blancs frisés, brosse cendré roux. Ailes assez limpides; point calleux testacé, noire au centre; côte, nervures brunes ”.</p><p>The description matches the specimen in key aspects, specifically the quite strongly punctured scutum, the hairs above red-brown (the probable meaning of roux in this case), those on the mesepisternum paler, metasoma with the discs of T 2–3 quite densely punctate, apical margins of T 2–4 with short hairbands of snow-white hair, and legs dark with the mid and hind tibiae ferruginous. Given these consistencies, and the known presence of Dours material in the MSNG collection (Wood and Bossert 2025), the specimen is hereby designated as the lectotype, and considered not to be conspecific with A. griseobalteata due to this sparser scutal punctation. As A. meridionalis is the oldest name in the langadensis - group of species, the challenge becomes 2) Where precisely was A. meridionalis described from? 2) Should all taxa in this group be lumped under the name A. meridionalis ? and 3) Are multiple good species involved?</p><p>Examination of the relevant type material (MSNG; OÖLM) shows that the MSNG lectotype of A. meridionalis is morphologically most similar to A. langadensis due to the relatively long tergal hairbands that are poorly defined and laterally exceed the length of the marginal areas before continuing onto the base of the following tergum (Figs 33 F, 34 D), the brownish pubescence of the head and (to a lesser extent) mesosoma (Figs 33 B, 33 C, 34 B, 34 C), and moderately dense tergal punctation and brown terminal fringe (Figs 33 F, 34 D). The problem is that A. langadensis s. str. is only known from the southern Balkan Peninsula, and the taxon found in the West Mediterranean ( ssp. albipila) has never been confidently recorded from Algeria, only Spain and Morocco (Gusenleitner and Schwarz 2002; Fig. 42).</p><p>There is therefore an incongruence between the reported collecting locality and the morphology presented by the lectotype of A. meridionalis . Given that Dours reported the species from “ Iles de l’archipel grec. Algérie ”, numerous Andrena species he described from “ Iles de l’archipel grec ” were based on material from Corfu (see Wood and Bossert 2025, section on Subgenus Ulandrena Warncke, 1968 below), and A. langadensis s. str. is known from Corfu based on material from multiple collectors (see Suppl. material 2), it is entirely plausible that this specimen is actually from Corfu (or at least Greece) and is simply mislabelled as coming from Algeria. Given this combined evidence, the position is taken that A. meridionalis was described from Greece as fixed by lectotype designation, and that it is senior to A. langadensis syn. nov.</p><p>The issue now becomes, what is the status of the taxa described as subspecies of A. langadensis? Examination of material from Lesbos has produced a female specimen which is referable to A. langadensis ssp. clanga . It shows the characters given by Warncke (1965 b), namely the bright orange antennae (Fig. 38 B; ventral surface), narrow tergal hairbands, and more strongly punctate terga (Fig. 38 D). When considering A. meridionalis and A. langadensis ssp. clanga, the density of the tergal punctation consistently differs, suggesting specific status for each taxon. When including A. langadensis ssp. oblonga (melanic, lacking tergal hairbands) and A. langadensis ssp. albipila (tergal hairbands consistently narrower and pubescence brighter, terga less densely punctate than A. langadensis ssp. clanga), as well as considering their non-overlapping geographic distributions (Fig. 42), the conclusion is that four distinct allopatric species are involved. Consequently, A. albipila stat. nov., A. clanga stat. nov., and A. oblonga stat. nov. are all elevated to species-status. An identification key for the females is presented below; males are currently insufficiently characterised, but can be identified through geographic provenance and co-occurrence with females.</p><p>Finally, it is necessary to discuss the species Andrena (Notandrena) stellaris Warncke, 1965 . This species was described from Aksaray in Anatolian Turkey, and the published distribution is Central Anatolia (Aksaray, Niğde, Ürgüp, Yeşilhisar, Sultanhani / Kayseri; Warncke 1974 a; Gusenleitner and Schwarz 2002). This species was reported from Lesbos by Grace (2010) without any further information or precise specimen details, and then in Rasmont et al. (2017) with personal communication from Erwin Scheuchl. For Anatolian Andrena species, unless they are very widespread (and hence are present in Anatolia as only a small part of their range), they do not typically descend to the Mediterranean coast. The presence of A. stellaris, a species with a small distribution range on the high Central Anatolian steppe, on Mediterranean Lesbos is considered implausible given the complete lack of published records or examined specimens from Mediterranean Turkey. Given the presence of A. clanga on Lesbos, it is strongly suspected that these listings of A. stellaris are in fact misidentifications of A. clanga . The two species are quite similar, but in the female sex A. clanga can be recognised due to the denser scutal punctures which are separated by 0.5–1 puncture diameters (in A. stellaris with scutal punctures separated by 1–3 puncture diameters) and the denser tergal punctures, on the disc of T 2 with punctures separated by &lt;0.5–1 puncture diameters (in A. stellaris with the punctures of the disc of T 2 separated by 1–2 puncture diameters). Since no actual evidence of A. stellaris on Lesbos has ever been published, it is here excluded from the Andrena fauna of the Balkans pending publication of precise information including diagnosis against A. clanga; Quod gratis asseritur, gratis negatur.</p><p>Distributions (Fig. 42).</p><p>Andrena meridionalis: Albania and Greece (including Crete); Andrena albipila: Spain and Morocco; Andrena clanga: Greece (East Aegean Islands), Turkey, Cyprus, Syria, Israel, and Jordan; Andrena oblonga: Libya and Egypt.</p></div>	https://treatment.plazi.org/id/D7B8E22A917A5622BE24EF2BD8181F6B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
8A10C730121754129B5428B36E00EE7B.text	8A10C730121754129B5428B36E00EE7B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Orandrena) Warncke 1968	<div><p>Subgenus Orandrena Warncke, 1968</p><p>Genetics.</p><p>The situation for morphological identification in this subgenus is challenging. Warncke considered a widespread species that was originally described from Uzbekistan, Andrena oralis Morawitz, 1876 (see Astafurova et al. 2022: 148 for illustration of lectotype), as present across the Eurasian steppe to the Pannonian basin and also descending to Greece and Turkey (Gusenleitner and Schwarz 2002). He determined specimens from Delphi, presenting them in Warncke (1965 a: 40). He then later described Andrena gallinula Warncke, 1975 from Turkey (Warncke 1975: 33), very briefly diagnosing it against A. oralis . The two species are extremely similar morphologically, and Warncke did not give the key characters precisely. Specifically, the genital capsule of the male has the apical part of the gonostyli short and broad in A. gallinula (Fig. 43 E) whereas this is comparatively long and narrow in A. oralis (Fig. 43 F). In the female sex, I have only been able to find the ocelloccipital distance to have discriminative power some of the time, approximately 1.5–2 times the diameter of a lateral ocellus in A. oralis (Figs 44 B, 44 D) and being equal to the diameter of a lateral ocellus in A. gallinula (Fig. 44 F, H).</p><p>Examination of Andrena specimens from Greece has produced specimens showing both genital types (see Suppl. material 2 for full specimen details). In fact, A. gallinula is much more commonly encountered in Greece than A. oralis, with 55 and 11 examined specimens, respectively (Fig. 45). Most examined specimens were collected by H. Teunissen, and dissected males clearly show the short and broad apex of the gonostyli of A. gallinula . However, many ambiguous females can be found in these Greek collections which seem to show intermediate ocelloccipital distances and hence are difficult to place.</p><p>Generation of DNA barcodes does not offer clarity (Fig. 46). Five sequences are available, one from Austria (ABEE 105-17) which must be A. oralis, and four from Greece. BGENL 2355-24 is a dissected male from Karkalou in the Peloponnese showing a typical A. oralis genital capsule. The remaining three sequences are unfortunately difficult to determine female specimens, two of which (BGENL 2352-24 and BGENL 2353-24) were determined as A. oralis by K. Warncke and W. Grünwaldt, and one of which (BGENL 2354-24) was determined as A. gallinula by myself. However, the topology of the tree does not clearly show two taxa (Fig. 46). The one possible A. gallinula female sequence is nested with A. oralis sequences, and the one male sequence of A. oralis is placed at the base of the other four sequences. The highest average genetic distance of 4.40 % (range 4.06–5.41 %) was actually seen between the Austrian sequence and all Greek sequences, with the highest individual difference of 5.41 % between the Austrian sequence and the A. oralis male from Karkalou.</p><p>This pattern is challenging to understand given the clear presence of two genital forms in southern Greece. To illustrate this point, a male A. oralis was captured on 3 Apr. 2024 near the village of Georgitsi in the central Peloponnese (Fig. 47 A, C), and a male A. gallinula was captured on 9 Apr. 2024 on Mount Parnassus (Fig. 47 B, D). The genital capsules clearly and unambiguously differ. It is currently not possible to explain this incongruence between DNA barcodes and morphology. It may be that A. oralis and A. gallinula were historically separated, and have now come back into contact and are hybridising in southern Greece. It may be that incomplete lineage sorting has occurred. It may be that all the sequences presented here actually belong to A. oralis, and A. gallinula remains unsampled. In the absence of additional data which would allow for detailed investigation of these problems, a morphological species concept is followed which distinguishes the two species on the basis of the male genital capsule, and A. gallinula is considered to be present in southern Greece. The female ocelloccipital distance is tentatively given as an identification character, but it is clear that additional work is needed to full characterise Orandrena species in the East Mediterranean.</p></div>	https://treatment.plazi.org/id/8A10C730121754129B5428B36E00EE7B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
347BC511E8C65B01BE5241A861F1EA9B.text	347BC511E8C65B01BE5241A861F1EA9B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Poecilandrena) Hedicke 1933	<div><p>Subgenus Poecilandrena Hedicke, 1933</p><p>Genetics.</p><p>The Subgenus Poecilandrena has its greatest species richness in the East Mediterranean (Pisanty et al. 2018), with only two species known from the Iberian Peninsula (Wood 2023 b). DNA barcodes (Fig. 48) were recently produced for many East Mediterranean species for the first time (Pisanty et al. 2022 b; present work), allowing species concepts to be validated. For the species in the subgenus showing partially metallic integument, the relatively recently described Greek endemics Andrena neovirida Grünwaldt, 2005 and Andrena standfussorum Scheuchl, 2010 were strongly separated from the widespread Andrena viridescens Viereck, 1916 (Fig. 48). The results also further support the species status of Andrena veronicae Pisanty &amp; Wood, 2022 as distinct from the more northerly Andrena semirubra Morawitz, 1875 (see Pisanty et al. 2022 b for discussion), as sequences of A. veronicae from Israel were separated by an average genetic distance of 13.15 % (range 12.92–13.37 %) from A. semirubra sequences from Turkey and Armenia.</p><p>However, the results in the non-metallic clade of Poecilandrena were more complex. Although Andrena crassana Warncke, 1965 showed minimal differentiation of the “ subspecies ” inka Warncke, 1969, with sequences from the island of Rhodes intermediate between mainland Greek and Levantine sequences (implying separation by distance), three red-marked taxa showed a confusing picture. Specifically, these were Andrena labiata Fabricius, 1781, Andrena sphecodimorpha Hedicke, 1942, and Andrena hybrida Warncke, 1975 . Andrena labiata is very widespread, occurring from Iberia to Central Asia, whereas A. sphecodimorpha is restricted to the Balkans and Crimean Peninsula. Andrena hybrida is possibly a species complex, with two described subspecies; ssp. uralskia Warncke, 1975 from western Kazakhstan, southern European Russia, Ukraine, and Crimea; ssp. tauriensis Warncke, 1975 from southern Turkey and newly reported from western Turkey and the island of Lesbos (Fig. 49; see species entry below); and hybrida s. str. from Turkey, Armenia *, Iran, and Iraq (Gusenleitner and Schwarz 2002; Wood et al. 2024; see Suppl. material 2).</p><p>In the phylogenetic tree, these three main taxa were intermixed. Two clades of A. labiata were seen (Fig. 48), the first ( labiata # 1) with sequences from Portugal, Spain (including the Sierra Nevada), Italy, the United Kingdom, and Bulgaria, and the second ( labiata # 2) with sequences from France, Belgium, Denmark, Germany, Italy, Greece, Crimea, and Kyrgyzstan. The clades were separated by an average genetic distance of 4.53 % (range 3.43–5.47 %) and received bootstrap support of 87 and 97, respectively. labiata # 1 showed within-clade variation of 1.79 % (range 0.15–2.69 %) and labiata # 2 showed within clade variation of 1.04 % (range 0.17–2.07 %).</p><p>The two clades of A. labiata did not form a monophyletic clade, as labiata # 1 was closer to a clade of A. sphecodimorpha ( sphecodimorpha # 2) containing sequences from Greece (Central Greece and East Macedonia). A second clade ( sphecodimorpha # 1) contained sequences from the Peloponnese. sphecodimorpha # 2 showed within clade variation of 1.56 % (range 0.00–2.49 %) and was separated from sphecodimorpha # 1 by an average genetic distance of 6.72 % (range 6.01–7.62 %). Between these clades was the single available sequence of A. hybrida . However, it is important to stress that the topology of this part of the tree is very weakly supported; all of the key nodes have &lt;75 bootstrap support, and the branches are very short. The tree is therefore fragile, and subject to structural changes if sequences are added or removed.</p><p>Regarding A. labiata, understanding this pattern is not possible with only evidence from DNA barcodes. It may be that this is a result of incomplete lineage sorting, but it is also possible that speciation started to begin in geographic isolation, leading to the accumulation of differences in the COI gene. Populations may then have come back into contact, re-establishing gene flow, halting speciation, and leading to distinct mitochondrial lineages within a single biological species (e. g. Kearns et al. 2023 for an example in ravens). Based on the morphology of the male genital capsule, there is no reason at the present time to believe that these two mitochondrial lineages represent two different biological species in Europe, and the concept of A. labiata has been more-or-less stable for over 200 years (albeit with a few clarifications needed regarding the appropriate name for this species; Gusenleitner and Schwarz 2002). No taxonomic action is therefore taken here, pending the generation of additional barcodes and ideally nuclear DNA sequences.</p><p>The holotype of A. sphecodimorpha (from Sofia, 17 Apr. 1915, leg. A. K. Drenowski) is lost, as are all Hedicke types which were placed in his personal collection (Wood 2025 a). It is currently considered premature to designate a neotype, both due to the two observed clades of A. sphecodimorpha, and also because genetic data from Bulgaria (the terra typica) are lacking. Based on the geographic distribution of the sequences presented here, A. sphecodimorpha should fall into sphecodimorpha # 2, as this contains sequences from Eastern Macedonia in northern Greece which are closer to the terra typica (Fig. 49).</p></div>	https://treatment.plazi.org/id/347BC511E8C65B01BE5241A861F1EA9B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
032F55EDB4E7248CB4D2BF354F39F0AA.text	032F55EDB4E7248CB4D2BF354F39F0AA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Stenomelissa) Hirashima & LaBerge 1965	<div><p>Subgenus Stenomelissa Hirashima &amp; LaBerge, 1965</p><p>Andrena lonicera Warncke, 1973 and Andrena lonicerae Tadauchi &amp; Hirashima, 1988</p><p>The Subgenus Stenomelissa is traditionally considered to contain three species restricted to East Asia (the Russian Far East, China, the Korean Peninsula, and Japan) (Tadauchi and Hirashima 1988; Gusenleitner and Schwarz 2002; Xu and Tadauchi 2008). However, the species Andrena lonicera was originally described from North Macedonia in the Subgenus Hoplandrena Pérez, 1890 . It was listed as part of the Subgenus Stenomelissa by Grace (2010: 71), placed there in the framework of the STEP project (Rasmont et al. 2013), and is currently listed there by Ascher and Pickering (2025). These treatments probably derive from study of the type series by Erwin Scheuchl who observed striking similarities between the male of A. lonicera and the East Palaearctic Andrena (Stenomelissa) halictoides Smith, 1869; this information was communicated to Fritz Gusenleitner and Andrew Grace (E. Scheuchl, pers. comm.).</p><p>Andrena lonicera was described based on six females and one male collected by Herman Teunissen from an altitude of 1800 m on the Galičica Planina mountain in south-western North Macedonia. Warncke (1973: 35) described this species as an addendum to his main article after receiving specimens from Teunissen, and indicated that the specimens were collected from Lonicera ( Caprifoliaceae) in difficult conditions (unter Schwierigkeiten) at the edge of the tree line. The species was therefore named after this plant genus. Of the original seven specimens, two female paratypes have been located in the RMNH collection, but in the OÖLM collection only the female holotype (Fig. 50) and one female paratype could be found. As for A. caprimulga, unfortunately the single male paratype appears to be missing from the OÖLM collection, despite being indicated as present by Blank and Kraus (1994). As to the remaining two female paratypes, their location is unclear. No further specimens of A. lonicera have yet been encountered in field or museum collections.</p><p>Tadauchi and Hirashima (1988: 72) described Andrena lonicerae from eastern and southern Japan (type material Kyushu University, Fukuoka, Japan) where it is endemic (Xu and Tadauchi 2008). It predominantly visits Lonicera gracilipes Miq., on which it appears to be specialised (Shimizu et al. 2014). The name therefore also derives from the genus Lonicera . Although the names are very similar, they are both valid, as they differ by a single letter (ICZN Article 57.6), and A. lonicerae does not meet the exceptions to this rule laid out under Article 58.</p><p>It was possible to generate barcodes from the two female paratype specimens of A. lonicera in the RMNH collection (Fig. 48). In the barcode tree, A. lonicera clustered most closely to A. lonicerae (sequences from Genbank), the sequences separated by an average genetic distance of 5.68 % (range 5.58–5.78 %). Together, A. lonicera + A. lonicerae placed closer to sequences from the subgenera Warnckandrena Pisanty, 2024 and Uncandrena Pisanty, 2024 which are the most basal Andrena subgenera known (Pisanty et al. 2022 a; Pisanty and Wood 2024). Whilst ultra-conserved element genetic data for members of the Subgenus Stenomelissa are lacking, and ultimate confirmation or rejection of the phylogenetic placement of Stenomelissa must wait, the placement close to Warnckandrena + Uncandrena combined with the morphology of A. lonicera is highly suggestive. Importantly, A. lonicera has the propodeum nearly completely declivous, without defined horizontal and vertical parts. This is most strongly observed in the genus Cubiandrena Warncke, 1968, and then most strongly observed in the subgenera Warnckandrena and Uncandrena (Pisanty and Wood 2024).</p><p>Andrena lonicera is therefore considered to be part of the Subgenus Stenomelissa, and to represent an extremely isolated member. This is slightly reminiscent of the situation in Subgenus Warnckandrena, with the most easterly member found in northern Iran, with a seemingly closely related but isolated species found on the Tibetan Plateau in central China ( Andrena remota Wood, 2024; Pisanty and Wood 2024). The use of Lonicera by A. lonicera and A. lonicerae is also highly unusual for Andrena species, and this seemingly linked niche use requires further study if populations of A. lonicera can be located in the southern Balkans in the present day.</p></div>	https://treatment.plazi.org/id/032F55EDB4E7248CB4D2BF354F39F0AA	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
813E236B6513403649AF524E38D91121.text	813E236B6513403649AF524E38D91121.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Taeniandrena) Hedicke 1933	<div><p>Subgenus Taeniandrena Hedicke, 1933</p><p>This subgenus has received much taxonomic attention in the past few years (e. g. Wood et al. 2021; Praz et al. 2022; Wood 2022, 2023 b, 2023 c). New samples from the southern Balkans and the Caucasus (Fig. 51) do not greatly change our understanding relative to the situation presented in Praz et al. (2022), but they do further support those conclusions. The main findings are additional sequences of Andrena afzeliella (Kirby, 1802) (from Bulgaria), Andrena hova Warncke, 1975 (from Armenia), Andrena intermedia Thomson, 1870 (from Bulgaria and Greece), Andrena leucopsis Warncke, 1967 (from Greece), Andrena ovata Schenck, 1853 (from Bulgaria), Andrena producta Warncke, 1973 (from Bulgaria and Greece), Andrena solitaria Warncke, 1975 (from Armenia), and Andrena taedium Wood, 2023 (from Greece and Armenia). Both A. leucopsis and A. taedium (Fig. 52) are confirmed to be species distributed from the Balkan Peninsula to western Iran.</p><p>The greatest area of uncertainty surrounds Andrena russula Lepeletier, 1841 and Andrena phoenicura Warncke, 1975 . Andrena russula is widely distributed across the West Palaearctic (Praz et al. 2022), with sequences available from Finland, Cyprus, and Israel in the east (Fig. 51). However, two sequences from northern Greece (NATBE 158-25) and southern Bulgaria (NATBE 163-25) were very widely separated from the main A. russula clade by an average genetic distance of 5.76 % (range 4.97–6.53 %). Andrena russula remains very difficult to understand, as away from the main central clade of sequences (the clade presented in Praz et al. 2022 and here; Fig. 51), multiple “ cryptic ” lineages can be seen in south-western Iberia, Morocco, and now the southern Balkans (Praz et al. 2022; Wood 2023 b, 2023 c; Wood et al. 2024 a). The Greek + Bulgarian sequences do not match either of the West Mediterranean “ cryptic ” lineages.</p><p>No actions are taken here due to the number of names of incompletely understood Taeniandrena species that are available in Turkey. Warncke described Andrena phoenicura from Turkey, and this species was listed from Greece (Kuhlmann et al. 2014) and Bulgaria (Wood 2021), specifically from specimens from Sandanski and Slanĉev Brjag (Sunny Beach). Re-examination of some of these specimens published in 2021 show that they are actually Andrena taedium, a species described in 2023 (Wood 2023 a). It has not been possible to locate all of the specimens deposited in the OÖLM (Kocourek collection) that were presented in Wood (2021), as the material (including all Andrena specimens determined by myself between 2019–2024) is in the process of being curated into the main collection. However, the specimens retained in my personal collection conform to A. taedium . Given the new records of A. taedium, including many from Greece, the revised global distribution is presented in Fig. 52). It was not possible to directly examine any specimens of A. taedium from Georgia, but on BOLD the sequence GBAH 18598-19 which was originally identified as Andrena wilkella (Kirby, 1802) is clearly nested with A. taedium (Fig. 51), and is considered to represent the first record of the species from this country.</p><p>The type material of A. phoenicura (OÖLM) has now been carefully studied (Fig. 53), and it is a taxon close to A. russula . Barcodes were generated from possible A. russula females from Armenia, but these matched males of Andrena solitaria, a species described only in the male sex which has a highly distinctive genital capsule (Warncke 1975). The holotype of A. phoenicura is a male, and so it can be confidently concluded that it is not conspecific with A. solitaria, but the characters allowing determination of the females of these two species are not clear. Currently, no barcodes from the southern Balkans match sequences of A. solitaria (Fig. 51). It is considered unlikely that the two “ A. russula ” sequences from Greece and Bulgaria are actually A. phoenicura, as the specimen from Greece (NATBE 158-25) is a male which has a genital capsule which is slimmer than in A. phoenicura and lacks the more strongly flattened and spatulate apices of the gonostyli. It has A 3 slightly shorter than A 4, whereas in some A. russula specimens A 3 exceeds the length of A 4, but this has already been noted to be a “ variable ” character (Praz et al. 2022) and is difficult to characterise even in European specimens. At the present time, A. phoenicura is considered unproven from the southern Balkans and must be treated as a species inquirenda.</p></div>	https://treatment.plazi.org/id/813E236B6513403649AF524E38D91121	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
AB21C6C902325EBD9465B818AD6DD709.text	AB21C6C902325EBD9465B818AD6DD709.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Taeniandrena) taedium Wood 2023	<div><p>Andrena (Taeniandrena) taedium Wood, 2023</p><p>Material examined.</p><p>ALBANIA • 1 ♀; Kukës, W of Perollaj; 26 May 2024; W. Vertommen leg.; WVC ; ARMENIA • 1 ♀; Byurakan; 1309 m a. s. l.; 9 May 2023; V. Leclercq leg.; VLC ; BULGARIA • 1 ♀; N Panitchishte vill.; 1237 m a. s. l.; 27 Jun. 2016; M. Naumova leg.; IBER • 2 ♂, 1 ♀; Sandanski; 1–30 Jun. 1969; M. Kocourek leg.; TJWC ; CROATIA • 1 ♀; Split, Gornji Muc, malaise trap; 500 m a. s. l.; 14 May – 8 Jun. 2014; B. Kokan leg.; OÖLM ; IRAQ • 1 ♀; Duhok gov., 5 km E of Ashewa, Mt. Gara; 1750–1950 m a. s. l.; 30 May – 12 Jun. 2024; D. Baiocchi leg.; MSVI ; NORTH MACEDONIA • 1 ♂; Polog reg., Sirichino, quarry; 400 m a. s. l.; 12–17 Apr. 2021; V. Bureš leg.; JSPC • 2 ♀; Vardar reg., Papeliste, steppe; 130 m a. s. l.; 6–11 Apr. 2021; V. Bureš leg.; JSPC .</p><p>Notes.</p><p>See Suppl. material 2 for full records, including from Greece.</p><p>Global distribution.</p><p>Croatia *, Albania *, North Macedonia *, Greece, Bulgaria *, Turkey, Lebanon, Georgia *, Armenia *, Iraq *, Iran (Wood 2023 a) (Fig. 52).</p></div>	https://treatment.plazi.org/id/AB21C6C902325EBD9465B818AD6DD709	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
8CDB312BDAB753DDB2E7FDEB6A75C37B.text	8CDB312BDAB753DDB2E7FDEB6A75C37B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Truncandrena) tscheki Morawitz 1872	<div><p>Andrena (Truncandrena) tscheki Morawitz, 1872</p><p>Andrena nigrifrons Smith, 1853: 106, ♀ ♂ [Greece, OUMNH, lectotype by present designation] nec. Eversmann, 1852 (Fig. 56)</p><p>Andrena tscheki Morawitz, 1872: 366, ♀ [Austria, ZISP, examined by photograph]</p><p>Andrena tscheki spp. tritica Warncke, 1965 a: 36, ♀ ♂ [Greece, OÖLM, examined] (Fig. 57)</p><p>Material examined.</p><p>GREECE • 1 ♂, 1 ♀; 446. Alb nia; OUMNH (female lectotype of Andrena nigrifrons Smith by present designation, male paralectotype) .</p><p>Notes.</p><p>Gusenleitner and Schwarz (2002: 783) gave the NHMUK as the repository for the type of A. nigrifrons Smith, but the species was stated as being described from the collection of William Wilson Saunders, which was acquired by the OUMNH in 1873 (Anon 1879). A female and male labelled as “ 446. ♀. Alb nia ” and “ 446. ♂. Alb nia ” were collected by Sir Sidney Smith Saunders, and are syntypes. Sidney S. Saunders (1809–1884) was cousin of W. W. Saunders, and was appointed as British Consul in “ Albania ” between 1835–1859.</p><p>It is necessary to discuss what “ Albania ” actually constituted at this time. After the creation of the First Hellenic Republic in 1822, the Ottoman Empire retained control of what is now northern Greece, Albania, North Macedonia, and Bulgaria. Importantly, the Ottoman Empire retained control of the town of Preveza, which today is in modern Greece. Under the Ottomans, this town was the centre of the Sanjak of Preveza. This covered the period during which S. S. Saunders was appointed consul at “ Prevesa ” [= Preveza] (Wilson 2011), i. e. within Ottoman Albania. Greece did not formally acquire the territories including Preveza until the First Balkan War and the 1913 treaty of London. Consequently, most of the material collected by S. S. Saunders in “ Albania ” is likely to be from within the borders of modern Greece, and following Baker (1993) it is treated as such.</p><p>Baker (1993: 132) located this material and stated that he labelled the female as lectotype, but this is not a valid designation as Baker (1993) is an unpublished thesis and hence fails Articles 8 and 9 (ICZN 1999). Moreover, Baker did not label the specimens. The female is now finally designated as the lectotype. Whilst the name is not available due to homonymy, Smith’s types morphologically conform to what Warncke called A. tscheki spp. tritica; the apical margin of T 4 has a complete hairband, whereas in the nominate subspecies T 4 is entirely dark-haired.</p><p>Based on the analysis of the COI fragment, three subclades were found within a broad A. tscheki which do not agree with the observed morphology (Fig. 54). Specimens from Greece were strongly separated from both German and Levantine sequences, despite morphologically being closer to Levantine specimens. Purely based on the morphology (Fig. 58), A. tscheki s. str. can be found across Central Europe to Vitosha mountain by Sofia in Bulgaria, with specimens from lower altitudes confirming to A. tscheki spp. tritica across the southern Balkans, across Turkey (see Warncke 1974 a: 92 for additional localities), and to the Levant. Additionally, and there are two female specimens from southern Russia (Dagestan) in the ZSM which were determined as “ aff tscheki ” by Osytshnjuk (see Suppl. material 2) which do seem to meet the morphologically criteria for A. tscheki s. str. In short, there does not appear to be a clear way to split A. tscheki; if split, the Levantine taxon would have to be described, and structurally, there do not appear to be any differences in the genital capsule or integument across the range from eastern France to the Levant. As such, the broad A. tscheki species concept of Warncke is maintained in this work.</p></div>	https://treatment.plazi.org/id/8CDB312BDAB753DDB2E7FDEB6A75C37B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
BEDE3D9319735378B71D552E1749A88A.text	BEDE3D9319735378B71D552E1749A88A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Truncandrena) Warncke 1968	<div><p>Subgenus Truncandrena Warncke, 1968</p><p>Genetics.</p><p>The Subgenus Truncandrena is taxonomically challenging, with many “ subspecific ” names available due to Warncke’s broad species concepts and a high level of “ variation ” comprising different colour patterns (in both the integument and pubescence). This has led to the elevation of many of these subspecific concepts to full species status (e. g. Gusenleitner and Schwarz 2002; Pisanty et al. 2022 b; Wood 2023 b, 2024 a). In this context, many new DNA barcodes were available for the East Mediterranean fauna. For many taxa, they offered good resolution; although females are challenging to separate morphologically, Andrena caneae Strand, 1915 and Andrena serraticornis Warncke, 1965 were strongly separated genetically (Fig. 54).</p><p>Three major issues were detected in this subgenus. Firstly, Andrena tscheki Morawitz, 1872 sensu lato formed three distinct subclades, with sequences from 1) Greece, 2) Germany, and 3) Israel and Lebanon. Greek specimens were separated from German specimens by an average genetic distance of 9.63 % (range 9.57–9.73 %), and from Levantine sequences by an average of 11.21 % (range 11.09–11.25 %). German and Levantine sequences were separated by an average of 7.38 % (range 7.29–7.45 %). This issue is discussed below, along with morphological recognition of the taxa.</p><p>Secondly, Andrena schmiedeknechti Magretti, 1883 (described from Italy) formed two lineages, with sequences from Italy (Sicily) strongly separated from sequences from southern Greece by an average genetic distance of 6.28 % (range 5.87–7.12 %). Although smaller in absolute terms than the distances seen in A. tscheki sensu lato, this led to a topology where Italian A. schmiedeknechti were placed closer to members of the rufomaculata - group, and Greek A. schmiedeknechti were placed closer to Andrena urfanella Scheuchl &amp; Hazir, 2012 . The name Andrena schmiedeknechti ssp. flavopilis Warncke, 1965 is available for Greek populations, but additional study is required. The topology of the tree separating Italian and Greek sequences is unstable, with bootstrap support grouping Italian A. schmiedeknechti + rufomaculata - group of only 37. The addition of more sequences from this group (including A. schmiedeknechti ssp. nigrorum Warncke, 1967 from North Africa) could result in a monophyletic clade, as in A. tscheki sensu lato . There also do not appear to be structural differences between the genital capsules of A. schmiedeknechti sensu lato seen in Italy and Greece, and the only difference is in the colouration of the hair (Warncke 1965 a). As such, no action is taken here, and the broad species concept of Warncke is maintained.</p><p>The third issue surrounds the rufomaculata - group which contains the taxa Andrena rufomaculata Friese, 1921, Andrena paramythensis Mavromoustakis, 1957, and Andrena optata Warncke, 1975 . Warncke (1974 a, 1975) treated this as a broad taxon A. rufomaculata, with A. rufomaculata s. str. found in southern Turkey and the Levant (locus typicus: Turkey, Airan [an Armenian village, modern name not currently traceable] in the Taurus Mountains), A. rufomaculata ssp. paramythensis found in Cyprus (locus typicus: Cyprus, Paramytha), and A. rufomaculata ssp. optata found from central Anatolia across the Mediterranean coast to Greece, Albania, the Dalmatian coast, southern Hungary, and Crimea (locus typicus: Greece, Lamia). Although Warncke and Gusenleitner and Schwarz (2002) used this broad concept, subsequent authors (e. g. Hazir et al. 2014) have treated the three as distinct, though this has led to some confusion between A. paramythensis and A. optata, with the latter being reported as the former in some studies (e. g. Hazir et al. 2014; Pisanty et al. 2018).</p><p>Although Warncke considered these “ subspecies ” not to have overlapping ranges, A. optata and A. rufomaculata actually appear to have overlapping ranges in the Levant, with A. optata descending to Lebanon and northern Israel (Mount Hermon) (Pisanty et al. 2018, as A. paramythensis misidentification; Wood et al. 2020; Fig. 55). Whilst the morphological characters appear solid, available genetic sequences indicate that the differences within this group are very small. Three sequences of A. paramythensis showed very low intraspecific variation of 0.04 % (range 0.00–0.06 %). These were separated from two sequences of A. optata (Greece: Peloponnese) plus one sequence of A. rufomaculata (Lebanon) by an average genetic distance of 3.61 % (range 3.33–3.81 %). The sequences of A. optata were separated from A. rufomaculata by only 0.84 % (range 0.76–0.91 %). In absolute terms, this is a small genetic difference. Inclusion of additional sequences from Cyprus and the Levant actually show more complexity, with no consistent topological differences and sequences of all three taxa intermixed (G. Pisanty, pers. comm.). However, the ecological affinities of the two taxa suggest genuine species-level differences. In the Levant, A. optata is found at high altitude on Mount Hermon and the Mount Lebanon chain between 1200–2200 m a. s. l. whereas A. rufomaculata is much more thermophilic and is found at low to moderate altitudes. Specimens morphologically conforming to A. paramythensis are found only on the island of Cyprus. Consequently, as a temporary solution, A. optata is retained here as a good species despite the only weak to non-existent genetic separation from A. rufomaculata . This remains an area for future investigation using more powerful genetic tools to quantify realised gene flow in this group.</p><p>Overall, the genetic results from within Subgenus Truncandrena paint a challenging picture in which it is difficult to consistently translate high genetic distances into species-specific differences within existing morphological species concepts, and some existing morphological species concepts receive low to non-existent genetic support. Whilst the conclusions presented here are conservative, they maintain nomenclatural stability and allow morphological recognition of taxa which is beneficial for ecological study and monitoring of bee species pending further study.</p></div>	https://treatment.plazi.org/id/BEDE3D9319735378B71D552E1749A88A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
3AE81D87DAFF53FBB579A07476CE92EA.text	3AE81D87DAFF53FBB579A07476CE92EA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Ulandrena) concinna Smith 1853	<div><p>Andrena (Ulandrena) concinna Smith, 1853</p><p>Andrena concinna Smith, 1853: 106, ♀ [Greece, OUMNH, lectotype by present designation] (Fig. 59)</p><p>Material examined.</p><p>GREECE • 1 ♀; OUMNH (lectotype by present designation) .</p><p>Notes.</p><p>Wood (2024 a) discussed the concinna - group, and commented that the type of A. concinna was not in the NHMUK, the stated type repository (Gusenleitner and Schwarz 2002: 198). As for A. nigrifrons Smith, the species was described from the collection of W. W. Saunders. A single female labelled as “ A. 314 ” is from “ Albania ”, with the same caveat as applied above to A. nigrifrons Smith given the likely geographical place of capture. Baker (1993: 133) concluded that it was the holotype and stated that he labelled it as such, but no label was found. Following ICZN (1999) recommendation 73 F, I do not assume that it is the holotype (as Smith’s description does not indicate the number of specimens used to describe the species), and designate it instead as the lectotype. It conforms to the expected species, as A. concinna is the only member of this species complex that occurs in coastal parts of western Greece, though it geographically comes close to the more continental A. crecca Warncke, 1965 (Wood 2024 a; Fig. 60).</p><p>In addition to the morphological differences illustrated by Wood (2024 a), A. concinna shows an earlier phenology than A. crecca . Andrena concinna peaks between 16–31 May, with A. crecca peaking between 1–15 June (Fig. 61). Based on the median record, the respective dates are 19 May for A. concinna and 13 June for A. crecca . For this analysis, 89.0 % of the specimens used (see Suppl. material 2) were collected after 1980, suggesting that any confounding effects of climae change would be minor. Thus, despite having very similar ecological niches (specialists of Asteroideae ( Asteraceae), see section on the dietary niches of Balkan Andrena), they have slightly different climatic affinities and phenologies.</p></div>	https://treatment.plazi.org/id/3AE81D87DAFF53FBB579A07476CE92EA	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
5EC61E98D89A50843812638D4459CAF7.text	5EC61E98D89A50843812638D4459CAF7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Ulandrena) neocypriaca Mavromoustakis 1956	<div><p>Andrena (Ulandrena) waschulzi Strand, 1921 and Andrena (Ulandrena) neocypriaca Mavromoustakis, 1956</p><p>Material examined ( Andrena neocypriaca). BULGARIA • 3 ♀; Krumovgrad; 2 Jun. 1965; N. Atanassov leg.; NZMS • 1 ♂; Petrich env., steppes near Struma river; 100 m a. s. l.; 10 May 2018; D. Benda leg.; JSPC ; GREECE • 1 ♂; Agh. Konstantinos (Samos); 3 May 1977; H. Teunissen leg.; RMNH • 1 ♀; Faliraki, Rhodes; 23 Apr. 1976; H. Teunissen leg.; RMNH • 2 ♂, 2 ♀; Karfas / Chios; 11–15 May 1985; W. Perraudin leg.; OÖLM • 1 ♀; Lesvos, Eressos; 12–16 May 1988; K. M. Guichard leg.; RMNH • 1 ♂; Parakila (Lesbos); 14–17 May 1980; H. Teunissen leg.; RMNH • 2 ♀; Rhodes, fields 1 km WSW of Archipoli; 210 m a. s. l.; 28 May 2025; TJWC • 1 ♂; Rhodes, Kamiros; 21 Apr. 1970; v. Ooststroom leg.; RMNH • 1 ♀; Rhodes, Psinthos; 1 May 1976; H. Teunissen leg.; RMNH • 1 ♀; Rhodos, Mt. Profitis Ilias; 23 Apr. 1970; v. Oostroom leg.; RMNH • 1 ♀; Rodos, 5 km ZW v. Lindos; 29 Mar. 1970; A. C. &amp; W. N. Ellis leg.; RMNH • 1 ♀; Rodos, Lindos; 6 Apr. 1970; A. C. &amp; W. N. Ellis leg.; RMNH • 1 ♀; Samos, Kakkari [Kokkari], 1 km E; 7–19 Jun. 2019; J. Straka &amp; A. Straková leg.; JSPC • 1 ♀; Samos, Pythagorio, Umg. Hotel Maritsa Bay; 18–31 May 2006; J. Tiefenthaler leg.; OÖLM .</p><p>Notes. Andrena waschulzi was described from Crete (previously known as Andrena schulzi, see Wood 2024 a for discussion as to the correct spelling of his name), and A. neocypriaca was described from Cyprus. Warncke (1967 b: 230) described Andrena schulzi ssp. alba Warncke, 1967 from continental Greece, as the continental European population shows slight differences from the Cretan population. Warncke (1974 a: 89) indicated that A. schulzi was present on the island of Rhodes and probably the western coast of Turkey, and Hazir et al. (2012) reported A. waschulzi (as A. schulzi) from Çanakkale.</p><p>However, all examined material from the East Aegean Islands (Chios, Lesbos, Rhodes, and Samos) belongs to A. neocypriaca, and A. neocypriaca moves across western Turkey to southern Bulgaria, and it has not been possible to find any A. waschulzi from the eastern side of the Aegean sea (Fig. 65). Additional Bulgarian specimens were presented by Wood (2021). Andrena waschulzi is therefore considered to have its eastern range limit in continental Greece, and to display an allopatric distribution with A. neocypriaca . See Suppl. material 2 for full details on all examined material.</p></div>	https://treatment.plazi.org/id/5EC61E98D89A50843812638D4459CAF7	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
2E7665BCB3065AD1BE662DC37DFE9D26.text	2E7665BCB3065AD1BE662DC37DFE9D26.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena (Ulandrena) subglobosa Dours 1873	<div><p>Andrena (Ulandrena) subglobosa Dours, 1873 sp. resurr.</p><p>Campylogaster (Andrena) subglobosus Dours, 1873: 287, ♀ ♂ [Greece: Corfu, MSNG, lectotype by present designation] (Fig. 62)</p><p>Andrena punctatissima Kriechbaumer, 1873: 59, ♀ [Croatia, ZSM, examined] nec. Morawitz, 1866 (Fig. 63)</p><p>Andrena kriechbaumeri Schmiedeknecht, 1883: 569, nom. nov. pro Andrena punctatissima syn. nov.</p><p>Material examined.</p><p>CROATIA • 1 ♀; Lesina [= Croatian island of Hvar]; Heller leg.; ZSM (syntype of Andrena punctatissima Kriechbaumer) ; GREECE • 1 ♂; Corfu; MSNG (lectotype of Andrena subglobosa by present designation) . See Suppl. material 2 for additional examined material.</p><p>Notes.</p><p>The name Andrena kriechbaumeri has been used for many years to refer to an Ulandrena species found in the East Mediterranean which can rapidly be recognised due to the strongly postfurcal nervulus of the forewing. Andrena kriechbaumeri was created as a replacement name, as Andrena punctatissima Kriechbaumer is a junior primary homonym of Andrena punctatissima Morawitz (1866: 14) (see Astafurova et al. 2021). The species was described from what is now the island of Hvar in southern Croatia, and a single female syntype was present in the ZSM labelled with a typical Kriechbaumer label (Fig. 63). The species was described from two females, but it is not considered necessary to designate a lectotype at the present time.</p><p>The name Andrena subglobosa was created as Campylogaster subglobosus, but as Dours (1873: 286) created Campylogaster as a subgenus of Andrena, the name can be used in combination with Andrena without Dours’ name in parentheses. Andrena subglobosa was treated by Warncke (1967 b: 311) as a junior synonym of Andrena (Ulandrena) fulvitarsis Brullé, 1833 based only on the description. As for A. gravida Dours, the type material was considered lost, but a male specimen was located in the Gribodo collection (MSNG). It was labelled as “ Campylogaster subglobosus Typus Dours ”, and coming from Corfu (Fig. 62). The male description is scanty, but Dours indicates that the clypeus is yellow-marked. Overall, the description of the female is quite generic and does not allow recognition at a specific level, and so the synonymy by Warncke, whilst speculative, is understandable.</p><p>Nevertheless, this discovered specimen is not A. fulvitarsis, since the forewing shows the strongly postfurcal nervulus (Fig. 62 D), and the genital capsule is of the A. polita Smith, 1847 type (Fig. 62 F; shared with multiple species including A. kriechbaumeri). The specimen was examined by W. Grünwaldt and labelled as A. kriechbaumeri, but he either did not recognise its significance, or specifically did not recognise it as syntypic. Warncke (1967 b: 311) gave the Cyclades as the locus typicus, but as Dours (1873: 288) only indicates “ Iles de l’Archipel grec. ”, Warncke’s interpretation is not immediately justified. In Dours’ immediately preceding work, Dours (1872: 293) writes that specimens used for the description of Greek species come “ surtout dans les îles de l’Archipel grec: Syra, Tinos, Micanos, Corfou, etc ”. Following the reasoning of Wood and Bossert (2025), it is clear that Dours considered Corfu to be part of the Greek Archipelago, and since the specimen comes from Corfu and from the collection of Gribodo who was known to exchange specimens with Dours, this specimen is considered to be syntypic (as for taxa dealt with by Wood and Bossert 2025) and is hereby designated as the lectotype.</p><p>The purpose of the ICZN (1999) code is to promote nomenclatural stability. However, stability is not a universal criterion which overrides all other considerations. Although both A. subglobosa and A. punctatissima Kriechbaumer were described in the same year, the replacement name A. kriechbaumeri is junior to A. subglobosa by 10 years. Under Article 23, it would be beneficial to apply for Reversal of Precedence (23.9) to maintain the use of A. kriechbaumeri . However, A. kriechbaumeri fails Condition 23.9.1.1 (the senior synonym or homonym has not been used as a valid name after 1899) because Paganetti-Hummler (1912: 380) and Cockerell (1922: 549) use A. subglobosa as a valid name. Additionally, it also fails Condition 23.9.1.2 (the junior synonym or homonym has been used for a particular taxon, as its presumed valid name, in at least 25 works, published by at least 10 authors in the immediately preceding 50 years and encompassing a span of not less than 10 years) because I could find only 15 validly published works between 1976–2025 which mention A. kriechbaumeri in the main body of their text.</p><p>Given the limited use of the name A. kriechbaumeri in the literature (in the past 50 years, as defined by the ICZN), the position is taken that referring this case to the ICZN Commission under Article 23.9. 3 would be unlikely to be successful. Andrena subglobosa sp. resurr. is therefore returned to species status, and A. kriechbaumeri syn. nov. is synonymised with it.</p><p>Distribution.</p><p>Italy, Croatia, Bosnia &amp; Herzegovina, Montenegro, Albania, North Macedonia, Bulgaria, Greece, Turkey, Syria, and Iraq (Fig. 64; Gusenleitner and Schwarz 2002; Wood et al. 2024 b).</p></div>	https://treatment.plazi.org/id/2E7665BCB3065AD1BE662DC37DFE9D26	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
73E874EFD94A5F679CFF802492E86453.text	73E874EFD94A5F679CFF802492E86453.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Andrena lasithi Wood 2026	<div><p>Andrena (incertae sedis) lasithi sp. nov.</p><p>Figs 73, 74</p><p>Type material.</p><p>Holotype. GREECE • 1 ♂; Crete, <a href="https://tb.plazi.org/GgServer/search?materialsCitation.longitude=25.512&amp;materialsCitation.latitude=35.183" title="Search Plazi for locations around (long 25.512/lat 35.183)">Mesa Lasithi</a>; 35.1830°N, 25.5120°E; 27 Nov. 1986; H. Teunissen leg.; RMNH, RMNH.INS.1716262.</p><p>Paratypes. GREECE • 2 ♂; Crete, Mesa Lasithi; 3 Oct. 1985; H. Teunissen leg.; RMNH • 10 ♂, 12 ♀; Crete, Mesa Lasithi; 1 Nov. 1985; H. Teunissen leg.; OÖLM / RMNH / SMNHTAU • 6 ♂, 3 ♀; Crete, Mesa Lasithi; 3 Nov. 1985; H. Teunissen leg.; RMNH • 1 ♂, 12 ♀; Crete, Mesa Lasithi; 4 Nov. 1985; H. Teunissen leg.; OÖLM / RMNH • 5 ♀; Crete, Mesa Lasithi; 11 Nov. 1985; H. Teunissen leg.; RMNH • 5 ♀; Crete, Mesa Lasithi; 12 Nov. 1985; H. Teunissen leg.; RMNH • 4 ♀; Crete, Mesa Lasithi; 14 Nov. 1985; H. Teunissen leg.; RMNH • 9 ♀; Crete, Mesa Lasithi; 15 Nov. 1985; H. Teunissen leg.; RMNH • 10 ♀; Crete, Mesa Lasithi; 17 Nov. 1985; H. Teunissen leg.; RMNH / TJWC • 2 ♂, 14 ♀; Crete, Mesa Lasithi; 19 Nov. 1985; H. Teunissen leg.; RMNH • 8 ♀; Crete, Mesa Lasithi; 20 Nov. 1985; H. Teunissen leg.; RMNH • 1 ♀; Crete, Mesa Lasithi; 29 Nov. 1985; H. Teunissen leg.; RMNH • 1 ♂, 1 ♀; Crete, Mesa Lasithi; 19 Oct. 1986; H. Teunissen leg.; RMNH • 9 ♂, 13 ♀; Crete, Mesa Lasithi; 21 Oct. 1986; H. Teunissen leg.; OÖLM / RMNH / SMNHTAU • 1 ♂; Crete, Mesa Lasithi; 26 Oct. 1986; H. Teunissen leg.; RMNH • 8 ♂, 10 ♀; Crete, Mesa Lasithi; 27 Oct. 1986; H. Teunissen leg.; OÖLM / RMNH / TJWC • 15 ♀; Crete, Mesa Lasithi; 28 Oct. 1986; H. Teunissen leg.; OÖLM / RMNH / SMNHTAU • 1 ♂, 6 ♀; Crete, Mesa Lasithi; 29 Oct. 1986; H. Teunissen leg.; RMNH • 10 ♀; Crete, Mesa Lasithi; 30 Oct. 1986; H. Teunissen leg.; RMNH • 1 ♂, 1 ♀; Crete, Mesa Lasithi; 31 Oct. 1986; H. Teunissen leg.; RMNH • 1 ♀; Crete, Mesa Lasithi; 1 Nov. 1986; H. Teunissen leg.; RMNH • 23 ♀; Crete, Mesa Lasithi; 3 Nov. 1986; H. Teunissen leg.; RMNH / TJWC • 19 ♀; Crete, Mesa Lasithi; 4 Nov. 1986; H. Teunissen leg.; RMNH • 4 ♂; Crete, Plati – Lasithiplain; 11 Nov. 1985; H. Teunissen leg.; RMNH • 1 ♂; Crete, Plati – Lasithiplain; 12 Nov. 1985; H. Teunissen leg.; RMNH • 1 ♂; Kriti, Nom. Iraklion, Malia, 34 km OZO v. Iraklion; 29 Oct. 1972; A. C. &amp; W. N. Ellis leg.; RMNH • 5 ♂; Lasithi, Lasithi; 850 m a. s. l.; 7 Nov. 1984; H. Teunissen leg.; RMNH .</p><p>Description.</p><p>Female. Body length: 10–11 mm (Fig. 73 A). Head: Dark, 1.1 times wider than long (Fig. 73 B). Clypeus very weakly domed, more-or-less flattened over majority of surface, finely microreticulate in basal 1 / 2, weakly shining, becoming polished and shining in apical 1 / 2; surface punctate, punctures separated by 0.5–1 puncture diameter with exception of impunctate longitudinal midline. Process of labrum broadly trapezoidal, short, three times wider than long. Mouthparts long, at maximum extent exceeding length of head (Fig. 73 A); surface of galea finely microreticulate, weakly shining. Mouthparts with seven maxillary palpi, six labial palpi (Fig. 73 C). Gena slightly exceeding width of compound eye; ocelloccipital distance 1.5 times diameter of lateral ocellus. Foveae dorsally occupying 1 / 3 space between compound eye and lateral ocellus, of uniform width, equalling diameter of flagellum; facial foveae filled with dark brown hairs. Head covered with long black hairs, longest equalling length of scape, with few whitish hairs on posterior part of vertex. Antennae dark, A 4–12 slightly lightened ventrally by presence of greyish-brown scales; A 3 exceeding A 4 + 5, slightly shorter than A 4 + 5 + 6.</p><p>Mesosoma: Scutum with regular granular microreticulation, weakly shining, irregularly and shallowly punctate, punctures separated by 1–3 puncture diameters. Scutellum posteriorly with granular microreticulation, dull, anteriorly becoming polished and shining; more regularly punctate, punctures separated by 1–2 puncture diameters. Pronotum with humeral angle. Mesepisternum and dorsolateral parts of propodeum with regular dense microreticulation, weakly shining. Propodeal triangle laterally delineated with weak carinae, internal surface with weak rugae in basal 1 / 2 (Fig. 73 D). Mesepisternum with long black hairs, exceeding scape in length, hairs becoming slightly shorter on scutum and scutellum, with occasional intermixed pale hairs. Propodeal corbicula lacking anterior fringe, dorsal fringe composed of loose long black plumose hairs, internal surface with simple brownish hairs. Legs dark, apical tarsal segments lightened brownish, pubescence dark brown to blackish. Flocculus complete, composed of dark brown plumose hairs; femoral and tibial scopae composed of finely plumose hairs, femoral scopae and ventral part of tibial scopae whitish, dorsal part of tibial scopae dark brown (Fig. 73 E). Hind tarsal claws with inner subapical tooth. Wings hyaline, stigma and venation dark brown, nervulus interstitial to postfurcal.</p><p>Metasoma: Terga dark, apical rims narrowly lightened hyaline brown. Disc of T 1 polished and shining, disc of T 2–4 finely shagreened, weakly shining. Disc of T 1 with scattered punctures, punctures separated by 2–4 puncture diameters, discs of T 2–4 with punctures separated by 2–3 punctures; punctures weakly extending onto tergal margins, here becoming obscure. Tergal discs with sparse obscure dark hairs, tergal margins with weak obscure dark hairs, not forming apical hairbands and not obscuring underlying surface. T 5 and hairs flanking pygidial plate dark brown; pygidial plate rounded triangular, lateral margins slightly raised, internal surface extremely densely punctate, punctures confluent, surface dull.</p><p>Male. Body length: 9–10 mm (Fig. 74 A). Head: Dark, 1.3 times wider than long (Fig. 74 B). Clypeus more-or-less flattened, with weak longitudinal depression along apical margin; surface finely microreticulate, weakly shining, surface punctate, punctures separated by 0.5–1 puncture diameters, with unclear longitudinal impunctate midline. Process of labrum undifferentiated from labrum. Mouthparts as in female (Fig. 74 C). Gena strongly exceeding width of compound eye (Fig. 74 D); ocelloccipital distance 1.5 times diameter of lateral ocellus. Head covered with long dark hairs, longest equalling length of scape, becoming intermixed with white hairs on posterior part of vertex. Antennae dark, A 5–13 lightened ventrally by presence of greyish-brown scales; A 3 exceeding A 4 + 5, slightly shorter than A 4 + 5 + 6.</p><p>Mesosoma: Mesosoma structurally as in female. Mesosoma covered with long intermixed black and whitish hairs, longest hairs exceeding length of scape; pale hairs most abundant dorsally and ventrally, black hairs most abundant medially. Legs dark, apical tarsal segments lightened brown, pubescence light to dark brown; hind tarsal claw with inner subapical tooth. Wings hyaline, stigma dark brown, venation orange brown, nervulus interstitial to weakly postfurcal.</p><p>Metasoma: Metasoma structurally as in female (Fig. 74 E). Tergal discs with scattered short dark hairs, not obscuring underlying surface. S 8 columnar, ventral surface covered with thick fringe of dark brown hairs, apex with clear triangular emargination. Genital capsule elongate-rounded, gonocoxal teeth strongly produced into long rounded teeth, gonostyli strongly flattened and broadened apically, apices rounded (Fig. 74 F). Penis valves occupying slightly less than 1 / 2 space between gonostyli, laterally produced into sharp projecting points.</p><p>Diagnosis.</p><p>Andrena lasithi can quickly be recognised as a member of the undescribed subgenus best referred to as the crocusella - group (see Pisanty et al. 2022 a) due to the elongate palpi (Fig. 73 A), pronotum with a humeral angle (strongly produced in the male sex), barely indicated propodeal triangle (Fig. 73 D), oviform metasoma (Figs 73 F, 74 E), long and narrow foveae that remain a consistent width along their length, and elongate mandibles in the male sex (Fig. 74 B) which strongly cross apically (when closed), expanded gena which exceeds the width of the compound eye (Fig. 74 D), and genital capsule with the penis valves laterally produced into sharp projecting points (Fig. 74 F).</p><p>It can immediately be recognised due to the structure of the mouthparts; Andrena lasithi has seven maxillary palpi and six labial palpi (Figs 73 C, 74 C). Within the crocusella - group, Andrena grossella Grünwaldt, 1976 has nine maxillary palpi and nine labial palpi. All other members of the group (and essentially all other Andrena species) have just six maxillary palpi and four labial palpi. Andrena lasithi is also the only member of the crocusella - group currently reported from Crete; it may be the undescribed species from Crete mentioned by Pisanty et al. (2016: 514), but that was diagnosed as having seven maxillary palpi and seven labial palpi, but A. lasithi has seven maxillary palpi and only six labial palpi. There may be a second species of the crocusella - group present on Crete, but I have examined no specimens meeting this description.</p><p>Remarks.</p><p>Several specimens have large spherical yellow-orange pollen grains in their scopae (Fig. 73 E). These are strongly suspected to be Crocus ( Iridaceae) pollen – dietary specialisation on this plant genus would fit the behaviour (including autumn / early winter emergence) observed in A. grossella (Grünwaldt 1976) .</p><p>Etymology.</p><p>Named after the Lasithi Plateau in eastern Crete where Teunissen collected the species in numbers. It is a noun in apposition.</p><p>Distribution.</p><p>Greece (Crete).</p></div>	https://treatment.plazi.org/id/73E874EFD94A5F679CFF802492E86453	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Pensoft via Plazi	Wood, Thomas J.	Wood, Thomas J. (2026): The genus Andrena Fabricius, 1775 in the southern Balkans (Hymenoptera, Andrenidae). Journal of Hymenoptera Research 99: 249-604, DOI: 10.3897/jhr.99.179503
