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The new annotated checklist of the wild bees of Europe (Hymenoptera: Anthophila)

August 2023
Zootaxa 5327(1)

DOI:10.11646/zootaxa.5327.1.1

Authors:

Guillaume Ghisbain

Université de Mons

Paolo Rosa

Université de Mons

Petr Bogusch

University of Hradec Králové

Simone Flaminio

Council for Agricultural Research and Agricultural Economy Analysis

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At a time when nature conservation has become essential to ensure the long-term sustainability of our environment, it is widely acknowledged that conservation actions must be implemented within a solid taxonomic framework. In preparation for the upcoming update of the IUCN Red List, we here update the European checklist of the wild bees (sensu the IUCN geographical framework). The original checklist, published in 2014, was revised for the first time in 2017. In the present revision, we add one genus, four subgenera and 67 species recently described, 40 species newly recorded since the latest revision (including two species that are not native to Europe), 26 species overlooked in the previous European checklists and 63 published synonymies. We provide original records for eight species previously unknown to the continent and, as original taxonomic acts, we provide three new synonyms, we consider two names as nomina nuda, ten names as nomina dubia, three as species inquirenda, synonymize three species and exclude 40 species from the previous checklist. Around a hundred other taxonomic changes and clarifications are also included and discussed. The present work revises the total number of genera for IUCN Europe to 77 and the total number of species to 2,138. In addition to specifying the taxonomic changes necessary to update the forthcoming Red List of European bees, we discuss the sampling and taxonomic biases that characterise research on the European bee fauna and highlight the growing importance of range expansions and species invasions.

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https://doi.org/10.11646/zootaxa.5327.1.1

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ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)

Accepted by J. Gibbs: 13 Jun. 2023; published: 11 Aug. 2023

5327

ZOOTAXA

Magnolia Press

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Zootaxa 5327 (1): 001–147

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Licensed under Creative Commons Attribution-N.C. 4.0 International https://creativecommons.org/licenses/by-nc/4.0/

The new annotated checklist of the wild bees of Europe

(Hymenoptera: Anthophila)

GUILLAUME GHISBAIN

A1*+

, PAOLO ROSA

A2*+

, PETR BOGUSCH

, SIMONE FLAMINIO

A3,C

, ROMAIN

LE DIVELEC

, ACHIK DORCHIN

A5,D

, MAX KASPAREK

, MICHAEL KUHLMANN

, JESSE LITMAN

MAUD MIGNOT

, ANDREAS MÜLLER

, CHRISTOPHE PRAZ

, VLADIMIR G. RADCHENKO

, PIERRE

RASMONT

, STEPHAN RISCH

, STUART P.M. ROBERTS

, JAN SMIT

, THOMAS J. WOOD

, DENIS

MICHEZ

A9#

& SARA REVERTÉ

A10#

Laboratory of Zoology, Research Institute for Biosciences, University of Mons, Place du parc 20, 7000, Mons, Belgium.

Faculty of Science, University of Hradec Králové, Hradec Králové, Czech Republic.

�

bogusch.petr@gmail.com; https://orcid.org/0000-

0002-4554-6141;

Centro di Ricerca Agricoltura e Ambiente, (CREA) Consiglio per la Ricerca in Agricoltura e l’analisi dell’Economia

Agraria-via di Corticella 133, 40128 Bologna, Italy.

�

Simone.flaminio@crea.gov.it; https://orcid.org/0000-0002-5823-1202;

Musée

Royal de l’Afrique Centrale, Leuvensesteenweg 13, 3080 Tervuren, Belgium.

�

achik.dorchin@africamuseum.be; https://orcid.org/0000-

0003-1151-5926;

Mönchhofstr. 16, 69120 Heidelberg, Germany.

�

kasparek@t-online.de; https://orcid.org/0000-0002-5604-6791;

Zoological Museum, University of Kiel, Hegewischstr. 3, 24105 Kiel, Germany.

�

mkuhlmann@zoolmuseum.uni-kiel.de; https://orcid.

org/0000-0003-3664-6922;

Natural History Museum of Neuchâtel, Terreaux 14, 2000 Neuchâtel, Switzerland.

�

jessica.litman@unine.

ch; https://orcid.org/0000-0003-2481-5709;

ETH Zürich, Institute of Agricultural Sciences, Biocommunication and Entomology,

Schmelzbergstrasse 9/LFO, 8092 Zürich, Switzerland.

�

andreas.mueller@usys.ethz.ch; https://orcid.org/0000-0002-8322-1292;

Info

fauna Swiss Zoological Records Center, Avenue de Bellevaux 51, 2000 Neuchâtel, Switzerland. Institute of Biology, University of Neuchatel,

Rue Emile-Argand 16, 2000 Neuchâtel, Switzerland.

�

christophe.praz@unine.ch; https://orcid.org/0000-0003-2649-3141;

Institute for

Evolutionary Ecology of the National Academy of Sciences of Ukraine, acad. Lebedev, 37, 03143 Kiev, Ukraine.

�

rvg@nas.gov.ua;

https://orcid.org/0000-0002-8679-1362;

Imbacher Weg 59, D-51381 Leverkusen, Germany.

�

stephan.risch@t-online.de; https://orcid.

org/0000-0003-1628-0345;

Agroecology Lab,Université Libre de Bruxelles (ULB), Boulevard du Triomphe CP 264/02, 1050 Brussels,

Belgium.

�

spmr@msn.com; https://orcid.org/0000-0002-5473-4718;

Voermanstraat 14, 6921 Duiven, The Netherlands.

�

smit.jan@

hetnet.nl; https://orcid.org/ 0000-0001-9249-8739;

�

guillaume.ghisbain@umons.ac.be; https://orcid.org/0000-0003-2032-8081;

�

paolo.rosa@umons.ac.be; https://orcid.org/0000-0003-2919-5297;

�

simone.flaminio@umons.ac.be; https://orcid.org/0000-0002-

5823-1202;

�

romain.ledivelec@ umons.ac.be; https://orcid.org/0000-0002-0759-0344;

�

achik.dorchin@umons.ac.be; https://

orcid.org/0000-0003-1151-5926 ;

�

maud.mignot@umons.ac.be; https://orcid.org/0000-0002-4674-1811;

�

pierre.rasmont@umons.

ac.be; https://orcid.org/0000-0003-0891-2189;

�

thomasjames.wood@umons.ac.be; https://orcid.org/0000-0001-5653-224X;

�

denis.michez@umons.ac.be; https://orcid.org/0000-0001-8880-1838;

�

sara.reverte@umons.ac.be; https://orcid.org/0000-0002-2924-

3394;

Co-first authors;

Co-last authors;

Corresponding authors.

�

guillaume.ghisbain@umons.ac.be;

�

paolo.rosa@umons.ac.be;

GHISBAIN, ROSA ET AL.

GUILLAUME GHISBAIN, PAOLO ROSA, PETR BOGUSCH, SIMONE FLAMINIO, ROMAIN LE

DIVELEC, ACHIK DORCHIN, MAX KASPAREK, MICHAEL KUHLMANN, JESSE LITMAN,

MAUD MIGNOT, ANDREAS MÜLLER, CHRISTOPHE PRAZ, VLADIMIR G. RADCHENKO,

PIERRE RASMONT, STEPHAN RISCH, STUART P.M. ROBERTS, JAN SMIT, THOMAS J. WOOD,

DENIS MICHEZ & SARA REVERTÉ

THE NEW ANNOTATED CHECKLIST OF THE WILD BEES OF EUROPE (HYMENOPTERA:

ANTHOPHILA)

(Zootaxa 5327)

147 pp.; 30 cm.

11 Aug. 2023

ISBN 978-1-77688-838-2 (paperback)

ISBN 978-1-77688-839-9 (Online edition)

FIRST PUBLISHED IN 2023 BY

Magnolia Press

P.O. Box 41-383

Auckland 1041

New Zealand

e-mail: magnolia@mapress.com

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This authorization does not extend to any other kind of copying, by any means, in any form, and for any

purpose other than private research use.

ISSN 1175-5326 (Print edition)

ISSN 1175-5334 (Online edition)

Table of Contents

Abstract ...................................................................................................7

Introduction ................................................................................................7

Material and methods ........................................................................................8

Results ..................................................................................................10

Taxonomic update of the wild bee fauna of IUCN Europe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

Family ANDRENIDAE Latreille, 1802 .........................................................................10

Tribe Andrenini Latreille, 1802 ...............................................................................10

Species recently described as new to science ....................................................................10

Published synonymies .......................................................................................15

Synonymic notes ...........................................................................................17

Taxonomic changes .........................................................................................18

Taxonomic acts and clarifications ..............................................................................21

Species recorded in Europe after 2017 ..........................................................................22

New species for Europe .....................................................................................22

Species overlooked in the previous European checklists ............................................................22

Species to be excluded from the European checklist ...............................................................23

Tribe Panurgini Leach, 1815 ..................................................................................23

Genus described ...........................................................................................23

Species recently described as new to science .....................................................................23

Taxonomic changes .........................................................................................23

Species overlooked in the previous European checklists ............................................................25

Family APIDAE Latreille, 1802 ...............................................................................25

Tribe Ammobatini Handlirsch, 1925 ...........................................................................25

Published synonymies .......................................................................................25

Tribe Ancylaini Michener, 1944 ...............................................................................25

Species to be excluded from the European checklist ...............................................................25

Tribe Anthophorini Dahlbom, 1835 ............................................................................25

Published synonymies .......................................................................................25

New synonymies ...........................................................................................26

Taxonomic acts and clarifications ..............................................................................26

Species recorded in Europe after 2017 ..........................................................................27

New species for Europe .....................................................................................27

Species to be excluded from the European checklist ...............................................................28

Tribe Melectini Westwood, 1839 ..............................................................................28

Taxonomic changes .........................................................................................28

Species recorded in Europe after 2017 ..........................................................................29

Species overlooked in the previous European checklists ............................................................29

Species to be excluded from the European checklist ...............................................................29

Tribe Bombini Latreille, 1802 ................................................................................29

Species recently described as new to science .....................................................................29

Taxonomic changes .........................................................................................30

Taxonomic acts and clarifications ..............................................................................31

Tribe Ceratinini Latreille, 1802 ...............................................................................33

Species to be excluded from the European checklist ...............................................................33

Tribe Epeolini Linsley & Michener, 1939 .......................................................................34

Species recently described as new to science ....................................................................34

Taxonomic acts and clarifications ..............................................................................34

Species recorded in Europe after 2017 ..........................................................................35

Tribe Eucerini Latreille, 1802 .................................................................................35

Species recently described as new to science .....................................................................35

Published synonymies .......................................................................................36

Taxonomic changes .........................................................................................37

Species recorded in Europe after 2017 ..........................................................................38

Species overlooked in the previous European checklists ............................................................39

Species to be excluded from the European checklist ...............................................................39

Tribe Nomadini Latreille, 1802 ...............................................................................40

Species recently described as new to science .....................................................................40

Published synonymies .......................................................................................44

Taxonomic acts and clarifications ..............................................................................45

Species recorded in Europe after 2017 ..........................................................................46

Species overlooked in the previous European checklists ............................................................47

New species for Europe .....................................................................................47

GHISBAIN, ROSA ET AL.

Species to be excluded from the European checklist ...............................................................47

Family COLLETIDAE Lepeletier, 1841 ........................................................................47

Tribe Colletini Lepeletier, 1841 ...............................................................................47

Species recently described as new to science ....................................................................47

Species recorded in Europe after 2017 ..........................................................................48

Species overlooked in the previous European checklists ............................................................48

Tribe Hylaeini Viereck, 1916 .................................................................................48

Published synonymies .......................................................................................48

Taxonomic acts and clarifications ..............................................................................49

Species recorded in Europe after 2017 ..........................................................................50

Species overlooked in the previous European checklists ............................................................51

Species to be excluded from the European checklist ...............................................................51

Family HALICTIDAE Thomson, 1869 .........................................................................51

Subfamily Halictinae Thomson, 1869 ..........................................................................52

Species recently described as new to science ....................................................................52

Published synonymies .......................................................................................52

Taxonomic changes .........................................................................................53

Taxonomic acts and clarifications ..............................................................................53

Species overlooked in the previous European checklists ............................................................54

Species to be excluded from the European checklist ...............................................................54

Subfamily Nomiinae ........................................................................................55

Species recently described as new to science ....................................................................55

Taxonomic acts and clarifications ..............................................................................55

Species recorded in Europe after 2017 ..........................................................................56

Species overlooked in the previous European checklists ............................................................56

Subfamily Rophitinae Schenk, 1866 ............................................................................56

Taxonomic acts and clarifications ..............................................................................56

Species to be excluded from the European checklist ...............................................................56

Family MEGACHILIDAE Latreille, 1802 .......................................................................56

Tribe Anthidiini Ashmead, 1899 ...............................................................................56

Species recently described as new to science ....................................................................56

Published synonymies .......................................................................................58

Taxonomic changes .........................................................................................58

Taxonomic acts and clarifications ..............................................................................58

Species recorded in Europe after 2017 ..........................................................................60

Species overlooked in the previous European checklists ............................................................60

Species to be excluded from the European checklist ...............................................................61

Tribe Megachilini Latreille, 1802 ..............................................................................61

Published synonymies .......................................................................................61

New synonymies ...........................................................................................63

Taxonomic acts and clarifications ..............................................................................63

Species recorded in Europe after 2017 ..........................................................................66

Species overlooked in the previous European checklists ............................................................67

Species to be excluded from the European checklist ...............................................................67

Tribe Osmiini Newman, 1834 .................................................................................68

Species recently described as new to science ....................................................................68

Published synonymies .......................................................................................68

Taxonomic changes .........................................................................................68

Taxonomic acts and clarifications ..............................................................................69

Species recorded in Europe after 2017 ..........................................................................70

New species for Europe .....................................................................................70

Species overlooked in the previous European checklists ............................................................70

Species to be excluded from the European checklist ...............................................................70

Family MELITTIDAE Schenck, 1860 ..........................................................................72

Tribe Dasypodaini Sagemehl, 1882 ............................................................................72

Species recently described as new to science ....................................................................72

Synonymic notes ...........................................................................................72

Species overlooked in the previous European checklists ............................................................72

Updated checklist of the wild bee fauna of Europe ................................................................74

Family Andrenidae Latreille, 1802 .............................................................................74

Tribe Andrenini Latreille, 1802 ...............................................................................74

Genus Andrena Fabricius, 1775 ...............................................................................74

Genus Cubiandrena Warncke, 1968 ............................................................................85

Tribe Melliturgini Newman, 1834 .............................................................................85

Genus Melitturga Latreille, 1809 ..............................................................................85

Tribe Panurgini Leach, 1815 ..................................................................................85

Genus Camptopoeum Spinola, 1843 ...........................................................................85

Genus Clavipanurgus Warncke, 1972 ..........................................................................85

Genus Flavipanurgus Warncke, 1972 ..........................................................................86

Genus Halopanurgus Wood, Patiny & Bossert, 2022 ..............................................................86

Genus Panurginus Nylander, 1848 ............................................................................86

Genus Panurgus Panzer, 1806 ................................................................................86

Genus Simpanurgus Warncke, 1972 ...........................................................................87

Family Apidae Latreille, 1802 ................................................................................87

Tribe Ammobatini Handlirsch, 1925 ...........................................................................87

Genus Ammobates Latreille, 1809 .............................................................................87

Genus Chiasmognathus Engel, 2006 ...........................................................................87

Genus Parammobatodes Popov, 1931 ..........................................................................87

Genus Pasites Jurine, 1807 ...................................................................................87

Tribe Ammobatoidini Michener, 1944 ..........................................................................87

Genus Ammobatoides Radoszkowski, 1867 ......................................................................87

Genus Schmiedeknechtia Friese, 1896 .........................................................................88

Tribe Ancylaini Michener, 1944 ...............................................................................88

Genus Ancyla Lepeletier, 1841 ................................................................................88

Genus Tarsalia Morawitz, 1895 ...............................................................................88

Tribe Anthophorini Dahlbom, 1835 ............................................................................88

Genus Amegilla Friese, 1897 .................................................................................88

Genus Anthophora Latreille, 1803 .............................................................................88

Genus Habropoda Smith, 1854 ...............................................................................90

Tribe Apini Latreille, 1802 ...................................................................................90

Genus Apis Linnaeus, 1758 ..................................................................................90

Tribe Biastini Linsley & Michener, 1939 ........................................................................91

Genus Biastes Panzer, 1806 ..................................................................................91

Tribe Bombini Latreille, 1802 ................................................................................91

Genus Bombus Latreille, 1802 ................................................................................91

Tribe Ceratinini Latreille, 1802 ...............................................................................93

Genus Ceratina Latreille, 1802 ...............................................................................93

Tribe Epeolini Robertson, 1903 ...............................................................................93

Genus Epeolus Latreille, 1802 ................................................................................93

Genus Triepeolus Robertson, 1901 ............................................................................94

Tribe Epeoloidini Linsley & Michener, 1939 ....................................................................94

Genus Epeoloides Giraud, 1863 ...............................................................................94

Tribe Eucerini Latreille, 1802 .................................................................................94

Genus Eucera Scopoli, 1770 .................................................................................94

Genus Tetralonia Spinola, 1839 ...............................................................................96

Tribe Melectini Westwood, 1839 ..............................................................................96

Genus Melecta Latreille, 1802 ................................................................................96

Genus Thyreus Panzer, 1806 .................................................................................97

Tribe Nomadini Latreille, 1802 ...............................................................................97

Genus Nomada Scopoli, 1770 ................................................................................97

Tribe Xylocopini Latreille, 1802 .............................................................................102

Genus Xylocopa Latreille, 1802 ..............................................................................102

Family Colletidae Lepeletier, 1841 ............................................................................102

Tribe Colletini Latreille, 1802 ...............................................................................102

Genus Colletes Latreille, 1802 ...............................................................................102

Tribe Hylaeini Viereck, 1916 ................................................................................104

Genus Hylaeus Fabricius, 1793 ..............................................................................104

Family Halictidae Thomson, 1869 ............................................................................106

Subfamily Halictinae Thomson, 1869 .........................................................................106

Genus Halictus Latreille, 1804 ...............................................................................106

Genus Lasioglossum Curtis, 1833 ............................................................................107

Genus Seladonia Robertson, 1918 ............................................................................ 111

Genus Sphecodes Latreille, 1804 ............................................................................. 112

Genus Thrincohalictus Blüthgen, 1955 ........................................................................113

Subfamily Nomiinae Robertson, 1904 .........................................................................113

Genus Nomiapis Cockerell, 1919 .............................................................................113

Genus Pseudapis Kirby, 1900 ................................................................................113

Subfamily Nomioidinae Börner, 1919 .........................................................................113

GHISBAIN, ROSA ET AL.

Genus Ceylalictus Strand, 1913 ..............................................................................113

Genus Nomioides Schenck, 1867 .............................................................................113

Subfamily Rophitinae Schenck, 1866 .........................................................................114

Genus Dufourea Lepeletier, 1841 ............................................................................ 114

Genus Rhophitoides Schenck, 1861 ...........................................................................114

Genus Rophites Spinola, 1808 ...............................................................................114

Genus Systropha Illiger, 1805 ............................................................................... 115

Family Megachilidae Latreille, 1802 ..........................................................................115

Tribe Anthidiini Ashmead, 1899 ..............................................................................115

Genus Afranthidium Michener, 1948 .........................................................................115

Genus Anthidiellum Cockerell, 1904 ..........................................................................115

Genus Anthidium Fabricius, 1804 ............................................................................115

Genus Eoanthidium Popov, 1950 .............................................................................116

Genus Icteranthidium Michener, 1948 ........................................................................ 116

Genus Pseudoanthidium Friese, 1898 ......................................................................... 116

Genus Rhodanthidium Isensee, 1927. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116

Genus Stelis Panzer, 1806 ...................................................................................117

Genus Trachusa Panzer, 1804 ............................................................................... 117

Tribe Dioxyini Cockerell, 1902 .............................................................................. 118

Genus Aglaoapis Cameron, 1901 .............................................................................118

Genus Dioxys Lepeletier & Serville, 1825 ......................................................................118

Genus Ensliniana Cameron, 1901 ............................................................................ 118

Genus Metadioxys Popov, 1947 ..............................................................................118

Genus Paradioxys Mocsáry, 1894 ............................................................................118

Tribe Lithurgini Newman, 1834 ..............................................................................118

Genus Lithurgus Latreille, 1825 .............................................................................118

Tribe Megachilini Latreille, 1802 .............................................................................118

Genus Coelioxys Latreille, 1809 ..............................................................................118

Genus Megachile Latreille, 1802 ............................................................................. 119

Tribe Osmiini Newman, 1834 ................................................................................121

Genus Chelostoma Latreille, 1809 ............................................................................121

Genus Haetosmia Popov, 1952 ...............................................................................122

Genus Heriades Spinola, 1808 ...............................................................................122

Genus Hofferia Tkalců, 1984 ................................................................................122

Genus Hoplitis Klug, 1807 ..................................................................................122

Genus Osmia Panzer, 1806 ..................................................................................125

Genus Protosmia Ducke, 1900 ...............................................................................127

Genus Stenoheriades Tkalců, 1984 ...........................................................................128

Family Melittidae Schenck, 1860 .............................................................................128

Tribe Dasypodaini Sagemehl, 1882 ...........................................................................128

Genus Dasypoda Latreille, 1802 ..............................................................................128

Tribe Macropidini Robertson, 1904 ...........................................................................128

Genus Macropis Panzer, 1809 ...............................................................................128

Tribe Melittini Schenck, 1860 ...............................................................................128

Genus Melitta Kirby, 1802 ..................................................................................128

Discussion ...............................................................................................129

Acknowledgements ........................................................................................131

References ...............................................................................................134

Abstract

At a time when nature conservation has become essential to ensure the long-term sustainability of our environment, it is

widely acknowledged that conservation actions must be implemented within a solid taxonomic framework. In preparation

for the upcoming update of the IUCN Red List, we here update the European checklist of the wild bees (sensu the IUCN

geographical framework). The original checklist, published in 2014, was revised for the first time in 2017. In the present

revision, we add one genus, four subgenera and 67 species recently described, 40 species newly recorded since the latest

revision (including two species that are not native to Europe), 26 species overlooked in the previous European checklists

and 63 published synonymies. We provide original records for eight species previously unknown to the continent and, as

original taxonomic acts, we provide three new synonyms, we consider two names as nomina nuda, ten names as nomina

dubia, three as species inquirenda, synonymize three species and exclude 40 species from the previous checklist. Around

a hundred other taxonomic changes and clarifications are also included and discussed. The present work revises the total

number of genera for IUCN Europe to 77 and the total number of species to 2,138. In addition to specifying the taxonomic

changes necessary to update the forthcoming Red List of European bees, we discuss the sampling and taxonomic biases

that characterise research on the European bee fauna and highlight the growing importance of range expansions and

species invasions.

Key words: Biodiversity, conservation, continental checklist, decline, expansion, new synonym, pollinators, species

inquirenda, status resurrectus, nomen dubium, non-natives species, nomen nudum, taxonomy

Introduction

Biotic losses are accelerating at an alarming rate globally (Del Claro & Dirzo 2021; Turvey & Crees 2019). Millions

of hectares of natural habitats are cleared each year, a large part of the Earth’s suitable land has already been

converted to agriculture, and the global climate is changing, causing unprecedented disruptions for wildlife (Curtis

et al. 2018; Dirzo et al. 2014; Hautier et al. 2015; IPBES 2019; Raven & Wagner 2021; Storch et al. 2022).

There is overwhelming evidence that the combination of these global changes has induced a major biodiversity

crisis, compared by some authors to a mass extinction (Ceballos et al. 2020; Cowie et al. 2022). Among the clades

most severely affected by the ongoing biodiversity crisis are insects and a flurry of reports has drawn attention to

significant declines in their abundance, biomass, diversity and spatial distribution (Forister et al. 2019; Hallman et

al. 2020; Lister & Garcia 2018; Loboda et al. 2018; Wagner 2020; Wagner et al. 2021).

Wild bees (Hymenoptera: Anthophila), with over 20,000 species described globally (Michener 2007; Ascher &

Pickering 2022), are one example of a highly emblematic insect group for which conservation is tightly linked to

both human welfare and ecosystem health (Matias et al. 2017; Potts et al. 2016). Bees are not only key to the yield

of ~85% of cultivated crop species globally, but also to the sexual reproduction of hundreds of thousands of other

plant species, making them critical providers of ecosystem services (Garibaldi et al. 2013; Ollerton et al. 2011).

In the last two decades however, high-profile reports have highlighted steep declines in both bee abundance and

species diversity, most clearly in Europe and North America (Biesmeijer et al. 2006; Duchenne et al. 2020; LeBuhn

& Vargas Luna 2021; Rasmont et al. 2021; Scheper et al. 2014; Zattara & Aizen 2021). Although a large part of the

research on non-domesticated bees has focused on bumblebees (Cameron & Sadd 2020; Ghisbain 2021), there is

mounting evidence that the patterns of decline are affecting many other bee genera (Nieto et al. 2014; Rasmussen

et al. 2021; Zattara & Aizen 2021).

In a time when nature conservation has become key to ensuring the long-term sustainability of our environment,

it is acknowledged that conservation actions must be taken within a robust, unambiguous taxonomic background

(Garnett & Christidis 2017; Mace 2004; Orr et al. 2021). This is especially true for clades like bees, for which

substantial taxonomic revisions are still underway across most genera and families globally (e.g. Bossert et al.

2022; Dorchin et al. 2018; Ferrari et al. 2020; Müller 2020; Onuferko et al. 2019; Orr et al. 2018; Williams et al.

2020). Although the current knowledge of the European bee fauna is substantial (Michez et al. 2019), new species

are frequently described (e.g. Praz et al. 2019; Radchenko 2017; Wood et al. 2021), and other major nomenclatural

changes still occur after more than 250 years of research on the continent (e.g. the description of a new genus

by Wood et al. 2022a). Such updates, fundamental to refine the knowledge of the bee fauna of the continent,

also demonstrate that many uncertainties persist regarding the taxonomy and distribution of European wild bees

(Rasmont et al. 2017). These uncertainties not only hinder our ability to identify accurately both museum specimens

GHISBAIN, ROSA ET AL.

and freshly collected material, but also impede our understanding of their temporal and spatial distributions, limiting

the efficiency of action plans. This issue was exemplified in the first Red List of European Bees in which ~55% of

all bees reported in the continent had to be classified as “Data Deficient” (DD) (Nieto et al. 2014). In this work,

although ~9% of bees were considered threatened, the real percentage of threatened taxa would have been between

4% (if none of the DD species was threatened) and ~60% (if all of the DD species were threatened). With data from

other animal groups suggesting that unassessed and DD species are more likely to be threatened with extinction

than their fully assessed counterparts (Caetano et al. 2022; Howard & Bickford 2014), it is clear that more work

on the taxonomy and natural history of wild bee species is urgently required to accurately implement adequate

conservation strategies for the European bee fauna (Potts et al. 2020).

In May 2019, the European Commission mandated a group of experts to develop a proposal for monitoring

pollinators (including wild bees) and to analyse the capacity for its implementation. A report evaluating the possibility

of an EU Pollinator Monitoring Scheme (‘the EU-PoMS’) was published in October 2020 and pointed out several

gaps for its implementation, including insufficient taxonomic resources for bees (Potts et al. 2020). Following this, a

series of projects were implemented (i) to strengthen taxonomic capacity in EU Member States regarding wild bees,

(ii) to better understand the causes of decline of wild bees on the continent and (iii) to develop capacities to allow

for their effective monitoring and conservation. The present work is therefore at the hub of the ongoing European

projects ORBIT (Developing resources for European bee inventory and taxonomy, aiming to create and centralise

taxonomic information about all the European bee species, 2021-2024), SPRING (Strengthening pollinator recovery

through indicators and monitoring, aiming to train researchers on pollinator identification and refine sampling

protocols to start a European-wide monitoring of pollinators, 2021-2023), SAFEGUARD (Safeguarding European

wild pollinators, aiming to expand current assessments of the status and trends of European wild pollinators, 2021-

2025), and PULSE (Providing technical and scientific support in measuring the pulse of European biodiversity

using the Red List Index, aiming to update the European Red List of Bees, 2022-2023).

Within the context of the aforementioned projects, we provide a new, updated and annotated checklist of bee

species occurring in the spatial area defined by the European borders proposed by the IUCN (henceforth IUCN

Europe). The first available update was published by Rasmont et al. (2017), who revised the total number of bee

species in Europe from 1,965 (in Nieto et al. 2014) to 2,051, increased the number of recognised genera from 75 to

77, and included changes to taxonomy and nomenclature. The present work is a new revision that includes the most

recent taxonomic advances and revisions in order to stabilise the taxonomic backbone of the European bee fauna for

the forthcoming European country records of wild bees (Reverté et al. in prep.), the new IUCN Red List (Boustani

et al. in prep.) and the online ORBIT platform.

Material and methods

Bringing together new literature records and taxonomic updates for this work was made possible by (i) an exhaustive

review of the literature published since the last update of the IUCN checklist of European bees (Rasmont et al.

2017), (ii) an in-depth revision of the literature omitted in the latter work and (iii) original information provided

by the authors of the present work. For the present checklist, we did not consider the list provided by Michez et

al. (2019), as the latter was not built as an update to the previous IUCN checklists and additions. This new list is

mostly based on material directly examined by taxonomists and does not include data published online that has not

otherwise been verified by European experts (e.g. observations reported on iNaturalist, Discover Life, GBIF).

Authors shared the work based on their own expertise on bee taxonomy: Petr Bogusch and Romain Le Divelec

for Ammobatini, Ammobatoidini, Epeolini and Sphecodes; Thomas J. Wood for Andrenidae and Thyreus; Pierre

Rasmont for Anthophorini; Jessica Litman and Max Kasparek for Anthidiini; Guillaume Ghisbain for Bombini;

Michael Kuhlmann and Romain Le Divelec for Colletidae; Achik Dorchin for Eucerini; Simone Flaminio and

Vladimir G. Radchenko for Halictidae; Christophe Praz for Megachilini; Guillaume Ghisbain, Denis Michez and

Vladimir G. Radchenko for Melittidae; Maud Mignot and Jan Smit for Nomada; Andreas Müller for Osmiini.

Authors keep the authority on the original taxonomic updates they apply here to their group(s) of expertise.

European borders

The geographical scope for this study is the territory considered in the regional assessment of Europe according to

IUCN (Fig. 1). The area includes the European mainland, which includes the European part of Turkey (Thrace).

The natural limits on the eastern side are the Ural Mountains, which are included, and the Caucasus, which is

not included. The European part of Russia includes the following Republics and administrative provinces: North

(Murmansk Prov., Arkhangelsk Prov., Karelian Rep., Vologda Prov., Komi Rep.); North-West (Kaliningrad Prov.,

Leningrad Prov., Pskov Prov., Novgorod Prov.); Centre (Tver Prov., Yaroslavl Prov., Kostroma Prov., Smolensk

Prov., Moscow Prov., Vladimir Prov., Ivanovo Prov., Nizhny Novgorod Prov., Kaluga Prov., Tula Prov., Ryazan

Prov., Mordovian Rep., Bryansk Prov., Orel Prov., Lipetsk Prov., Tambov Prov., Penza Prov., Kursk Prov., Belgorod

Prov., Voronezh Prov.); East (Kirov Prov., Udmurt Rep., Mari El Rep., Chuvash Rep., Tatar Rep., Ulyanovsk Prov.,

Samara Prov., Saratov Prov.); South (Rostov Prov., Volgograd Prov., Kalmyk Rep., Astrakhan Prov.).

European Island territories such as Great Britain, Ireland, Iceland, and Cyprus are included as well as the Macaronesian

islands that are part of European countries (Azores, Madeira, Canary Islands). A detailed discussion on the European

borders is given in Rasmont et al. (2017).

FIGURE 1. Map of IUCN Europe, corresponding to the geographical framework of this study.

How to read the catalogue

The species are ordered by family, subfamily and tribe presented in alphabetical order, and listed alphabetically

within the following sections:

- Species recently described as new to science (i.e. new species described after 2017);

- Published synonymies (i.e. synonymies published after 2017);

- New synonymies (i.e. new synonymies proposed in this manuscript);

- Synonymic notes (i.e. notes related to future synonymisations in articles currently in preparation);

- Taxonomic changes (i.e. relevant changes published after 2017, such as new combinations, taxa upgraded to

species rank or downgraded to subspecies rank);

- Taxonomic acts and clarifications (i.e. taxonomic acts here proposed and clarifications of interesting cases

which generally led to changes in the new checklist of the European bees);

- Species recorded in Europe after 2017 (i.e. new to Europe but not new to science);

- Species overlooked in the previous European checklists (i.e. species recorded in Europe before 2017 but not

included in the update of Rasmont et al. 2017);

- New species for Europe (new entries presented in this article for the first time);

GHISBAIN, ROSA ET AL.

- Species to be excluded from the European checklist (discussions and explanations of the exclusions of

certain species from the new checklist).

The systematics at family, subfamily and tribe levels are mainly based on the hypotheses reviewed by Danforth et

al. (2013) and followed by Michez et al. (2019). We considered here the tribes of Andrenidae proposed by Bossert

et al. (2022), and the subfamilies of Apidae used in Bossert et al. (2019).

The following abbreviations are used for museums and private collections:

GGPC—Private collection of Gérard Le Goff, Barentin, (France)

IZKP—Institute of Systematic and Experimental Zoology, Polish Academy of Sciences, Kraków (Poland)

MNCN—Museo Nacional de Ciencias Naturales, Madrid (Spain)

MHNN—Muséum d’histoire naturelle de Neuchâtel (Switzerland)

MKPC—Private collection of Max Kasparek, Heidelberg (Germany)

MNHN—Muséum national d’Histoire naturelle, Paris (France)

MSPC—Private collection of Maximilian Schwarz, Ansfelden (Austria)

NHMUK—Natural History Museum, London (United Kingdom)

OÖLM—Oberösterreich Landesmuseum, Biologiezentrum, Linz (Austria)

PRUN—Research collection of Christophe Praz, University of Neuchatel (Switzerland)

RMNH—Naturalis Biodiversity Center, Leiden (the Netherlands)

SMF—Senckenberg Museum, Frankfurt (Germany)

SDEI—Senckenberg Deutsches Entomologisches Institut, Müncheberg (Germany)

SMNH—Steinhardt Museum of Natural History, University of Tel Aviv (Israel)

UAL—University of Almería, La Cañada de San Urbano Almería (Spain)

UCSI—Department of Agriculture, Food and Environment, University of Calabria, Rende (Italy)

UMONS—Laboratory of Zoology, University of Mons (Belgium)

ZISP—Zoological Institute, St. Petersburg (Russia)

ZMHB –Museum für Naturkunde, Berlin (Germany)

ZMKU—Research collection of Michael Kuhlmann, Zoological Museum, University of Kiel (Germany)

ZSM—Zoologische Staatssammlung München (Germany)

Results

Taxonomic update of the wild bee fauna of IUCN Europe

Family ANDRENIDAE Latreille, 1802

Tribe Andrenini Latreille, 1802

Species recently described as new to science

Andrena (Euandrena) amieti Praz, Müller & Genoud, 2019

Andrena (Euandrena) amieti Praz, Müller & Genoud, 2019: 20. Holotype ♀; Switzerland: Bernese Oberland: Oeschinensee

(MHNN).

Distribution. France, Switzerland, Germany, Italy, Austria.

Andrena (Taeniandrena) antonellae Praz & Genoud, 2022

Andrena (Taeniandrena) antonellae Praz & Genoud in Praz et al., 2022: 390. Holotype ♀; Italy: Sardinia, Buggerru, Cala

Domestica (PRUN).

Distribution. France (Corsica) and Italy (Sardinia).

Andrena (Lepidandrena) baetica Wood, 2020

Andrena (Lepidandrena) baetica Wood in Wood et al., 2020a: 202. Holotype ♀; Portugal: Alto Alentejo, Portalegre, Vaiamonte

(OÖLM).

Distribution. Portugal, Spain.

Andrena (Taeniandrena) benoisti Wood & Praz, 2021

Andrena (Taeniandrena) benoisti Wood & Praz in Wood et al., 2021: 162. Holotype ♀; Portugal: Minho, Confurco, Várzea Cova

(OÖLM).

Distribution. Portugal, Spain.

Andrena (Taeniandrena) contracta Wood, 2022

Andrena (Taeniandrena) contracta Wood, 2022: 2. Holotype ♂; Spain: Sierra Nevada, Puerto de La Ragua, Barranco Maja Caco

(OÖLM).

Distribution. Spain.

Andrena (Avandrena) erodiorum Wood & Ortiz-Sánchez, 2022

(Figs 2A, B)

Andrena (Avandrena) erodiorum Wood & Ortiz-Sánchez, 2022: 115. Holotype ♀; Spain: Castilla–La Mancha, Albacete, Torre

de Gorgojí, Reolid (OÖLM).

Distribution. Spain.

Andrena (Notandrena) foeniculae Wood, 2020

Andrena (Notandrena) foeniculae Wood in Wood et al., 2020a: 209. Holotype ♀; Portugal: Ribatejo, Tomar, Aqueduto do

Convento de Cristo (OÖLM).

Distribution. Portugal, Spain.

Andrena (Euandrena) fortipunctata Wood, 2021

Andrena (Euandrena) fortipunctata Wood in Wood et al., 2021: 165. Holotype ♂; Spain: in between Asturias and Castilla y

León, Puerto de Pajares (RMNH).

Distribution. Spain.

GHISBAIN, ROSA ET AL.

Andrena (Suandrena) gades Wood & Ortiz-Sánchez, 2022

Andrena (Suandrena) gades Wood & Ortiz-Sánchez, 2022: 118. Holotype ♂; Spain: Andalusia, Cádiz, Rota, Punta Candor

(OÖLM).

Distribution. Spain.

FIGURE 2. A. Andrena erodiorum Wood & Ortiz-Sánchez, 2022 female, habitus in lateral view. The species was recently

described in Wood & Ortiz- Sánchez (2022) based on individuals collected in Spain. B. A. erodiorum female, head in oblique

view. C. Andrena juliana Wood 2021, habitus in lateral view. The species was recently described in Wood et al. (2021) based on

individuals collected in Portugal and Spain. D. A. juliana female, head in oblique view. E. Andrena levante Wood & Praz 2021

male, habitus in lateral view. The species was recently described by Wood et al. (2021) based on individuals collected in Spain.

F. A. levante male, head in oblique view. Pictures by Paolo Rosa.

Andrena (Truncandrena) ghisbaini Wood, 2023

Andrena (Truncandrena) ghisbaini Wood, 2023c: 344. Holotype ♀; Spain: Málaga, PN Sierra de las Nieves, mountain peak S

of Pinsapo Escalereta (OÖLM).

Distribution. Spain.

Andrena (Ulandrena) graciliata Wood, 2023

Andrena (Ulandrena) graciliata Wood, 2023a: 51. Holotype ♂; Cyprus: Limassol, Yermasoyia [Germasogeia] (OÖLM).

Distribution. Cyprus.

Andrena (Euandrena) isolata Wood, 2023

Andrena (Euandrena) isolata Wood, 2023c: 339. Holotype ♀; Spain: Granada, Sierra Nevada, Trevélez to Refugio La Campiñuela

(OÖLM).

Distribution. Spain.

Andrena (Avandrena) juliae Wood, 2023

Andrena (Avandrena) juliae Wood, 2023c: 336. Holotype ♀; Cádiz, Parque Natural Los Alcornocales, Las Algamitas, Finca

Murtas (OÖLM).

Distribution. Spain.

Andrena (Notandrena) juliana Wood, 2021

(Figs 2C, D)

Andrena (Notandrena) juliana Wood in Wood et al., 2021: 174. Holotype ♀; Spain: Andalusia, Málaga, San Julián, 8 km SW

of Málaga (RMNH).

Distribution. Portugal, Spain.

Andrena (Simandrena) kocoureki Wood, 2021

Andrena (Simandrena) kocoureki Wood, 2021: 5. Holotype ♀; Bulgaria: Blagoevgrad, Sandanski (OÖLM).

Distribution. Bulgaria.

Andrena (Taeniandrena) laevicorpus Wood, 2023

Andrena (Taeniandrena) laevicorpus Wood, 2023a: 41. Holotype ♀; Cyprus: W of Polis [Polis Chrysochous], E of Cedar Valley

(OÖLM).

Distribution. Cyprus.

GHISBAIN, ROSA ET AL.

Andrena (Taeniandrena) levante Wood & Praz, 2021

(Figs 2E, F)

Andrena (Taeniandrena) levante Wood & Praz in Wood et al., 2021: 156. Holotype ♂; Spain: Valencia, 80 km SW of Valencia,

Reserva de Muela de Cortes (OÖLM).

Distribution. Spain.

Andrena (Taeniandrena) lusitania Wood & Ortiz-Sánchez, 2022

Andrena (Taeniandrena) lusitania Wood & Ortiz-Sánchez, 2022: 120. Holotype ♂; Portugal: Castelo Branco, Fundão, Vale Praz

(OÖLM).

Distribution. Portugal, Spain.

Andrena (Micrandrena) omnilaevis Wood, 2020

Andrena (Micrandrena) omnilaevis Wood in Wood et al., 2020a: 206. Holotype ♂; Portugal: Braga, Quinta do Confurco, Várzea

Cova (OÖLM).

Distribution. Portugal, Spain.

Andrena (Micrandrena) ortizi Wood, 2023

Andrena (Micrandrena) ortizi Wood, 2023c: 342. Holotype ♀; Spain: Sierra Nevada, Mirador Monte Ahí de Cara (OÖLM).

Distribution. Spain.

Andrena (Euandrena) pelagonia Wood, 2021

Andrena (Euandrena) pelagonia Wood, 2021: 14. Holotype ♀; North Macedonia: Mount Kožuf, Smrdliva Voda (OÖLM).

Distribution. North Macedonia.

Andrena (Micrandrena) pirinia Wood, 2021

Andrena (Micrandrena) pirinia Wood, 2021: 11. Holotype ♀; Bulgaria: Blagoevgrad, Popina Luka (OÖLM).

Distribution. Bulgaria.

Andrena (?Planiandrena) ramosa Wood, 2022

Andrena (?Planiandrena) ramosa Wood in Wood et al., 2022b: 8. Holotype ♀; Spain: Andalusia, Sevilla, Aznalcázar

(OÖLM).

Distribution. Spain.

Andrena (Taeniandrena) taedium Wood, 2023

Andrena (Taeniandrena) taedium Wood, 2023a: 44. Holotype ♀; Lebanon: Horch Ehden, Ain al Bayada Gate (OÖLM).

Distribution. Greece, Turkey, Lebanon, Iran.

Published synonymies

Andrena (Aciandrena) astrella Warncke, 1975

Synonymised with Andrena fulica Warncke, 1974 (Figs 3A, B), which is the senior synonym according to Wood et

al. (2020b: 66).

Andrena (incertae sedis) breviscopa Pérez, 1895

This taxon was misinterpreted by Warncke, and is a synonym of A. numida Lepeletier (Wood 2023c: 298).

Andrena (Andrena) bulgariensis Warncke, 1965

Synonymised with Andrena inconstans Morawitz, 1877, which is the senior synonym according to Wood (2023a:

58).

Andrena (Melandrena) creberrima Pérez, 1895

Synonymised with Andrena discors Erichson, 1841, which is the senior synonym according to Wood (2023c:

267).

Andrena (Micrandrena) curtula Pérez, 1895

Synonymised with A. spreta Pérez, 1895, which is the senior synonym according to Wood (2023c: 287).

Andrena (Truncandrena) espanola Warncke, 1967

Synonymised with A. nigropilosa Warncke, 1967, which is the senior synonym according to Wood (2023c: 306).

Andrena (Melandrena) gallica Schmiedeknecht, 1883

Synonymised with Andrena assimilis Radoszkowski, 1876 (Figs 3C, D), which is the senior synonym according to

Wood & Monfared (2022).

Andrena (Melandrena) hispania Warncke, 1967

Synonymised with Andrena morio Brullé, 1832, which is the senior synonym according to Wood (2023c: 263).

GHISBAIN, ROSA ET AL.

FIGURE 3. A. Andrena fulica Warncke, 1974 male, habitus in lateral view. The taxonomic review of Wood et al. (2020b) shows

that the taxon Andrena astrella Warncke, 1975 is a junior synonym of this species. B. A. fulica male, frontal view of the head.

C. Andrena assimilis Radoszkowski, 1876 female, habitus in lateral view. The taxonomic review of Wood & Monfared (2022)

shows that the taxon Andrena gallica Schmiedeknecht, 1883 is a junior synonym of this species. D. A. assimilis female, frontal

view of the head. E. Andrena bimaculata (Kirby, 1802) female, habitus in lateral view. Unpublished data from T.J. Wood agrees

with the baseline of Gusenleitner & Schwarz (2002) and indicates that the taxon Andrena oligotricha Mavromoustakis, 1952 is

a junior synonym of this species. F. A. bimaculata female, frontal view of the head. Pictures by Paolo Rosa.

Andrena (Euandrena) impressa Warncke, 1967

Synonymised with Andrena (Euandrena) lavandulae Pérez, 1902, which is the senior synonym according to Wood

(2023c: 261).

Andrena (Leimelissa) ispida Warncke, 1965

Synonymised with Andrena fallax Eversmann, 1852, which is the senior synonym according to Astafurova et al.

(2022).

Andrena (Avandrena) siciliana Warncke, 1980

Synonymised with Andrena heterodoxa Pérez, 1903, which is the senior synonym according to Wood (2023a: 57).

Andrena (Taeniandrena) similis Smith, 1847

Synonymised with Andrena russula Lepeletier, 1841, which is the senior synonym according to Praz et al. (2022).

Andrena (incertae sedis) toelgiana Friese, 1921

Synonymised with A. limbata Eversmann, 1852 which is the senior synonym according to Wood (2023: 325).

Andrena (Melandrena) vachali Pérez, 1895

Synonymised with Andrena discors Erichson, 1841, which is the senior synonym according to Wood (2023c:

267).

Synonymic notes

Andrena (Plastandrena) cypricola Mavromoustakis, 1952

Synonym of Andrena tibialis (Kirby, 1802). Andrena cypricola was listed at the species rank by Gusenleitner &

Schwarz (2002) and Varnava et al. (2020) but we do not consider this to be justified given the lack of variation in the

genital capsule. Moreover, given the problems of introgression within the subgenus Plastandrena, detailed genetic

work is needed before to be confident that additional taxa exist outside of the ‘core’ Plastandrena taxa.

Andrena (incertae sedis) iohannescaroli Nobile, 2000

In the original description (Nobile 2000), the name and location of the collection housing the holotype are omitted,

thereby not complying with Article 16.4.2 of the International Code of Zoological Nomenclature (ICZN, 1999).

Consequently, because this species is not correctly described, we consider the name a nomen nudum.

Andrena (Plastandrena) oligotricha Mavromoustakis, 1952

Andrena oligotricha was considered to be a valid species by Varnava et al. (2020). However, we consider it a

synonym of Andrena bimaculata (Kirby, 1802) (Figs 3E, F) based on genetic data (Wood 2023b), the lack of

GHISBAIN, ROSA ET AL.

morphological differentiation from A. bimaculata, particularly in the genital capsule, the presence of this red colour

form in Turkey and the Levant, and we thus return to the hypothesis proposed by Gusenleitner & Schwarz (2002).

Andrena (Simandrena) palumba Warncke, 1974

This taxon will be synonymised with Andrena rhypara Pérez, 1903 as part of a revision of the Moroccan fauna

conducted by Wood (in prep.).

Taxonomic changes

Cubiandrena Warncke, 1968

Andrena (Cubiandrena) Warncke, 1968: 72. Type species: Andrena cubiceps Friese, 1914, by original designation.

Cubiandrena: Dubitzky et al. 2010: 137. Elevated to genus rank and recently validated by Pisanty et al. (2022) by means of

molecular analyses.

Andrena (Micrandrena) acuta Warncke, 1968

Andrena wollastoni acuta Warncke, 1968: 77.

Andrena acuta: Kratochwil 2020: 171. Upgraded to species rank.

Andrena (Taeniandrena) afzeliella (Kirby, 1802)

Melitta afzeliella Kirby, 1802: 169.

Andrena ovatula sensu auctorum: Warncke 1967: 248. Synonymised.

Andrena afzeliella: Praz et al. 2022: 383. Resurrected to species status.

Andrena (Micrandrena) alma Warncke, 1975

Andrena mariana alma Warncke, 1975c: 299

Andrena alma: Wood 2023c: 276. Upgraded to species rank.

Andrena (Micrandrena) ampla Warncke, 1967

Andrena proxima ampla Warncke, 1967: 229. Holotype ♀, Spain: Alberche (OÖLM).

Andrena ampla: Schmid-Egger 2005: 1038. Upgraded to species rank (overlooked by Nieto et al. 2014 and Rasmont et al.

2017).

Andrena (Micrandrena) catula Warncke, 1968

Andrena wollastoni catula Warncke, 1968: 77.

Andrena catula: Kratochwil 2020: 180. Upgraded to species rank.

Andrena (Simandrena) cilissaeformis Pérez, 1895

Andrena cilssaeformis Pérez, 1895: 42.

Andrena numida Warncke, 1967: 186. Synonymised (incorrectly).

Andrena cilissaeformis Lepeletier, 1841: Wood 2023c: 298. Resurrected to species status.

Andrena (Euandrena) croatica Friese, 1887

Andrena croatica Friese, 1887: 85.

Andrena bicolor Fabricius, 1775: Warncke 1967: 264. Synonymised.

Andrena croatica: Praz et al. 2019: 31. Resurrected to species status.

Andrena (Taeniandrena) croceiventris Morawitz, 1871

Andrena croceiventris Morawitz, 1871: 219.

Andrena russula croceiventris: Warncke 1967: 264.

Andrena croceiventris: Praz et al. 2022: 396. Returned to species status.

Andrena (Taeniandrena) eversmanniana Osytshnjuk, 1994

Andrena fulva Eversmann, 1852: 31, nom. praeocc., nec Andrena fulva (Müller, 1776).

Andrena marginata Fabricius, 1776: Warncke 1967: 273. Synonymised (incorrectly).

Andrena eversmanniana Osytshnjuk, 1994: 35, repl. name for Andrena fulva Eversmann.

Andrena eversmanniana: Astafurova et al. 2022: 398. Resurrected to species status.

Andrena (Leimelissa) fallax Eversmann, 1852

Andrena fallax Eversmann, 1852: 20.

Andrena chrysosceles (Kirby, 1802): Warncke 1967: 269. Synonymised (incorrectly).

Andrena fallax: Astafurova et al. 2022: 396. Returned to species status.

Andrena ispida Warncke, 1965: Astafurova et al. 2022: 396. Synonymised.

Andrena (Micrandrena) gomerensis Warncke, 1993

Andrena wollastoni gomerensis Warncke, 1993: 762.

Andrena gomerensis: Kratochwil 2020: 190. Upgraded to species rank.

Andrena (Euandrena) lavandulae Pérez, 1902

Andrena lavandulae Pérez, 1902: 176

Andrena bicolor Fabricius, 1775: Warncke 1967: 283. Synonymised (incorrectly).

Andrena angustior impressa Warncke, 1967: 234.

Andrena impressa: Wood et al. 2021: 151. Upgraded to species rank.

Andrena lavandulae Wood 2023c: 260. Resurrected to species status.

Andrena (Micrandrena) lecana Warncke, 1975

Andrena niveata lecana Warncke, 1975c: 298.

Andrena lecana Wood 2023c: 283. Upgraded to species rank.

Andrena (Truncandrena) nigropilosa Warncke, 1967

Andrena truncatilabris nigropilosa Warncke, 1967: 225

Andrena truncatilabris espanola Warncke, 1967: 224

Andrena espanola Nieto et al. 2014: 44.

Andrena nigropilosa Wood 2023c: 306. Upgraded to species rank, including A. espanola as a junior subjective synonym.

GHISBAIN, ROSA ET AL.

Andrena (Plastandrena) nigrospina Thomson, 1872

Andrena nigrospina Thomson, 1872: 80.

Apis carbonaria Linnaeus, 1767, auct.

Andrena pilipes Fabricus, 1781: Gusenleitner & Schwarz 2002: 594.

Andrena nigrospina: Schmid-Egger & Patiny 1997: 37. Recognised as a valid taxon.

Notes. A two-taxon model based on the work of Schmid-Egger & Patiny (1997) is followed here with a bivoltine A.

pilipes that has a more southerly distribution and a univoltine A. nigrospina that has a more northerly distribution.

Additional detail is given in Wood (2023b; 2023c)

Andrena (Micrandrena) obsoleta Pérez, 1895

Andrena obsoleta Pérez, 1895: 44.

Andrena obsoleta auctorum: Warncke 1967: 186.

Andrena mariana solda Warncke, 1974: 40.

Andrena obsoleta Wood 2023c: 273. Recognised as a valid taxon.

Notes. This taxon was used in a sensu auctorum by Warncke; it is actually a valid species (Wood 2023c: 273), and

in a European context it is found only in Sicily.

Andrena (Taeniandrena) ovata Schenck, 1853

Andrena ovata Schenck, 1853: 133.

Andrena ovatula (Kirby, 1802): Warncke 1967: 295. Synonymised.

Andrena ovata: Praz et al. 2022: 399. Resurrected to species status.

Andrena (Pruinosandrena) parata Warncke, 1967

Andrena pruinosa parata Warncke, 1967: 233.

Andrena parata: Wood 2023c: 311. Upgraded to species status.

Andrena (Euandrena) pileata Warncke, 1975

Andrena allosa pileata Warncke, 1975a: 85.

Andrena pileata: Praz et al. 2019: 31. Upgraded to species rank.

Andrena (Suandrena) portosanctana Cockerell, 1922

Andrena portosanctana Cockerell, 1922: 32.

Andrena maderensis portosanctana Warncke, 1967: 208.

Andrena portosanctana: Kratochwil et al. 2014: 1540. Returned to species status (overlooked by Nieto et al. 2014 and Rasmont

et al. 2017).

Andrena (Taeniandrena) poupillieri Dours, 1872

Andrena poupillieri Dours, 1872: 430.

Andrena ovatula poupillieri Warncke 1967: 176.

Andrena poupillieri: Wood 2023c: 300. Returned to species status.

Andrena (Micrandrena) tenostra Warncke, 1975

Andrena mariana tenostra Warncke, 1975c: 300

Andrena tenostra: Wood 2023c: 273. Upgraded to species rank.

Andrena (Notandrena) varuga Warncke, 1975

Andrena varuga reperta Warncke, 1974: 48.

Andrena varuga Warncke, 1975c: 312.

Andrena reperta varuga Gusenleitner & Schwarz 2002: 633.

Andrena varuga: Wood 2023c: 292. Upgraded to species rank (distinct from A. reperta).

Andrena (Taeniandrena) vocifera Warncke, 1975

Andrena gelriae vocifera Warncke, 1975b: 136.

Andrena vocifera: Praz et al. 2022: 406. Upgraded to species rank.

Cubiandrena cubiceps (Friese, 1914)

Andrena cubiceps Friese, 1914: 223.

Cubiandrena cubiceps: Dubitzky et al. 2010: 144.

Taxonomic acts and clarifications

Andrena (Hoplandrena) carantonica Pérez, 1902

Considered to be a nomen dubium by Wood et al. (2022c).

Andrena (Notandrena) erythrocnemis Morawitz, 1870 sensu auctorum

This taxon was used incorrectly by Warncke (Gusenleitner & Schwarz 2002; Proshchalykin et al. 2017) as Andrena

erythrocnemis sensu auctorum, and the name is replaced in the European list by A. griseobalteata Dours, 1872.

Wood (2023c: 361) designated a neotype for A. griseobalteata from southern France.

Andrena (Truncandrena) oulskii Radoszkowski, 1867 sensu auctorum

Listed by Nieto et al. (2014). This identification was based on a misapplication of the type concept: the true Andrena

oulskii belongs to the subgenus Truncandrena, whereas European material is a member of the subgenus Ulandrena.

The type material of Andrena oulskii was clarified by Wood (2021); the correct name for European material is A.

(Ulandrena) biguttata Friese, 1923.

Andrena (Ulandrena) osychniukae Osytshnjuk, 1977 and Andrena (Ulandrena) polemediana Mavromoustakis,

1956

In the new checklist of the European species these two taxa are included within Andrena abbreviata Dours, 1873

sensu lato. A comprehensive molecular revision of these taxa is required for clarity due to ongoing taxonomic

confusion.

GHISBAIN, ROSA ET AL.

Andrena (Hoplandrena) scotica Perkins, 1916

This name replaces the use of Andrena carantonica sensu auctorum, Andrena carantonica Pérez, 1902 is treated as

a nomen dubium (see above).

Species recorded in Europe after 2017

Andrena (Notandrena) falcinella Warncke, 1975

Distribution. First recorded for Europe by Wood & Monfared (2022) from Limassol in Cyprus. Outside Europe

known from Turkey, Israel and Iran.

Andrena (incertae sedis) laurivora Warncke, 1974

Distribution. First recorded for Europe by Wood et al. (2021) from Huelva and Sevilla in Spain. Outside Europe

known only from Morocco.

Andrena (Avandrena) melacana Warncke, 1967

Distribution. First recorded for Europe by Wood & Ortiz-Sánchez (2022) from Albacete, Cádiz, Granada, and

Málaga in Spain. Outside Europe known from Morocco, Algeria, and Tunisia.

Andrena (Truncandrena) varia Pérez, 1895

Distribution. First recorded for Europe by Ortiz-Sánchez (2020) from Córdoba in Spain. Outside Europe known

from Morocco, Algeria, and Tunisia.

New species for Europe

Andrena (Notandrena) hebescens Wood, 2020

Distribution. New record (!) SPAIN: ♀, Canary Is., Fuerteventura, La Costilla, 12.iii.1935, (OÖLM). Outside

Europe known only from Morocco.

Andrena (Chrysandrena) henotica Warncke, 1975

Distribution. New record (!) CYPRUS: ♀, Nicosia, Kykkos, 800 m, 11.v.2014, leg. M. Kafka (OÖLM). Outside

Europe known from Turkey and Israel.

Species overlooked in the previous European checklists

Andrena (Suandrena) portosanctana Cockerell, 1922

Considered as a subspecies of Andrena maderensis by Warncke (1967), returned to species rank by Kratochwil et

al. (2014) but overlooked in the last update of Rasmont et al. (2017).

Species to be excluded from the European checklist

Andrena (Trucandrena) derbentina Morawitz, 1886

Distribution. The only verified records of this species are from the Caucasus.

Andrena (Melandrena) grandilabris Pérez, 1903

Distribution. This species is present only in eastern Turkey and Iran (see Wood & Monfared 2022). European

citations from Cyprus refer to misidentified material of Andrena elmaria Gusenleitner, 1998. Not present in

Europe.

Andrena (Euandrena) majalis Morawitz, 1876

Distribution. The only verified records of this species are from Central Asia.

Andrena (incertae sedis) wolfi Gusenleitner & Scheuchl, 2000

Distribution. Levant only, from Israel and to Syria and Jordan (Wood unpublished data). Not present in Europe.

Tribe Panurgini Leach, 1815

Genus described

Halopanurgus Wood, Patiny & Bossert, 2022

(Figs 4A, B)

Halopanurgus Wood, Patiny & Bossert in Wood et al. 2022a: 190. Type species: Camptopoeum baldocki Wood & Cross, 2017,

by original designation.

Species recently described as new to science

Halopanurgus baldocki (Wood & Cross, 2017)

(Figs 4A, B)

Camptopoeum baldocki Wood & Cross, 2017: 286. Holotype ♂; Portugal: Algarve, Cacela Velha (NHMUK).

Halopanurgus baldocki: Wood et al. 2022a: 190. New combination.

Distribution. Portugal, Spain.

Taxonomic changes

Halopanurgus fuzetus (Patiny, 1999)

(Figs 4C, D)

Flavipanurgus fuzetus Patiny, 1999: 58.

Halopanurgus fuzetus: Wood et al. 2022a: 194. New combination.

GHISBAIN, ROSA ET AL.

FIGURE 4. A. Halopanurgus baldocki (Wood & Cross, 2017) male, habitus in lateral view. The species was originally described

under the genus Camptopoeum Spinola, 1843, and then served as the type species for the newly described genus Halopanurgus

Wood, Patiny & Bossert, 2022 (Wood et al. 2022a). B. H. baldocki male, oblique view of the head. C. Halopanurgus fuzetus

(Patiny, 1999) female, habitus in lateral view. The species was originally described as Flavipanurgus fuzetus by Patiny 1999

but was later shown to belong to the genus Halopanurgus. D. H. fuzetus female, oblique view of the head. E. Flavipanurgus

kastiliensis (Warncke, 1987) male, habitus in lateral view. The species was initially considered to be a subspecies of Flavipanurgus

ibericus (Warncke, 1972) before being upgraded to species rank in Wood & Cross (2018). F. F. kastiliensis male, frontal view

of the head. Pictures by Paolo Rosa.

Flavipanurgus kastiliensis (Warncke, 1987)

(Figs 4E, F)

Panurgus ibericus kastiliensis Warncke, 1987: 86.

Flavipanurgus kastiliensis: Cross & Wood 2018: 564. Upgraded to species rank.

Species overlooked in the previous European checklists

Panurginus alticolus Morawitz, 1875

Distribution. Russia (Samara Prov., Volgograd Prov., Orenburg Prov., Bashkir Rep.) (Romankova & Astafurova

2011).

Family APIDAE Latreille, 1802

Tribe Ammobatini Handlirsch, 1925

Published synonymies

Ammobates globosus Mavromoustakis, 1954

Synonymised with Ammobates biastoides (Friese, 1895), which is the senior synonym according to Warncke (1983:

294). This change was overlooked in Nieto et al. (2014) and Rasmont et al. (2017).

Tribe Ancylaini Michener, 1944

Species to be excluded from the European checklist

Ancyla nitida Friese, 1922

Distribution. Given that we could not locate specimens to verify the old and doubtful records of this species from

Cyprus, we deleted this taxon from the checklist of the European species (Varnava et al. 2020).

Ancyla oraniensis Lepeletier, 1841

Distribution. North African species, not present in Europe.

Tribe Anthophorini Dahlbom, 1835

Published synonymies

Amegilla magnilabris (Fedtschenko, 1875)

In Nieto et al. (2014) as a valid species. This species was synonymised with Amegilla savignyi (Lepeletier, 1841) by

Brooks (1988), yet never revalidated. In the present list we follow the interpretation of the latter author until further

taxonomic work is done.

GHISBAIN, ROSA ET AL.

Anthophora salviae (Panzer, 1805)

Synonymised with Anthophora crinipes Smith, 1854, which is the valid name according to Maghni et al. (2017).

The latter authors and Scheuch & Willner (2016) considered the basionym Lasius salviae Panzer, 1805 a nomen

dubium.

Anthophora thomsonii (Saunders, 1882)

Synonymised with Anthophora atriceps Pérez, 1879, which is the senior synonym, according to Baldock et al.

(2018). This taxon was described as Podalirius thomsonii Saunders, 1882, but the name in use was thomsoni (with

only one –i), which is an incorrect subsequent spelling.

New synonymies

Anthophora senicula Pérez, 1902

We synonymise here Anthophora senicula Pérez, 1902, syn. nov. with Anthophora (Pyganthophora) balearica

(Friese, 1896), the name with priority. Type series were revised in MNHN and SMF, respectively.

Taxonomic acts and clarifications

Anthophora andalusica Pérez, 1902

Morphologically, the type specimen of this taxon (examined in MNHN) is an abraded female belonging to the

subgenus Pyganthophora Brooks, 1988 that may be synonymous with Anthophora retusa meridionalis Pérez, 1879.

However, given the degraded nature of the specimen, it is safer to regard A. andalusica a nomen dubium.

Anthophora cincrea (Friese, 1896)

This taxon was described by Friese as Podalirius cincreus. The name is a misspelling of the Latin adjective cinereus

[= grey], referring to the body hair coloration. An original incorrect spelling can be emended, but the emendation

would generate a secondary homonym of A. cinerea Eversmann, 1852, thus creating instability of the system.

We therefore prefer to keep the original misspelling cincreus, nevertheless the name is an adjective and must be

modified in accordance with the feminine genus gender of Anthophora. For this reason, in the new checklist we use

the name Anthophora cincrea (Friese, 1896).

Anthophora (Pyganthophora) erschowi Fedtschenko, 1875

The type series was revised in ZISP. The specimens of the series are only females, all belonging to the difficult

group of Anthophora aestivalis (Panzer, 1801), in which generally only males can be reliably identified. The name

Anthophora erschowi is therefore considered as a species inquirenda and removed from the present checklist.

Material under this name requires revision.

Anthophora (Pyganthophora) rubricrus Dours, 1869

The type series of Anthophora rubricrus from Greece was destroyed during the WWI bombing of the museum of

Amiens (France). Only one species is likely to correspond to A. rubricrus Dours which is found in Syros today, and

therefore corresponds with the type locality (J. Devalez, pers. comm.). However, without type material and reliably

identified specimens from the taxon author, we consider Anthophora rubricrus a species inquirenda and remove it

from the present checklist.

Anthophora (Paramegilla) segnis Eversmann, 1852

Eversmann (1852) gives as locus typicus “in prov. Orenburg. Australi, Saratoviensi et Astrachanensi”. Proshchalykin

et al. (2019), without revision of the type series, suggest a likely synonymy with Anthophora podagra Lepeletier, 1841.

However, after examination of the type series in IZKP, the species looks more related to Anthophora prshewalskyi

Morawitz, 1880. Anthophora segnis is here considered as the senior synonym of Anthophora prshewalskyi syn.

nov.

Anthophora (Pyganthophora) ventilabris Lepeletier, 1841

In Nieto et al. (2014) erroneously as Anthophora ventrilabris Lepeletier, 1841, an incorrect subsequent spelling.

Species recorded in Europe after 2017

Anthophora (Lophanthophora) cinerascens Lepeletier, 1841

Distribution. Russia (Orenburg Prov.) (Proshchalykin et al. 2019). Outside Europe known from north Africa, Israel,

Pakistan, Central Asia.

Anthophora (Paramegilla) segnis Eversmann, 1852

Distribution. Crimea. Outside Europe known from Turkey, Turkmenistan, Kazakhstan and China (Levchenko et

al. 2017).

New species for Europe

Anthophora (Paramegilla) balassogloi (Radoszkowski, 1877)

Distribution. New record (!) RUSSIA: 3♂ and 5♀, Bashkortostan (Adzanov), 3.vii.1955, leg. Nikiforuk (ZISP);

7♀, idem, 8.vii.1956; 1♀, idem 9.vii.1956; 2♀, idem, 10.vii.1956; 5♂ and 1♀, idem, 12.vii.1956; 6♂ and 1♀, idem,

13.vii.1956; 1♂ and 2♀, Isyangulovo, 10.vii.1956, leg. Nikiforuk (ZISP); 1♂, idem 13.vii.1956. Outside Europe

known from the Caucasus, Ural Mountains, Turkey, Armenia (Ascher & Pickering 2022).

Anthophora (Lophanthophora) crysocnemis Morawitz, 1877

Distribution. New record (!) RUSSIA: 1♀, Sarepta [=Volgograd], leg. Bakker (ZISP). Outside Europe known

from Armenia and Kazakhstan (!): 1♀, Yanvartsevo, 21.v.1949, leg. Rudolf (ZISP).

Remarks. Brooks (1988) erroneously classified this species into the subgenus Anthomegilla Marikovskaya,

1976. This species undoubtedly belongs to the subgenus Lophanthophora Brooks, 1988.

GHISBAIN, ROSA ET AL.

Anthophora (incertae sedis) raddei Morawitz, 1875

Distribution. New record (!) BULGARIA: 1♂ and 4♀, Plovdiv, 6–20.vi.1909, leg. A. Gutbier (ZISP). Outside

Europe known from Armenia (Morawitz 1875) and Iran (Alfken 1935).

Remarks. This species was classified by Brooks (1988) in the subgenus Paramegilla Friese, 1897 but the

species is difficult to classify in the current system.

Species to be excluded from the European checklist

Anthophora (Anthophora) lanata (Klug, 1845)

Remarks. Listed by Nieto et al. (2014), but the status of this species is doubtful, and it might be a subspecies of A.

(Anthophora) canescens Brullé, 1832. Waiting for a proper taxonomic revision, we choose to exclude it from the

present checklist and to consider it a species inquirenda.

Tribe Melectini Westwood, 1839

Taxonomic changes

Melecta baerii (Radoszkowski, 1865)

Melecta baerii (Radoszkowski, 1865) was previously misspelt as Melecta baeri (Radoszkowski, 1865) in the Red

List of Nieto et al. (2014).

Thyreus aberrans (Morawitz, 1875)

This taxon has been confused, as the location of the type material is uncertain. According to Proshchalykin (pers.

comm.) the type could be located in Moscow. The species was described from Uzbekistan as Crocisa aberrans, and

its identity is unclear. Lieftinck (1968) notes that in the original description, the scutellum has the posterior margin

truncate, with the apex produced into a median lobe. This does not fit any known Thyreus species, and is reminiscent

of a Dioxyine bee. Without a type, the concept is unclear, and the name must be treated as a nomen dubium. Records

from the European part of Russia must therefore be considered to be unclear due to this taxonomic uncertainty. We

therefore remove this taxon from the European checklist.

Thyreus piceus (Meyer, 1921)

This taxon is known only from the type specimen which is a female collected from the island of Poros in Greece.

The type appears to be lost, as it cannot be located in the Berlin collection (Lieftinck 1968). Without a type, this

name must be considered a nomen dubium given the impossibility to conclusively conclude on its identity based

only on the description.

Thyreus plumatus (Meyer, 1921)

This taxon is known only from the type specimen which is a male collected from Milan in Italy. The type appears to

be lost, as it cannot be located in the Berlin collection (Lieftinck 1968). Without a type, this name must be considered

a nomen dubium given the impossibility to conclusively conclude on its identity based only on the description.

Species recorded in Europe after 2017

Melecta amanda Lieftinck, 1980

Distribution. Recorded for the first time by Levchenko et al. (2017) from the south of the European part of Russia.

Outside Europe known from Iran.

Species overlooked in the previous European checklists

Melecta alcestis Lieftinck, 1980

Distribution. Described from the European part of Russia (Orenburg). Only known from the type specimens

(unpublished).

Melecta diacantha Eversmann, 1852

Distribution. Described from the European part of Russia (Urals) (Levchenko et al. 2017).

Melecta eversmanni Radoszkowski, 1893

Distribution. Described from the European part of Russia (Orenburg) (Radoszkowski 1893; Proshchalykin et al.

2019). Outside Europe known from Uzbekistan.

Melecta rutenica Radoszkowski, 1893

Distribution. The type locality given by Radoszkowski (1893) is Ciechocinek, Poland. However, as discussed by

Lieftinck (1980), this is an error. The lectotype specimens is labelled as “Nickon: p. Stani” which is an unclear

locality. However, the specimen is labelled by Radoszkowski and was considered to be a valid lectotype by Lieftinck

(1980). In Europe, the species has only confidently been recorded from Ukraine (Kirill ravine, Kiev).

Species to be excluded from the European checklist

Thyreus tricuspis (Pérez, 1883)

Listed by Nieto et al. (2014) for Europe, but only present in north Africa.

Tribe Bombini Latreille, 1802

Species recently described as new to science

Bombus (Melanobombus) bisiculus Lecocq, Biella, Martinet & Rasmont, 2019

Bombus (Melanobombus) bisiculus Lecocq, Biella, Martinet & Rasmont, 2019: 7. Holotype ♂; Italy: Sicily, Isnello

(UMONS).

Individuals of the taxon described recently with the name Bombus bisiculus have been recorded traditionally as B.

lapidarius decipiens Pérez, 1879, because individuals of the new taxon share a yellow-banded colour pattern that

GHISBAIN, ROSA ET AL.

has been given by some the status of a subspecies within B. lapidarius (Linnaeus, 1758). But based on its divergence

from Bombus lapidarius in northern Europe in COI barcodes and CLGS (cephalic labial gland secretions, believed

to function as sex-specific pheromones in bumblebees), Lecocq et al. (2019) described the new taxon from southern

Italy and Sicily as a separate species, B. bisiculus Lecocq, Biella, Martinet & Rasmont, 2019 (the name decipiens

Pérez was applied originally to yellow-banded individuals of B. lapidarius from the Pyrenees and the Iberian

Peninsula).

In a revision of the Melanobombus bumblebees world-wide, Williams et al. (2020) noted from a re-analysis of

COI barcodes based in part on sequences provided by Lecocq et al. (2015), that some individuals (‘lapidarius SE

Europe’) were grouped in the tree together with the new taxon Bombus bisiculus. This group of individuals was not

explicitly reported or discussed by Lecocq et al. (2019) when they formally described B. bisiculus. These SE Europe

sequences came from unbanded specimens from Slovakia, the Czech Republic, Serbia, Bulgaria, and Hungary that

are phenotypically identical to the typical B. lapidarius (Linnaeus, 1758), not to the yellow-banded taxon bisiculus

of Lecocq et al. (2019).

More significantly, Williams et al. (2020) highlighted that the SE Europe samples also produce CLGS mixtures

(as reported by Lecocq et al. 2015), that are typical of B. lapidarius, not of B. bisiculus. Williams et al. (2020)

therefore suggested that the definition of Bombus bisiculus by Lecocq et al. (2019) as a yellow-banded bumblebee

species diverging in both COI barcodes and CLGS from B. lapidarius appears to be contradicted by the dataset of

Lecocq et al. (2015) when the whole of Europe is considered. Williams et al. (2020) therefore concluded that there

is disagreement between the two independent lines of evidence (COI and CLGS) from Lecocq et al. (2013, 2015,

2019), not the corroboration that would be required to support species status for the taxon bisiculus: the SE Europe

bees have the COI of bisiculus but the CLGS of lapidarius.

In the absence of corroboration between COI and CLGS (a required combination for deserving a species status

following Lecocq et al. 2013, 2015, 2019), we here consider the taxon bisiculus as a subspecies of B. lapidarius.

Further genetic sequencing and semio-chemical data from Eastern Europe and the Balkans are required to understand

more in detail the evolutionary history of these lineages.

Distribution. Southern Italy. Records from Bulgaria, the Czech Republic, Hungary, Serbia, and Slovakia need

confirmation.

Taxonomic changes

Bombus (Megabombus) reinigiellus (Rasmont, 1983)

Megabombus reinigiellus Rasmont, 1983: 43.

Bombus hortorum reinigiellus: Ghisbain et al. 2021b: 8. Downgraded to subspecies rank.

The taxon reinigiellus Rasmont, 1983 was re-assessed as a subspecies of Bombus hortorum (Linnaeus, 1761) by

Ghisbain et al. (2021b) based on genetic and semio-chemical analyses.

Bombus (Pyrobombus) konradini Reinig, 1965

Bombus lapponicus konradini Reinig, 1965:105.

Bombus konradini: Martinet et al. 2018a: 205. Upgraded to species rank.

The taxon konradini Reinig, 1965 was re-assessed as a valid species by Martinet et al. (2018a) based on genetic and

semio-chemical analyses.

Bombus (Thoracobombus) mocsaryi Kriechbaumer, 1877

(Figs 5A, B)

Bombus mocsaryi Kriechbaumer, 1877: 253.

Bombus laesus mocsaryi: Brasero et al. 2021: 10. Downgraded to subspecies rank.

The taxon mocsaryi Kriechbaumer, 1877 (Figs 5A, B) was re-assessed as a subspecies of Bombus laesus Morawitz

(1875) (Figs 5C, D) by Brasero et al. (2021) based on genetic and semio-chemical analyses.

FIGURE 5. A. Bombus laesus mocsaryi Kriechbaumer, 1877 female, habitus in lateral view. The taxon mocsaryi was recently

shown to be conspecific with the taxon laesus Morawitz, 1875 by Brasero et al. (2021). B. B. laesus mocsaryi female, head in

oblique view. C. Bombus laesus laesus female, habitus in lateral view. D. B. laesus laesus female, head in oblique view. Pictures

by Paolo Rosa.

Taxonomic acts and clarifications

Bombus (Alpinobombus) polaris Curtis, 1835 and Bombus (Alpinobombus) pyrrhopygus Friese, 1902

There is an ongoing debate as to whether the Nearctic taxon polaris Curtis, 1835 and Palaearctic taxon pyrrhopygus

Friese, 1902 should be considered as conspecific or heterospecific.

Based on a lack of statistical differentiation in CLGS, Martinet et al. (2018b) consider these taxa as conspecific,

grouping them under the oldest available name Bombus (Alpinobombus) polaris Curtis, 1835. These authors also

base their argument on the fact that a previous study by Williams et al. (2015) showed no differentiation in the

slowly-evolving PEPCK nuclear gene between these two taxa (despite showing a significant differenciation based

on COI).

However, a detailed revision of the world Alpinobombus Skorikov, 1914 species by Williams et al. (2019)

concluded that Bombus polaris Curtis, 1835 and Bombus pyrrhopygus Friese, 1902 could be considered as two

differentiated species. The decision of Williams et al. (2019) is based on the following evidence: (i) a species

coalescent in the COI marker demonstrated by the Poisson-tree-process procedure (cf. Zhang et al. 2013); (ii)

diagnostic differences in the 16S gene; (iii) differences in morphology and colour patters between both taxa.

Furthermore, according to Williams et al. (2019), the lack of statistical differentiation in CLGS cannot be used as a

GHISBAIN, ROSA ET AL.

convincing argument of conspecificity, as the absence of co-occurrence between both taxa might have removed any

selective pressure that might otherwise have enhanced barriers to interbreeding by driving evolutionary divergence

in sex pheromones. Williams et al. (2019) also note that a significant difference in CLGS would not be required to

consider both taxa as conspecific following the unified species concept of de Queiroz (2007) because (i) the status

of both taxa as independently evolving lineages (EILs) is automatically maintained by the wide sea barrier and (ii)

the existence of these two separate EILs is directly evidenced by their two species’ coalescents in the COI gene.

Here we follow the latter interpretation and consider the Nearctic taxon polaris Curtis, 1835 and Palaearctic taxon

pyrrhopygus Friese, 1902 as distinct species, with only Bombus pyrrhopygus occurring in IUCN Europe. Additional

work is required to better understand the evolutionary history of this highly interesting species complex.

Bombus (Thoracobombus) muscorum pereziellus (Skorikov, 1922)

(Figs 6A, B)

Agrobombus pereziellus Skorikov, 1922: 150.

Bombus muscorum pereziellus: Lecocq et al. 2014: 243. Downgraded to subspecies rank.

The taxon pereziellus (Figs 6A, B) was re-assessed as a subspecies of Bombus muscorum (Linnaeus, 1758) (Figs

6C, D) by Lecocq et al. (2014) based on genetic and semio-chemical analyses. This taxonomic update was omitted

in Rasmont et al. (2017).

FIGURE 6. A. Bombus muscorum pereziellus Skorikov, 1922 female, habitus in lateral view. The taxon pereziellus was shown

to be conspecific with the taxon muscorum (Linnaeus, 1758) by Lecocq et al. (2014). This taxonomic update was omitted in

Rasmont et al. (2017). B. B. muscorum pereziellus female, head in oblique view. C. Bombus muscorum pereziellus female,

habitus in lateral view. D. B. muscorum pereziellus female, head in oblique view. Pictures by Paolo Rosa.

Bombus (Psithyrus) vestalis perezi (Schulthess-Rechberg, 1886)

(Figs 7A, B)

Psithyrus perezi Schulthess-Rechberg, 1886: 275.

Bombus vestalis perezi: Lecocq et al. 2014: 243. Downgraded to subspecies rank.

The taxon perezi (Figs 7A, B) was re-assessed as a subspecies of Bombus vestalis Geoffroy (1785) (Figs 7C, D) by

Lecocq et al. (2014) based on genetic and semio-chemical analyses. This taxonomic update was omitted in Rasmont

et al. (2017).

FIGURE 7. A. Bombus vestalis perezi Schulthess-Rechberg, 1886 female, habitus in lateral view. The taxon perezi was shown

to be conspecific with the taxon vestalis Geoffroy, 1785 by Lecocq et al. (2014). This taxonomic update was omitted in Rasmont

et al. (2017). B. B. vestalis perezi female, head in oblique view. C. Bombus vestalis vestalis female, habitus in lateral view. D.

B. vestalis vestalis female, head in oblique view. Pictures by Paolo Rosa.

Tribe Ceratinini Latreille, 1802

Species to be excluded from the European checklist

Ceratina (Euceratina) zwakhalsi Terzo & Rasmont, 1998

Remarks. Listed by Nieto et al. (2014) yet only recorded from SW-Turkey. This species is therefore removed from

the present list.

GHISBAIN, ROSA ET AL.

Tribe Epeolini Linsley & Michener, 1939

Species recently described as new to science

Epeolus ibericus Bogusch, 2018

(Figs 8A, B)

Epeolus ibericus Bogusch in Bogusch & Hadrava, 2018: 28. Holotype ♀; Portugal: Caparica, 18.vii.1982, leg. K. Guichard

(NHMUK).

Distribution. Portugal and Spain (Bogusch & Hadrava 2018). Outside Europe known from Morocco (Bogusch

2021).

FIGURE 8. A. Epeolus ibericus Bogusch, 2018 female, habitus in dorsal view. The species was recently described by Bogusch

in Bogusch & Hadrava (2018). In Europe, it is found in Portugal and Spain. B. E. ibericus male, habitus in dorsal view. Pictures

by Petr Bogusch.

Taxonomic acts and clarifications

Epeolus julliani Pérez, 1884

(Figs 9A, B)

This taxon was recently synonymised with Epeolus transitorius Eversmann, 1852 by Bogusch & Hadrava (2018).

However, it is now recognized as a distinct species (Le Divelec 2021b; Astafurova & Proshchalykin 2022). Both

Epeolus julliani and E. transitorius occur in Europe.

FIGURE 9. A. Epeolus julliani Pérez, 1884 female, habitus in lateral view. The taxon had been synonymised with Epeolus

transitorius Eversmann, 1852 by Bogusch & Hadrava (2018) but is now recognized as a distinct species based on the work of

Le Divelec (2021b). B. E. julliani female, head in oblique view. Pictures by Paolo Rosa.

Epeolus minutus Radoszkowski, 1888

This name is here considered to be a nomen dubium as no type specimens are preserved, and no further specimens

have been attributed to this species (Bogusch & Hadrava 2018).

Species recorded in Europe after 2017

Epeolus bischoffi (Mavromoustakis, 1954)

Distribution. First recorded for Europe by Bogusch & Hadrava (2018) from Cyprus. Outside Europe known from

Turkey, Syria, Lebanon, Israel and Jordan.

Tribe Eucerini Latreille, 1802

Species recently described as new to science

Eucera (Eucera) dafnii Dorchin, 2019

Eucera (Eucera) dafnii Dorchin, 2019: 465. Holotype ♂; Israel: Tel Yizhaq south NR (SMNH).

Distribution. Bulgaria, Greece, North Macedonia. Outside Europe known from Iran, Israel, Palestine, Syria and

Turkey (Dorchin 2019).

Tetralonia gennargentui (Nobile, Catania & Bella, 2021)

Eucera (Tetralonia) gennargentui Nobile, Catania & Bella in Catania et al. 2021: 5. Holotype ♀; Italy: Sardinia, Nuoro, Fonni,

Gennargentu Massif, Bruncu Spina (UCSI).

Distribution. Italy (Sardinia).

GHISBAIN, ROSA ET AL.

Published synonymies

Eucera (Synhalonia) alternans (Brullé, 1832)

Eucera rufa (Lepeletier, 1841), which is the junior synonym, is retained by Dorchin (2023) as the valid name for

this species under the principle of name stability.

Eucera (Synhalonia) cressa (Tkalců, 1984)

Synonymised with Eucera tricincta Erichson, 1835, which is the senior synonym according to Kuhlmann et al.

(2022).

Eucera (Eucera) decolorata Gribodo, 1924

Synonymised with Eucera confinis Pérez, 1895, which is the senior synonym according to Dorchin (2023).

Eucera (Eucera) eucnemidea Dours, 1873

Synonymised with Eucera grisea Fabricius, 1793, which is the senior synonym according to Dorchin (2023).

Eucera (Synhalonia) fedtschenkoi Dalla Torre, 1896

Synonymised with Eucera intermedia (Morawitz, 1875), which is the original name (nec Melissodes intermedia

Cresson, 1872 (= Eucera belfragei (Cresson 1872), synonymy in: Ascher & Pickering (2022)), listed as Eucera in

Dalla Torre (1896)).

Eucera (Eucera) graeca Radoszkowski, 1876

Synonymised with Eucera proxima Morawitz, 1875, which is the senior synonym according to Dorchin (2023).

Eucera (Eucera) hispaliensis Pérez, 1902

Synonymised with Eucera longicornis (Linnaeus, 1758), which is the senior synonym according to Dorchin

(2023).

Eucera (Synhalonia) lucasi Gribodo, 1893

Synonymised with Eucera obscura (Brullé, 1832) (= Macrocera obscura Brullé, 1832), which is the senior synonym

according to Dorchin (2023).

Eucera (Eucera) maxima Tkalců, 1987

Synonymised with Eucera taurea Vachal, 1907, which is the senior synonym according to Dorchin (2023).

Eucera (Eucera) obsoleta Pérez, 1910

Synonymised with Eucera terminata Pérez, 1895, which is the senior synonym according to Dorchin (2023).

Eucera (Synhalonia) radoszkovskyi (Morawitz, 1872)

Synonymised with Eucera alborufa (Radoszkowski, 1871), which is the senior synonym according to Augul

(2018).

Eucera (Synhalonia) zeta Dalla Torre, 1896

Synonymised with Eucera melectoides (Radoszkowski, 1893), which is the senior synonym [= Macrocera melectoides

Radoszkowski, 1893, nec Tetralonia melectoides Smith, 1879 = Florilegus melectoides (Smith, 1879)].

Taxonomic changes

Based on Dorchin (2023) and Freitas et al. (in press), the following name changes are proposed in Dorchin (2023):

Tetralonia Spinola, 1838 is re-established as genus, including Tetraloniella Ashmead, 1899 (Dorchin et al. 2018),

Cubitalia Friese, 1911 is established as subgenus of Eucera Scopoli, 1770 and Synhalonia Patton, 1879 is retained

as subgenus of Eucera as in Michener (2000). Therefore, the following 19 combinations established for species

previously included in the genus Tetraloniella Ashmead, 1899 (Nieto et al. 2014) are now transferred to the genus

Tetralonia Spinola, 1839.

Tetralonia alticincta (Lepeletier, 1841), T. cinctella (Saunders, 1908), T. dentata (Germar, 1839), T. fulvescens

Giraud, 1863, T. glauca (Fabricius, 1775), T. graja (Eversmann, 1852), T. hohmanni Tkalců, 1993, T. iberica

Dusmet y Alonso, 1926, T. inulae Tkalců, 1979, T. julliani (Pérez, 1879), T. lanzarotensis Tkalců, 1993, T. lyncea

Mocsáry, 1879, T. nana Morawitz, 1873, T. pollinosa (Lepeletier, 1841), T. ruficornis (Fabricius, 1804), T. salicariae

(Lepeletier, 1841), T. scabiosae Mocsáry, 1881, T. strigata (Lepeletier, 1841), and T. vicina Morawitz, 1875.

Eucera (Eucera) pollinosa Smith, 1854

This species was previously referred to as Eucera chrysopyga Pérez, 1879 (Nieto et al. 2014), as when Eucera and

Tetralonia were treated as a single genus Eucera pollinosa Smith became a junior homonym of Eucera pollinosa

Lepeletier, 1841. Now that Tetralonia is restored as a genus, Eucera pollinosa Lepeletier is moved to Tetralonia,

and Eucera pollinosa Smith is no longer a junior homonym and becomes the senior synonym of Eucera chrysopyga

Pérez. Eucera pollinosa Smith was made a nomen protectum by Dorchin (2023).

Eucera (Eucera) palaestinae Friese, 1922

This taxon, originally proposed in the combination Eucera notata var. palaestinae Friese, 1922, was previously

misspelt as Eucera palestinae Friese, 1922.

Eucera (Synhalonia) ruficollis (Brullé, 1832)

This name was resurrected from synonymy with Eucera alternans (Brullé, 1832) after nearly 200 years (Dorchin

2023), the latter which itself was incorrectly interpreted (see above).

GHISBAIN, ROSA ET AL.

Species recorded in Europe after 2017

Eucera (Eucera) aequata Vachal, 1907

(Fig. 10)

Distribution. First recorded for Europe by Dorchin (2019) from Cyprus (Akrotiri and Limassol). Outside Europe

known from Turkey, Syria, Israel and Palestine (Dorchin 2019).

FIGURE 10. A. Eucera aequata Vachal, 1907 female, habitus in lateral view. The species was newly recorded for Europe (in

Cyprus) by Dorchin et al. (2019). B. E. aequata female, frontal view of the head. C. Eucera aequata male, habitus in lateral

view. D. E. aequata male, frontal view of the head. Pictures by Paolo Rosa.

Eucera (Cubitalia) breviceps Friese, 1911

Distribution. Listed as Eucera aff. breviceps Friese, 1911 in the key to Eucerini species of France in Aubert (2020).

Reported from northern Italy and south-eastern France by M. Aubert (pers. comm.). Outside Europe known from

Turkey, Syria, Georgia.

Eucera (Eucera) ferghanica Morawitz, 1875

Distribution. Recorded for the European part of Russia (south of European part; Levchenko et al. 2017). Outside

Europe known from Iran and Uzbekistan.

Eucera (Eucera) punctatissima Pérez, 1895

This name was incorrectly interpreted as synonymous with the unrelated species Eucera impressiventris Pérez, 1895

from north-western Africa (Dorchin 2023). It was recently found in southern Portugal by Thomas Wood (reported

as E. decolorata by Baldock et al. 2018).

Eucera (Cubitalia) tristis Morawitz, 1875

Distribution. Eucera tristis Morawitz, 1875 as Cubitalia tristis was recorded from Crimea (Levchenko et al. 2017).

Outside Europe it is known from Russia (Dagestan), Georgia and Turkey.

Species overlooked in the previous European checklists

Eucera (Eucera) atriceps Morawitz, 1877

The taxonomic status of Eucera atriceps is unclear. Further work is needed to evaluate its relationships with E.

nigripes Morawitz.

Distribution. Eucera atriceps Morawitz, 1877 was recorded from Crimea (Friese 1896; Sitdikov & Pesenko

1988; Levchenko et al. 2017). Outside Europe it is known from Armenia and Kazakhstan.

Species to be excluded from the European checklist

Eucera commixta Dalla Torre & Friese, 1895

Eucera commixta Dalla Torre & Friese (nomen novum proposed for Tetralonia nigrifacies Dours, 1873 nec Eucera

nigrifacies Lepeletier, 1841), listed as such by Nieto et al. (2014), was originally described from Algeria and south

France. The type series is considered to be lost and specimens described from France are likely not conspecific

(Dorchin, 2023).

Distribution. This taxon occurs in Algeria and other reported localities are doubtful due to uncertain identification

and confusion with similar species.

Eucera (Synhalonia) distinguenda (Morawitz, 1875)

Distribution. The data records from Romania (Iuga 1958) are doubtful as the species is only known from dry

environments from central Asia.

Eucera (Eucera) nigripes Klug, 1845

Listed from Greece in https://westpalbees.myspecies.info and from Italy in www.discoverlife.it, but according to S.

Risch (pers. comm.) it does not occur there.

Distribution. Turkey, Azerbaijan, Lebanon, Israel and Palestine.

Eucera (Eucera) sogdiana Morawitz, 1875

Distribution. No verified occurrence data of this species in Europe. The nearest record is from central Anatolia.

GHISBAIN, ROSA ET AL.

Eucera (Synhalonia) spectabilis (Morawitz, 1875)

Available records from Europe are probably erroneous.

Distribution. Turkey, Georgia, Uzbekistan, Kazakhstan, Kyrgyzstan, Iran, Pakistan.

Tribe Nomadini Latreille, 1802

Species recently described as new to science

Nomada achaica Schwarz & Smit, 2020

Nomada achaica Schwarz & Smit, 2020: 684. Holotype ♀; Greece: Peloponnese, Chelmos Mt., Piste W Avgo, ± 1900 m,

4.vi.2008, leg. A.W. Ebmer (MSPC).

Distribution. Greece (Peloponnese).

Nomada acutispina Schwarz & Smit, 2018

Nomada acutispina Schwarz & Smit in Smit, 2018: 106. Holotype ♀; Turkey: Antalya, 4.vi.1985, leg. M. Schwarz (MSPC).

Distribution. Greece (Crete). Outside Europe known from Turkey and Israel (Smit 2018).

Nomada aeginaica Schwarz & Smit, 2018

Nomada aeginaica Schwarz & Smit in Smit, 2018: 108. Holotype ♀; Greece: Peloponnese, Chelmos Mt., 1900 m, 2.vi.1962,

leg. M. Schwarz (MSPC).

Distribution. Greece (Peloponnese). Outside Europe known from Armenia and Turkey (Smit 2018).

Nomada breviceps Schwarz, Smit & Ockermüller, 2019

Nomada breviceps Schwarz, Smit & Ockermüller in Schwarz et al., 2019: 6. Holotype ♀; Graecia: Peloponnese, Trikala,

17.iv.1963, leg. Kl. Warncke (MSPC).

Distribution. Greece (Peloponnese). Outside Europe known from Turkey (Schwarz et al. 2019).

Nomada breviscapa Schwarz & Smit, 2018

Nomada breviscapa Schwarz & Smit in Smit, 2018: 122. Holotype ♂; Greece: Samos, Kerkis SE-Anstieg, 1200 m, 20.iv.1999,

leg. A.W. Ebmer (MSPC).

Distribution. Greece (Samos) (Smit 2018).

Nomada crenulata Schwarz & Smit, 2018

Nomada crenulata Schwarz & Smit in Smit, 2018: 132. Holotype ♀; Turkey: Konya, 27.v.1965, leg. M. Schwarz (MSPC).

Distribution. Greece (Alexandropolis) (Smit 2018) and Limnos (leg. Devalez 2012). Outside Europe known from

Azerbaijan and Turkey.

Nomada ebmeri Schwarz & Smit, 2018

Nomada ebmeri Schwarz & Smit in Smit, 2018: 140. Holotype ♂; Greece: Phocis, Delphi, 12.iv.1963, leg, W. Grünwaldt

(MSPC).

Distribution. Greece. Outside Europe known from Turkey and Syria (Smit 2018).

Nomada elsei Schwarz & Smit, 2018

Nomada elsei Schwarz & Smit in Smit, 2018: 142. Holotype ♀; Spain: Andalusia, Vélez Blanco, 3.vi.72, leg. W. Gross

(MSPC).

Distribution. Spain.

Nomada filicornis Schwarz & Smit, 2018

Nomada filicornis Schwarz & Smit in Smit, 2018: 149. Holotype ♀; Turkey: Antalya, 7.vi.1965, leg. M. Schwarz (MSPC).

Distribution. Italy (Sicily), Greece (Crete), Cyprus. Outside Europe known from Turkey, Syria and Jordan (Smit

2018).

Nomada gageae Schwarz & Smit, 2018

Nomada gageae Schwarz & Smit in Smit, 2018: 158. Holotype ♀; Cyprus: Limassol, 25.i.1948, leg. Mavromoustakis

(MSPC).

Distribution. Cyprus.

Nomada halophila Wood, 2022

Nomada halophila Wood, 2022: 5. Holotype ♀; Spain: Andalusia, Málaga, Campillos, Laguna Dulce, 4.vi.2021, leg. T. J. Wood

(OÖLM).

Distribution. Spain.

Nomada lapillula Schwarz & Smit, 2018

Nomada lapillula Schwarz & Smit in Smit, 2018: 174. Holotype ♂; Greece: Thessaly, Larissa, 15.v.1962, leg. Kl. Warncke

(MSPC).

Distribution. Greece. Outside Europe known from Turkey and Israel (Smit 2018).

Nomada legoffi Dufrêne, 2021

Nomada legoffi Dufrêne, 2021: 438. Holotype ♀; France: Corsica, Saint-Julien valley, east of Bonifacio, 50 m, 7.x.1900, C.

Ferton (MNHN).

Distribution. France (Corsica).

GHISBAIN, ROSA ET AL.

Nomada luteipes Schwarz & Smit, 2018

Nomada luteipes Schwarz & Smit in Smit, 2018: 181. Holotype ♀; Turkey: Hakkari, Tanin-Pass, 2300–2600 m, 3.vi.1980, leg.

M. Schwarz (MSPC).

Distribution. Bulgaria and Greece (Crete). Outside Europe known from Turkey and Iran (Smit 2018).

Nomada maxschwarzi Smit, 2018

Nomada maxschwarzi Smit, 2018: 185. Holotype ♀; Greece: Lesvos, Agiassos, 600–700 m, 5.vi.1992, leg. J.P. Duffels

(RMNH).

Distribution. Greece (Lesvos).

Nomada montarco Álvarez Fidalgo, 2023

Nomada montarco Álvarez Fidalgo, 2023: 9. Holotype ♀; Spain: Montarco, 15.vi.1924, leg. P. Álvarez Fidalgo (MNCN).

Distribution. Spain (Montarco).

Nomada nigrifrons Schwarz & Smit, 2018

Nomada nigrifrons Schwarz & Smit in Smit, 2018: 193. Holotype ♀; Turkey: Hakkari, Suvari-Halil-Pass, 2500 m, 2.vi.1980,

leg. M. Schwarz (MSPC).

Distribution. Greece (Lesvos) (Smit 2018). Outside Europe known from Turkey and Israel.

Nomada nigrilabris Schwarz & Smit, 2018

Nomada nigrilabris Schwarz & Smit in Smit, 2018: 194. Holotype ♀; Turkey: Mersin, Namrun, 14.v.1967, leg. F. Resse

(MSPC).

Distribution. Greece. Outside Europe known from Turkey and Israel (Smit 2018).

Nomada opaciformis Schwarz & Smit, 2018

Nomada opaciformis Schwarz & Smit in Smit, 2018: 200. Holotype ♀; Turkey: Tuncell, 13.vi.1973, leg. Kl. Warncke

(MSPC).

Distribution. Greece (Rhodes). Outside Europe known from Turkey and Israel (Smit 2018).

Nomada ottomanensis Schwarz & Smit, 2018

Nomada ottomanensis Schwarz & Smit in Smit, 2018: 203. Holotype ♀; Turkey: Mersin, Namrun, 10.v.–3.vi.1963, leg. F.

Schubert (MSPC).

Distribution. Greece (Lesvos, Rhodes). Outside Europe known from Turkey and Israel (Smit 2018).

Nomada pilosa Schwarz & Gusenleitner, 2017

Nomada pilosa Schwarz & Gusenleitner, 2017: 980. Holotype ♀; Turkey: Konya, 5.vi.1967, leg. J. Gusenleitner (MSPC)

Distribution. Ukraine, Crimea and southern Russia. Outside Europe known from Turkey and Iran (Smit 2018).

Nomada pyrgosica Schwarz & Smit, 2018

Nomada pyrgosica Schwarz & Smit in Smit, 2018: 214. Holotype ♀; Cyprus: Gazimagusa Dipkarpaz, Kiles point, 29.iii.2012,

leg. Schwenninger (MSPC).

Distribution. Cyprus. Outside Europe known from Turkey (Smit 2018).

Nomada simulatrix Schwarz & Smit, 2018

Nomada simulatrix Schwarz & Smit in Smit, 2018: 227. Holotype ♀; Greece: Phocis, Delphi, 8.iv.1963, leg, W. Grünwaldt

(MSPC).

Distribution. Greece.

Nomada smiti Schwarz, 2018

Nomada smiti Schwarz in Smit, 2018: 229. Holotype ♀; Spain: Madrid, Casa de Campo, 4.v.1965, leg. V. Llorente (MSPC).

Distribution. Spain.

Nomada tarsalis Schwarz & Smit, 2018

Nomada tarsalis Schwarz & Smit in Smit, 2018: 235. Holotype ♀; Israel: Elom, 27.iii.1946, leg. Bytinski-Salz (MSPC).

Distribution. Bulgaria, East Aegean Islands. Outside Europe known from Turkey, Israel and Iran (Smit 2018).

Nomada teunisseni Schwarz & Smit, 2018

Nomada teunisseni Schwarz & Smit in Smit, 2018: 237. Holotype ♂; Cyprus: Akanthou, 12.iii.1981, leg. H. Teunissen

(MSPC).

Distribution. Cyprus.

Nomada tuberculifera Schwarz & Smit, 2018

Nomada tuberculifera Schwarz & Smit in Smit, 2018: 241. Holotype ♀; Greece: Crete, Sitia, 18.v.1963, leg. M. Schwarz

(MSPC).

Distribution. Greece (Peloponnese, Crete). Outside Europe known from Turkey.

GHISBAIN, ROSA ET AL.

Nomada unica Schwarz & Smit, 2018

Nomada unica Schwarz & Smit in Smit, 2018: 243. Holotype ♀; Turkey: Konya, 5.viii.1987, leg. J. Gusenleitner (MSPC).

Distribution. Greece. Outside Europe known from Turkey and Israel (Smit 2018).

Nomada warnckei Schwarz & Smit, 2018

Nomada warnckei Schwarz & Smit in Smit, 2018: 247. Holotype ♀; Turkey: Hakkari, Suvari–Halil–Pass, 2500 m, 2.vi.1980,

leg. M. Schwarz (MSPC).

Distribution. Greece (Lesvos). Outside Europe known from Turkey (Smit 2018).

Nomada yermasoyiae Schwarz, Smit & Gusenleitner 2018

Nomada yermasoyiae Schwarz, Smit & Gusenleitner, 2018: 1422. Holotype ♀; Cyprus: Cherkes, 23.iii.1950, leg. Mavromoustakis

(MSPC).

Distribution. Cyprus. Outside Europe known from Israel (Smit 2018).

Published synonymies

Nomada ferghanica Morawitz, 1875

Synonymised with Nomada numida Lepeletier, 1841, which is the senior synonym according to Smit (2018).

Nomada lagrecai Nobile, 1990

Synonymised with Nomada hungarica Dalla Torre & Friese, 1894, which is the senior synonym according to Smit

(2018).

Nomada longipalpis Schwarz & Smit, 2020

Described as a new species in Schwarz & Smit. (2020), then synonymised with Nomada kriesteni Schwarz &

Gusenleitner, 2013, which is the senior synonym according to Schwarz & Smit (2021).

Nomada obscuriceps Schwarz & Levchenko, 2017

Synonymised with Nomada mitaii Proshchalykin, 2010, which is the senior synonym according to Proshchalykin

et al. (2019).

Nomada transitoria Schmiedeknecht, 1882

Synonymised with Nomada corcyraea Schmiedeknecht, 1882, which is the senior synonym according to Schwarz

& Gusenleitner (2015).

Taxonomic acts and clarifications

Nomada cypriaca (in Schwarz, 1999)

Incorrect subsequent spelling of Nomada cypria Mavromoustakis, 1952 and N. cypricola Mavromoustakis, 1957

(Schwarz 1999).

Nomada glabella Thomson, 1870

More cryptic species are being found in the Nomada panzeri group, and future changes are expected based on

ongoing studies. The name Nomada glabella sensu Stöckert (1954) does not match the lectotype (Falk et al. 2022)

and additional work is needed on this species complex. For now Nomada glabella is considered as a species

inquirenda and therefore not included in the European checklist.

Nomada jaramense Dusmet y Alonso, 1913

Incorrect subsequent spelling of Nomada jaramensis Dusmet y Alonso, 1913.

Nomada minuscula Noskiewicz, 1930

For some authors and in Nieto et al. (2014), Nomada minuscula is a subspecies of Nomada sheppardana (Kirby,

1802) (Smit 2018). Here we follow the opinion Scheuchl (2000) and Scheuchl & Willner (2016) and consider both

taxa as heterospecific. Both are therefore listed in the present checklist.

Distribution. Portugal, Spain, France, Germany, Switzerland (extinct), Austria, Italy (mainland, Sicily),

Slovenia, Czech Republic, Greece, Serbia, Hungary, Poland, Ukraine and European part of Russia (Smit 2018;

Baldock et al. 2018). Outside Europe known from Morocco, Algeria, Tunisia.

Nomada rufa Rossi, 1790

Species originally described from Italy by Rossi (1790). It was considered as a valid species by Balzan et al. (2016).

We here consider the name Nomada rufa a nomen dubium; while it likely has priority on other names currently in

use, it cannot be correctly attributed to any taxon as the type series is lost and the original description is ambiguous.

Material under this name should be revised, and the taxon is for now excluded from the European checklist.

Nomada siciliensis Dalla Torre & Friese, 1894

The status of this taxon is uncertain. The description of this species was based on a single male collected in Sicily

(Santa Ninfa, Trapani province) (Dalla Torre & Friese 1894). The holotype was destroyed, and no other specimen

of this taxon is known (Smit 2018; M. Schwarz, pers. comm.). The name was considered a nomen dubium by Smit

(2018) and is therefore not considered in the present checklist.

Nomada subcornuta (Kirby, 1802)

A recent genetic study suggests that it is a distinct species from Nomada fulvicornis Fabricius, 1793 (Falk et al.

2017). Both species are therefore considered as valid and are included in the present checklist.

Distribution. United Kingdom, Belgium, Netherlands, Germany, Czech Republic, Hungary, Estonia, Finland,

Sweden, Norway, Russia.

GHISBAIN, ROSA ET AL.

Nomada tormentillae Alfken, 1901

Nomada tormentillae Alfken, 1901 is treated as a valid species by some authors (Scheuchl & Willner 2016) and as

a subspecies or form of N. roberjeotiana Panzer, 1799 by others (Amiet et al. 2007; Smit 2018; M. Schwarz, pers.