taxonID	type	description	language	source
1EFE51B476275C6B8C125E74B87501D3.taxon	description	Figs 66 A – C, 66 E, 66 G, 67 A – E	en	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.taxon	description	Fig. 68	en	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.taxon	description	Fig. 69	en	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.taxon	description	Fig. 70 A – C, E, F	en	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.taxon	description	Figs 71, 72 A, B	en	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.taxon	discussion	Andrena lonicera Warncke, 1973 and Andrena lonicerae Tadauchi & Hirashima, 1988 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.). 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. 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. 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). 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.	en	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.taxon	discussion	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. 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. 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. 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.	en	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.taxon	materials_examined	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. & W. N. Ellis leg.; RMNH • 1 ♀; Rodos, Lindos; 6 Apr. 1970; A. C. & W. N. Ellis leg.; RMNH • 1 ♀; Samos, Kakkari [Kokkari], 1 km E; 7 – 19 Jun. 2019; J. Straka & A. Straková leg.; JSPC • 1 ♀; Samos, Pythagorio, Umg. Hotel Maritsa Bay; 18 – 31 May 2006; J. Tiefenthaler leg.; OÖLM.	en	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.taxon	discussion	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. 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.	en	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.taxon	description	Figs 73, 74	en	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
