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Biodiversity Data Journal 12: e117172 doi: 10.3897/BDJ.12.e117172 Data Paper The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees Thomas James Wood , Hugo Gaspar , Romain Le Divelec , Andreia Penado , Teresa Luísa Silva , Vanessa A. Mata , Joana Veríssimo , Denis Michez , Sílvia Castro , João Loureiro , Pedro Beja , Sónia Ferreira ‡ Naturalis Biodiversity Center, Leiden, Netherlands § Centre for Functional Ecology, Associate Laboratory TERRA, Department of Life Sciences, University of Coimbra, Calçada Martim de Freitas, 3000-456, Coimbra, Portugal | University of Mons, Research Institute for Biosciences, Laboratory of Zoology, Place du parc 20, 7000, Mons, Belgium ¶ Ciência Viva - Agência Nacional para a Cultura Científica e Tecnológica, Largo José Mariano Gago n.º1, Parque das Nações, 1990-073, Lisboa, Portugal # BIOPOLIS Program in Genomics, Biodiversity and Land Planning, CIBIO, Campus de Vairão, 4485–661 Vairão, Vila do Conde, Portugal ¤ CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, InBIO Laboratório Associado, Campus de Vairão, Universidade do Porto, 4485-661 Vairão, Vila do Conde, Portugal « University of Mons, Bruxelles, Belgium » EBM, Estação Biológica de Mertola, Praça Luis de Camoes, Mertola, Mertola, Portugal Corresponding author: Sónia Ferreira ([email protected]) Academic editor: Dominique Zimmermann Received: 11 Dec 2023 | Accepted: 14 Feb 2024 | Published: 05 Mar 2024 Citation: Wood T, Gaspar H, Le Divelec R, Penado A, Silva TL, Mata VA, Veríssimo J, Michez D, Castro S, Loureiro J, Beja P, Ferreira S (2024) The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees. Biodiversity Data Journal 12: e117172. https://doi.org/10.3897/BDJ.12.e117172 Abstract Background Bees are important actors in terrestrial ecosystems and are recognised for their prominent role as pollinators. In the Iberian Peninsula, approximately 1,100 bee species are known, with nearly 100 of these species being endemic to the Peninsula. A reference collection of DNA barcodes, based on morphologically identified bee specimens, representing 514 Iberian species, was constructed. The "InBIO Barcoding Initiative Database: DNA Barcodes of Iberian bees" dataset contains records of 1,059 sequenced specimens. The species of this dataset correspond to about 47% of Iberian bee species diversity and 21% ‡ § | ¶ #,¤ #,¤ #,¤ « § § #,¤ #,¤,» © Wood T et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
of endemic species diversity. For peninsular Portugal only, the corresponding coverage is 71% and 50%. Specimens were collected between 2014 and 2022 and are deposited in the research collection of Thomas Wood (Naturalis Biodiversity Center, The Netherlands), in the FLOWer Lab collection at the University of Coimbra (Portugal), in the Andreia Penado collection at the Natural History and Science Museum of the University of Porto (MHNC-UP) (Portugal) and in the InBIO Barcoding Initiative (IBI) reference collection (Vairão, Portugal). New information Of the 514 species sequenced, 75 species from five different families are new additions to the Barcode of Life Data System (BOLD) and 112 new BINs were added. Whilst the majority of species were assigned to a single BIN (94.9%), 27 nominal species were assigned to multiple BINs. Although the placement into multiple BINs may simply reflect genetic diversity and variation, it likely also represents currently unrecognised species-level diversity across diverse taxa, such as Amegilla albigena Lepeletier, 1841, Andrena russula Lepeletier, 1841, Lasioglossum leucozonium (Schrank, 1781), Nomada femoralis Morawitz, 1869 and Sphecodes alternatus Smith, 1853. Further species pairs of Colletes, Hylaeus and Nomada were placed into the same BINs, emphasising the need for integrative taxonomy within Iberia and across the Mediterranean Basin more broadly. These data substantially contribute to our understanding of bee genetic diversity and DNA barcodes in Iberia and provide an important baseline for ongoing taxonomic revisions in the West Palaearctic biogeographical region. Keywords Hymenoptera, occurrence records, species distributions, peninsular Portugal, peninsular Spain, DNA barcode, cytochrome c oxidase subunit I (COI), pollinator Introduction Bees are a diversified lineage of Hymenopteran insects, with nearly 21,000 species recognised globally (Ascher and Pickering 2023). The greatest bee diversity is found in parts of the world with a Mediterranean to xeric climate (Orr et al. 2021), most clearly south-western Australia, south-western North America, south-western South America, southern Africa and the Old World Mediterranean Basin. Within the Mediterranean Basin, the Iberian Peninsula dominates its western part and is made up of the two countries Portugal (mainland area 89,000 km ) and Spain (mainland area 492,000 km ). Peninsular Portugal hosts 725 species (Baldock et al. 2018, Wood et al. 2020, Gaspar et al. 2023, Reverté et al. 2023), of which only two bee species are currently recognised as endemic and peninsular Spain hosts approximately 1,100 species (Lhomme et al. 2020, OrtizSánchez 2020). 2 2 2Wood T et al
The total Iberian fauna is represented by approximately 1,100 species belonging to the six major bee families (Apidae, Andrenidae, Colletidae, Halictidae, Megachilidae and Melittidae) with approximately 96 species endemic to the Peninsula (8.7%); these values continue to vary due to ongoing taxonomic revisions which both describe and synonymise taxa, as well as adding new records and exclusions (e.g. Ghisbain et al. (2021b), Cross (2023), Dorchin (2023), Wood (2023), Ghisbain et al. (2023a)). Iberia, therefore, hosts slightly over half of the 2,138 bee species from Europe (Ghisbain et al. 2023b) and around one third of the approximately 3,500 bee species found in the entire West Palaearctic biogeographical region (Rasmont et al. 2017 and subsequent taxonomic works). Given the large size of the Iberian bee fauna, against the background of widely-reported bee declines (LeBuhn and Vargas Luna 2021, Zattara and Aizen 2021) and the global and regional importance of Iberian bees as pollinators of wild (e.g. Castro et al. (2013), Mendes et al. (2021)) and cultivated plants (e.g. Stefanescu et al. (2018), Gaspar et al. (2022), Mota et al. (2022)), a good understanding of this fauna is highly relevant to ensure appropriate conservation actions (LeBuhn and Vargas Luna 2021, Zattara and Aizen 2021). However, whilst the national totals for Portugal and Spain are now more or less complete, this somewhat masks how patchy the study of Iberian bees has been. Before 2018, knowledge of the Portuguese fauna was seriously incomplete (Baldock et al. 2018) and many taxonomic issues have plagued the understanding of the wider Iberian fauna. Some of the taxonomic issues remained unsolved or simply unresolved for hundreds of years (e.g. Pauly et al. (2019), Litman et al. (2021), Wood and Le Divelec (2022), Dorchin (2023), Praz and Bénon (2023), Wood (2023)) and some were only discovered and resolved much more recently (e.g. Wood et al. (2021), Wood (2022)). Many parts of Iberia remain poorly studied and much work is still required to fully understand the distribution of bees within Iberia (e.g. Álvarez Fidalgo et al. (2022), Gaspar et al. (2023)), certainly when compared to countries in Northern and Central Europe that have a long history of study and a large quantity of validated occurrence data (e.g. Peeters et al. (2012), Else and Edwards (2018), Drossart et al. (2019), Praz et al. (2023)). In addition to the incomplete study, the Iberian bee fauna has also been seldom studied using genetic techniques as a way to taxonomically delineate its fauna (as opposed to focused population genetic studies or taxonomic studies on specific species, for example, Chávez‐Galarza et al. (2015), Cejas et al. (2019), de Sousa et al. (2023)) until very recently (e.g. Ghisbain et al. (2021a), Ghisbain et al. (2021b), Wood et al. (2021), Kasparek et al. (2023), Wood (2023)). These studies have predominantly employed DNA barcodes targeting the cytochrome oxidase I subunit (COI) of the mitochondrial genome, a technique which is widely used in bee taxonomy to delineate species (e.g.Packer and Ruz 2017, Williams et al. 2020, Brasero et al. 2021, McLaughlin et al. 2022, Praz et al. 2022, Wood 2022, Gardner and Gibbs 2023, Praz and Bénon 2023, Wood 2023). However, it is important to note that this approach can produce ambiguous results that do not consistently separate certain taxa or can lead to conflict with morphological characters (e.g. Gibbs (2018)), which can require more powerful analyses to resolve (e.g. Gueuning et al. (2020)). The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 3
More broadly, however, the DNA barcoding approach is largely supportive for the majority of existing taxonomic concepts in bees, at least in Europe (Schmidt et al. 2015). Indeed, the work of Schmidt et al. (2015) is one of the few studies to attempt to systematically barcode the entire fauna of an European country, in this case Germany (though see also Magnacca and Brown (2012), Villalta et al. (2021)). In order to meaningfully increase our knowledge of the DNA barcodes of Iberian bees and, hence, also their taxonomic status and presence in Iberia, we attempt to systematically barcode this fauna, with a focus on the species present in peninsular Portugal. We generated barcodes for species representing 46.5% (n = 512) of the bees known from the Peninsula and 20.8% (n = 20) of known endemic Iberian species. This work was conducted within the framework of the InBIO Barcoding Initiative. General description Purpose: This dataset aims to provide a contribution to an authoritative DNA barcode sequences library for Iberian bees, documenting biodiversity of this functional group, harbouring relevant pollinators in a Mediterranean hotspot. Such a library aims to enable DNA-based identification of species for both traditional molecular studies and DNAmetabarcoding studies. Furthermore, it constitutes a relevant resource for taxonomic research on Iberian bees and their distributions within and outside the Iberian Peninsula. Additional information: A total of 1,059 bee specimens were sequenced (Suppl. materials 1, 2, 3, 4). Fig. 2 illustrates examples of the diversity of species that are part of the dataset of distribution data and DNA barcodes of Iberian Bees. A full-length barcode of 658 bp was obtained for 1,007 specimens (95%) (Table 1, Suppl. material 2). These specimens represent 514 (46.5%) of the approximately 1,100 described bee species known to occur in the Iberian Peninsula, one species of Melecta being identified only to the level of genus. Furthermore, 20 taxa are Iberian endemics, representing 20.8% of the total species endemic to the Iberian Penisnsula (n = 96, subject to continuous variation due to ongoing taxonomic revisions and, hence, changing species concepts and distributions, for example, Wood (2023)). For peninsular Portugal, the corresponding values represent 70.8% of the fauna (ca. 725 species) and 50% of the endemic species (one of the only two species endemic to mainland Portugal). The dataset includes specimens from all six bee families known to occur in the Iberian Peninsula (Apidae, Andrenidae, Colletidae, Halictidae, Megachilidae, Melittidae) and 51 of the 56 (91.1%) genera present in the fauna (treating Eucera and Tetralonia as distinct genera and retaining broad Halictus genus including Seladonia and Vestitohalictus), lacking only Simpanurgus (ne species in Iberia), Habropoda (one species), Camptopoeum (one species), Biastes (three species) and Rophites (two species). These data substantially contribute to our understanding of bee barcodes with 75 species represented by barcodes for the first time in the BOLD database. Currently, 91% of the Portuguese bee species have public DNA barcodes; nevertheless, 64 species remain unrepresented in the DNA barcode reference collections (Suppl. material 5). 4Wood T et al
Family Species IBI code BOLD code BOLD BIN GenBank Andrenidae Andrena aerinifrons Dours, 1873 INV12500 IBIHM471-21 BOLD:ADJ3594 OR795910 Andrenidae Andrena aerinifrons Dours, 1873 INV12633 IBIHM1034-22 BOLD:AET5046'OR796236 Andrenidae Andrena aerinifrons Dours, 1873 INV12827 IBIHM1228-22 BOLD:AET5046'OR795930 Andrenidae Andrena afzeliella (Kirby, 1802) INV12647 IBIHM1048-22 BOLD:AAP2754 OR795919 Andrenidae Andrena agilissima (Scopoli, 1770) INV12501 IBIHM472-21 BOLD:AEO1903 OR795937 Andrenidae Andrena agilissima (Scopoli, 1770) INV12502 IBIHM473-21 BOLD:AEO1903 OR796727 Andrenidae Andrena agilissima (Scopoli, 1770) INV13382 IBIHM952-21 BOLD:AEO1903 OR796085 Andrenidae Andrena alfkenella Perkins, 1914 INV12503 IBIHM474-21 BOLD:AAK0552 OR796780 Andrenidae Andrena alfkenella Perkins, 1914 INV13052 IBIHM623-21 BOLD:AAK0552 OR796891 Andrenidae Andrena alfkenella Perkins, 1914 INV13198 IBIHM769-21 BOLD:AAK0552 OR796142 Andrenidae Andrena alfkenella Perkins, 1914 INV13199 IBIHM770-21 BOLD:AAK0552 OR796444 Andrenidae Andrena alluaudi Benoist, 1961# INV12504 IBIHM475-21 BOLD:AEN9732'OR796776 Andrenidae Andrena alluaudi Benoist, 1961# INV12505 IBIHM476-21 BOLD:AEN9732'OR796724 Andrenidae Andrena alma Warncke, 1975 INV12828 IBIHM1229-22 BOLD:AEN9431'OR796777 Andrenidae Andrena alma Warncke, 1975 INV12829 IBIHM1230-22 BOLD:AEN9431'OR796133 Andrenidae Andrena alma Warncke, 1975 INV13377 IBIHM947-21 BOLD:AEN9431'OR796746 Andrenidae Andrena ampla Warncke, 1967 INV13168 IBIHM739-21 BOLD:ABA2611 OR796274 Andrenidae Andrena ampla Warncke, 1967 INV12635 IBIHM1036-22 BOLD:AES0278'OR796123 Andrenidae Andrena angustior (Kirby, 1802) INV12638 IBIHM1039-22 BOLD:AAE5176 OR796639 Andrenidae Andrena angustior (Kirby, 1802) INV13174 IBIHM745-21 BOLD:AAE5176 OR796475 Andrenidae Andrena angustior (Kirby, 1802) INV13175 IBIHM746-21 BOLD:AAE5176 OR795960 Andrenidae Andrena angustior (Kirby, 1802) INV13176 IBIHM747-21 BOLD:AAE5176 OR796180 Andrenidae Andrena angustior (Kirby, 1802) INV13180 IBIHM751-21 BOLD:AAE5176 OR796837 Andrenidae Andrena angustior (Kirby, 1802) INV13181 IBIHM752-21 BOLD:AAE5176 OR796285 Andrenidae Andrena angustior (Kirby, 1802) INV13182 IBIHM753-21 BOLD:AAE5176 OR796538 Andrenidae Andrena angustior (Kirby, 1802) INV13183 IBIHM754-21 BOLD:AAE5176 OR796374 Table 1. List of species that were collected and DNA barcoded for this project. "IBI code" is the Sample ID, while "BOLD code" is the Process ID attributed by BOLD. In the Species column, # indicates taxa for which no DNA barcode was available prior to this study; in the BOLD BIN column, ' indicates unique BINs. The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 5
Family Species IBI code BOLD code BOLD BIN GenBank Andrenidae Andrena angustior (Kirby, 1802) INV13184 IBIHM755-21 BOLD:AAE5176 OR796685 Andrenidae Andrena angustior (Kirby, 1802) INV13188 IBIHM759-21 BOLD:AAE5176 OR796929 Andrenidae Andrena angustior (Kirby, 1802) INV13192 IBIHM763-21 BOLD:AAE5176 OR796292 Andrenidae Andrena angustior (Kirby, 1802) INV13196 IBIHM767-21 BOLD:AAE5176 OR796082 Andrenidae Andrena angustior (Kirby, 1802) INV13226 IBIHM797-21 BOLD:AAE5176 OR796580 Andrenidae Andrena angustior (Kirby, 1802) INV13227 IBIHM798-21 BOLD:AAE5176 OR796304 Andrenidae Andrena antigana Pérez, 1895 INV12506 IBIHM477-21 BOLD:AEO4687 OR796250 Andrenidae Andrena antigana Pérez, 1895 INV12507 IBIHM478-21 BOLD:AEO4687 OR796852 Andrenidae Andrena assimilis Radoszkowski, 1876 INV12627 IBIHM1028-22 BOLD:AEU3679'OR796869 Andrenidae Andrena baetica Wood, 2020 INV12821 IBIHM1222-22 BOLD:AEN8421 OR796472 Andrenidae Andrena bellidis Pérez, 1895# INV12824 IBIHM1225-22 BOLD:AET0142'OR796068 Andrenidae Andrena bellidis Pérez, 1895# INV12825 IBIHM1226-22 BOLD:AET0142'OR796765 Andrenidae Andrena benoisti Wood, 2021 INV12637 IBIHM1038-22 BOLD:AEO0062 OR796387 Andrenidae Andrena bicolor Fabricius, 1775 INV13051 IBIHM622-21 BOLD:AAD0135 OR796905 Andrenidae Andrena bicolorata (Rossi, 1790) INV12628 IBIHM1029-22 BOLD:AEK9626 OR796512 Andrenidae Andrena bicolorata (Rossi, 1790) INV12826 IBIHM1227-22 BOLD:AEK9626 OR796382 Andrenidae Andrena bimaculata (Kirby, 1802) INV12508 IBIHM479-21 BOLD:AAY9952 OR796913 Andrenidae Andrena bimaculata (Kirby, 1802) INV13398 IBIHM968-21 BOLD:AAY9952 OR796445 Andrenidae Andrena cinerea Brullé, 1832 INV12509 IBIHM480-21 BOLD:AEM8298 OR796689 Andrenidae Andrena cinerea Brullé, 1832 INV12823 IBIHM1224-22 BOLD:AEM8298 OR796590 Andrenidae Andrena combinata (Christ, 1791) INV12634 IBIHM1035-22 BOLD:AAK0530 OR796412 Andrenidae Andrena combinata (Christ, 1791) INV12649 IBIHM1050-22 BOLD:AAK0530 OR796847 Andrenidae Andrena discors Erichson, 1841 INV12546 IBIHM517-21 BOLD:AEI2585 OR796311 Andrenidae Andrena djelfensis Pérez, 1895 INV12818 IBIHM1219-22 BOLD:AET0920 OR796145 Andrenidae Andrena doursana citreola Warncke, 1975 INV12822 IBIHM1223-22 BOLD:AET0918 OR796522 Andrenidae Andrena fabrella Pérez, 1903 INV12639 IBIHM1040-22 BOLD:AEO4550 OR796649 Andrenidae Andrena fabrella Pérez, 1903 INV12645 IBIHM1046-22 BOLD:AEO4550 OR796248 Andrenidae Andrena ferrugineicrus Dours, 1872 INV12512 IBIHM483-21 BOLD:AEN6074 OR795954 Andrenidae Andrena flavipes Panzer, 1799 INV12513 IBIHM484-21 BOLD:ACX2946 OR796075 Andrenidae Andrena flavipes Panzer, 1799 INV12817 IBIHM1218-22 BOLD:ACX2946 OR796258 6Wood T et al
Family Species IBI code BOLD code BOLD BIN GenBank Andrenidae Andrena flavipes Panzer, 1799 INV13161 IBIHM732-21 BOLD:ACX2946 OR796504 Andrenidae Andrena flavipes Panzer, 1799 INV13162 IBIHM733-21 BOLD:ACX2946 OR796762 Andrenidae Andrena flavipes Panzer, 1799 INV13163 IBIHM734-21 BOLD:ACX2946 OR796170 Andrenidae Andrena flavipes Panzer, 1799 INV13164 IBIHM735-21 BOLD:ACX2946 OR796714 Andrenidae Andrena flavipes Panzer, 1799 INV13165 IBIHM736-21 BOLD:ACX2946 OR796733 Andrenidae Andrena flavipes Panzer, 1799 INV13392 IBIHM962-21 BOLD:ACX2946 OR796291 Andrenidae Andrena florea Fabricius, 1793 INV12514 IBIHM485-21 BOLD:AAK0287 OR796059 Andrenidae Andrena florea Fabricius, 1793 INV13166 IBIHM737-21 BOLD:AAK0287 OR796159 Andrenidae Andrena florea Fabricius, 1793 INV13169 IBIHM740-21 BOLD:AAK0287 OR796279 Andrenidae Andrena florea Fabricius, 1793 INV13172 IBIHM743-21 BOLD:AAK0287 OR796755 Andrenidae Andrena florentina Magretti, 1883 INV12515 IBIHM486-21 BOLD:ADZ9682 OR795967 Andrenidae Andrena florentina Magretti, 1883 INV13193 IBIHM764-21 BOLD:ADZ9682 OR796604 Andrenidae Andrena florentina Magretti, 1883 INV13387 IBIHM957-21 BOLD:ADZ9682 OR796889 Andrenidae Andrena florentina Magretti, 1883 INV13388 IBIHM958-21 BOLD:ADZ9682 OR796719 Andrenidae Andrena florentina Magretti, 1883 INV13394 IBIHM964-21 BOLD:ADZ9682 OR796136 Andrenidae Andrena fulica Warncke, 1974 INV12629 IBIHM1030-22 BOLD:AEN6978 OR796196 Andrenidae Andrena fulica Warncke, 1974 INV13379 IBIHM949-21 BOLD:AEN6978 OR796041 Andrenidae Andrena fulvago (Christ, 1791) INV12516 IBIHM487-21 BOLD:AAK0296 OR795999 Andrenidae Andrena fuscipes (Kirby, 1802) INV15655 IBIHY003-22 BOLD:AAE1706 OR796101 Andrenidae Andrena fuscipes (Kirby, 1802) INV15677 IBIHY020-22 BOLD:AAE1706 OR796027 Andrenidae Andrena fuscosa Erichson, 1835 INV12517 IBIHM488-21 BOLD:AEO6270 OR796949 Andrenidae Andrena granulosa Pérez, 1902 INV12518 IBIHM489-21 BOLD:AAR3409 OR796195 Andrenidae Andrena haemorrhoa (Fabricius, 1781) INV12519 IBIHM490-21 BOLD:AAD8891 OR796468 Andrenidae Andrena hedikae Jaeger, 1934 INV12520 IBIHM491-21 BOLD:AEO3736 OR796737 Andrenidae Andrena hedikae Jaeger, 1934 INV12521 IBIHM492-21 BOLD:AEO3736 OR796601 Andrenidae Andrena hedikae Jaeger, 1934 INV13418 IBIHM988-21 BOLD:AEO3736 OR796558 Andrenidae Andrena hesperia Smith, 1853 INV12836 IBIHM1237-22 BOLD:AEN4997 OR796647 Andrenidae Andrena hesperia Smith, 1853 INV12841 IBIHM1242-22 BOLD:AEN4997 OR796360 Andrenidae Andrena hesperia Smith, 1853 INV12842 IBIHM1243-22 BOLD:AEO5653 OR796396 Andrenidae Andrena hesperia Smith, 1853 INV13376 IBIHM946-21 BOLD:AEO5653 OR796299 The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 7
Family Species IBI code BOLD code BOLD BIN GenBank Andrenidae Andrena humilis Imhoff, 1832 INV12524 IBIHM495-21 BOLD:AER4065 OR796904 Andrenidae Andrena hypopolia Schmiedeknecht, 1884 INV12630 IBIHM1031-22 BOLD:AAJ2215 OR796330 Andrenidae Andrena hypopolia Schmiedeknecht, 1884 INV12648 IBIHM1049-22 BOLD:AAJ2215 OR796871 Andrenidae Andrena hystrix Schmiedeknecht, 1883 INV12631 IBIHM1032-22 BOLD:AEO0162 OR796029 Andrenidae Andrena hystrix Schmiedeknecht, 1883 INV12632 IBIHM1033-22 BOLD:AEO0162 OR796383 Andrenidae Andrena impressa Warncke, 1967 INV12646 IBIHM1047-22 BOLD:AEO5002 OR796456 Andrenidae Andrena juliana Wood, 2021 INV12525 IBIHM496-21 BOLD:AEO3956'OR796882 Andrenidae Andrena labialis (Kirby, 1802) INV12526 IBIHM497-21 BOLD:AAK0232 OR796783 Andrenidae Andrena labialis (Kirby, 1802) INV13189 IBIHM760-21 BOLD:AAK0232 OR796394 Andrenidae Andrena labiata Fabricius, 1781 INV02352 IBIHM019-19 BOLD:ADS9137 OR796910 Andrenidae Andrena labiata Fabricius, 1781 INV02353 IBIHM020-19 BOLD:ADS9137 OR796320 Andrenidae Andrena labiata Fabricius, 1781 INV12527 IBIHM498-21 BOLD:ADS9137 OR796000 Andrenidae Andrena labiata Fabricius, 1781 INV13419 IBIHM989-21 BOLD:ADS9137 OR796244 Andrenidae Andrena lagopus Latreille, 1809 INV12528 IBIHM499-21 BOLD:AAK0222 OR796638 Andrenidae Andrena lagopus Latreille, 1809 INV13389 IBIHM959-21 BOLD:AAK0222 OR796802 Andrenidae Andrena lagopus Latreille, 1809 INV13396 IBIHM966-21 BOLD:AAK0222 OR795990 Andrenidae Andrena leptopyga Pérez, 1895 INV13371 IBIHM941-21 BOLD:AEO3338 OR796421 Andrenidae Andrena leucolippa Pérez, 1895 INV12529 IBIHM500-21 BOLD:AEO2473'OR796565 Andrenidae Andrena limata Smith, 1853 INV12530 IBIHM501-21 BOLD:AAE1815 OR796200 Andrenidae Andrena limata Smith, 1853 INV13386 IBIHM956-21 BOLD:AAE1815 OR796862 Andrenidae Andrena limata Smith, 1853 INV13384 IBIHM954-21 BOLD:AEX3903 OR796675 Andrenidae Andrena livens Pérez, 1895 INV12531 IBIHM502-21 BOLD:AEL4518 OR796809 Andrenidae Andrena livens Pérez, 1895 INV12532 IBIHM503-21 BOLD:AEL4518 OR796116 Andrenidae Andrena longibarbis Pérez, 1895 INV12533 IBIHM504-21 BOLD:AEO0110 OR795913 Andrenidae Andrena longibarbis Pérez, 1895 INV12534 IBIHM505-21 BOLD:AEO0110 OR796779 Andrenidae Andrena longibarbis Pérez, 1895 INV13422 IBIHM992-21 BOLD:AEO0110 OR796557 Andrenidae Andrena lusitania Wood & OrtizSánchez, 2022# INV13428 IBIHM998-21 BOLD:AET5292 OR796355 Andrenidae Andrena minutula (Kirby, 1802) INV12535 IBIHM506-21 BOLD:AAJ2143 OR796477 Andrenidae Andrena minutula (Kirby, 1802) INV12536 IBIHM507-21 BOLD:AAJ2143 OR796060 8Wood T et al
Family Species IBI code BOLD code BOLD BIN GenBank Andrenidae Andrena minutula (Kirby, 1802) INV12564 IBIHM535-21 BOLD:AAJ2143 OR796044 Andrenidae Andrena minutula (Kirby, 1802) INV13191 IBIHM762-21 BOLD:AAJ2143 OR796198 Andrenidae Andrena minutula (Kirby, 1802) INV13407 IBIHM977-21 BOLD:AAJ2143 OR796039 Andrenidae Andrena minutuloides Perkins, 1914 INV12537 IBIHM508-21 BOLD:AAE4947 OR796022 Andrenidae Andrena morio Brullé, 1832 INV12523 IBIHM494-21 BOLD:AAJ2141 OR795955 Andrenidae Andrena morio Brullé, 1832 INV13380 IBIHM950-21 BOLD:AAJ2141 OR796420 Andrenidae Andrena morio Brullé, 1832 INV13381 IBIHM951-21 BOLD:AER2061 OR796324 Andrenidae Andrena nana (Kirby, 1802) INV12538 IBIHM509-21 BOLD:AAR3413 OR796476 Andrenidae Andrena nana (Kirby, 1802) INV13404 IBIHM974-21 BOLD:AAR3413 OR796337 Andrenidae Andrena nana (Kirby, 1802) INV13406 IBIHM976-21 BOLD:AAR3413 OR795968 Andrenidae Andrena nigroaenea (Kirby, 1802) INV12539 IBIHM510-21 BOLD:AAC8068 OR796900 Andrenidae Andrena nigroaenea (Kirby, 1802) INV13173 IBIHM744-21 BOLD:AAC8068 OR796237 Andrenidae Andrena nigroaenea (Kirby, 1802) INV13186 IBIHM757-21 BOLD:AAC8068 OR796885 Andrenidae Andrena nigroolivacea Dours, 1873 INV12540 IBIHM511-21 BOLD:ADZ3313 OR796223 Andrenidae Andrena nigroolivacea Dours, 1873 INV13194 IBIHM765-21 BOLD:ADZ3313 OR796879 Andrenidae Andrena nigroolivacea Dours, 1873 INV13195 IBIHM766-21 BOLD:ADZ3313 OR795987 Andrenidae Andrena nitidula Pérez, 1903 INV12642 IBIHM1043-22 BOLD:AAJ2142 OR796203 Andrenidae Andrena nitidula Pérez, 1903 INV12814 IBIHM1215-22 BOLD:AAJ2142 OR795912 Andrenidae Andrena orana Warncke, 1975 INV12542 IBIHM513-21 BOLD:AEI8573 OR796921 Andrenidae Andrena orbitalis Morawitz, 1871# INV12543 IBIHM514-21 BOLD:AEO3428'OR796140 Andrenidae Andrena orbitalis Morawitz, 1871# INV13373 IBIHM943-21 BOLD:AEO3428'OR796432 Andrenidae Andrena orbitalis Morawitz, 1871# INV13374 IBIHM944-21 BOLD:AEO3428'OR796307 Andrenidae Andrena orbitalis Morawitz, 1871# INV13425 IBIHM995-21 BOLD:AEO3428'OR796096 Andrenidae Andrena ovatula (Kirby, 1802) INV12644 IBIHM1045-22 BOLD:AAK0399 OR796018 Andrenidae Andrena ovatula (Kirby, 1802) INV13414 IBIHM984-21 BOLD:AAK0399 OR796216 Andrenidae Andrena ovatula (Kirby, 1802) INV13426 IBIHM996-21 BOLD:AAK0399 OR796108 Andrenidae Andrena ovatula (Kirby, 1802) INV13429 IBIHM999-21 BOLD:AAK0399 OR796389 Andrenidae Andrena oviventris Pérez, 1895 INV12544 IBIHM515-21 BOLD:AEO5504 OR796772 Andrenidae Andrena oviventris Pérez, 1895 INV13378 IBIHM948-21 BOLD:AEO5504 OR796443 Andrenidae Andrena pandellei Pérez, 1895 INV12545 IBIHM516-21 BOLD:AEO5453 OR796629 The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 9
Family Species IBI code BOLD code BOLD BIN GenBank Apidae Ceratina chalybea Chevrier, 1872 INV12164 IBIHM235-21 BOLD:AAW6323 OR795927 Apidae Ceratina cucurbitina (Rossi, 1792) INV12163 IBIHM234-21 BOLD:AAO0285 OR796251 Apidae Ceratina cucurbitina (Rossi, 1792) INV13037 IBIHM608-21 BOLD:AAO0285 OR796824 Apidae Ceratina cucurbitina (Rossi, 1792) INV13214 IBIHM785-21 BOLD:AAO0285 OR796954 Apidae Ceratina cucurbitina (Rossi, 1792) INV15669 IBIHY012-22 BOLD:AAO0285 OR796797 Apidae Ceratina cyanea (Kirby, 1802) INV02339 IBIHM046-19 BOLD:AAE9812 OR796425 Apidae Ceratina cyanea (Kirby, 1802) INV12170 IBIHM241-21 BOLD:AAE9812 OR796723 Apidae Ceratina cyanea (Kirby, 1802) INV13034 IBIHM605-21 BOLD:AAE9812 OR796828 Apidae Ceratina cyanea (Kirby, 1802) INV13215 IBIHM786-21 BOLD:AAE9812 OR796552 Apidae Ceratina cyanea (Kirby, 1802) INV13216 IBIHM787-21 BOLD:AAE9812 OR796259 Apidae Ceratina cyanea (Kirby, 1802) INV13252 IBIHM822-21 BOLD:AAE9812 OR796823 Apidae Ceratina cyanea (Kirby, 1802) INV13254 IBIHM824-21 BOLD:AAE9812 OR795924 Apidae Ceratina dallatorreana Friese, 1896 INV12169 IBIHM240-21 BOLD:ADF8180 OR796186 Apidae Ceratina dallatorreana Friese, 1896 INV13035 IBIHM606-21 BOLD:ADF8180 OR796583 Apidae Ceratina dallatorreana Friese, 1896 INV13255 IBIHM825-21 BOLD:ADF8180 OR796684 Apidae Ceratina gravidula Gerstäcker, 1869 INV12161 IBIHM232-21 BOLD:AEA2549 OR796616 Apidae Ceratina mocsaryi Friese, 1896 INV12167 IBIHM238-21 BOLD:AEO3844 OR796766 Apidae Ceratina mocsaryi Friese, 1896 INV13253 IBIHM823-21 BOLD:AEO3844 OR796608 Apidae Ceratina mocsaryi Friese, 1896 INV13257 IBIHM827-21 BOLD:AEO3844 OR796376 Apidae Ceratina nigrolabiata Friese, 1896 INV12474 IBIHM1327-22 BOLD:ACG0777 OR796448 Apidae Ceratina nigrolabiata Friese, 1896 INV12475 IBIHM1328-22 BOLD:ACG0777 OR796678 Apidae Ceratina nigrolabiata Friese, 1896 INV13256 IBIHM826-21 BOLD:ACG0777 OR796868 Apidae Ceratina parvula Smith, 1854 INV12166 IBIHM237-21 BOLD:AEO2595'OR796596 Apidae Ceratina saundersi Daly, 1983# INV12176 IBIHM247-21 BOLD:AEO3675'OR796877 Apidae Epeolus cruciger marginatus Bischoff, 1930 INV12012 IBIHM083-21 BOLD:AAN3663 OR796946 Apidae Epeolus cruciger (Panzer, 1799) INV12462 IBIHM1315-22 BOLD:AAN3663 OR796164 Apidae Epeolus intermedius Pérez, 1884 INV12464 IBIHM1317-22 BOLD:AAJ0611 OR796091 Apidae Epeolus intermedius Pérez, 1884 INV12467 IBIHM1320-22 BOLD:AAJ0611 OR795988 Apidae Epeolus julliani Pérez, 1884 INV12463 IBIHM1316-22 BOLD:ACD1277 OR796659 Apidae Epeolus variegatus (Linnaeus, 1758) INV12466 IBIHM1319-22 BOLD:AAJ0611 OR796525 16 Wood T et al
Family Species IBI code BOLD code BOLD BIN GenBank Apidae Eucera barbiventris Pérez, 1902# INV12100 IBIHM171-21 BOLD:AEN2742 OR796562 Apidae Eucera barbiventris Pérez, 1902# INV13239 IBIHM809-21 BOLD:AEN2742 OR796912 Apidae Eucera chrysopyga Pérez, 1879 INV12114 IBIHM185-21 BOLD:ABV4745 OR796487 Apidae Eucera chrysopyga Pérez, 1879 INV13017 IBIHM588-21 BOLD:ABV4745 OR796235 Apidae Eucera chrysopyga Pérez, 1879 INV13021 IBIHM592-21 BOLD:ABV4745 OR796079 Apidae Eucera collaris Dours, 1873 INV12104 IBIHM175-21 BOLD:AEO1485'OR796413 Apidae Eucera collaris Dours, 1873 INV13237 IBIHM807-21 BOLD:AEO1485'OR796855 Apidae Eucera dalmatica Lepeletier, 1841 INV13235 IBIHM805-21 BOLD:ADE3334 OR795980 Apidae Eucera dalmatica Lepeletier, 1841 INV13236 IBIHM806-21 BOLD:ADE3334 OR796856 Apidae Eucera elongatula Vachal, 1907 INV12108 IBIHM179-21 BOLD:ABU9563 OR796076 Apidae Eucera elongatula Vachal, 1907 INV12109 IBIHM180-21 BOLD:ABU9563 OR796825 Apidae Eucera elongatula Vachal, 1907 INV13241 IBIHM811-21 BOLD:ABU9563 OR796516 Apidae Eucera elongatula Vachal, 1907 INV13242 IBIHM812-21 BOLD:ABU9563 OR796786 Apidae Eucera eucnemidea Dours, 1873 INV12110 IBIHM181-21 BOLD:AEO0071'OR796893 Apidae Eucera eucnemidea Dours, 1873 INV12111 IBIHM182-21 BOLD:AEO0071'OR796875 Apidae Eucera hispana Lepeletier, 1841# INV12116 IBIHM187-21 BOLD:AEO1591'OR796391 Apidae Eucera hungarica Friese, 1895 INV12099 IBIHM170-21 BOLD:AAL4610 OR796898 Apidae Eucera iberica (Dusmet y Alonso 1926) INV13251 IBIHM821-21 BOLD:AER1385'OR795982 Apidae Eucera interrupta Bär, 1850 INV12118 IBIHM189-21 BOLD:ABZ4791 OR796865 Apidae Eucera longicornis (Linnaeus, 1758) INV12115 IBIHM186-21 BOLD:ABZ4790 OR796805 Apidae Eucera longicornis (Linnaeus, 1758) INV12482 IBIHM1335-22 BOLD:ABZ4790 OR796183 Apidae Eucera longicornis (Linnaeus, 1758) INV12483 IBIHM1336-22 BOLD:ABZ4790 OR796957 Apidae Eucera longicornis (Linnaeus, 1758) INV13238 IBIHM808-21 BOLD:ABZ4790 OR796431 Apidae Eucera nigrescens Pérez, 1879 INV12105 IBIHM176-21 BOLD:ABZ2397 OR796395 Apidae Eucera nigrescens Pérez, 1879 INV13240 IBIHM810-21 BOLD:ABZ2397 OR796255 Apidae Eucera nigrescens Pérez, 1879 INV13246 IBIHM816-21 BOLD:ABZ2397 OR795989 Apidae Eucera nigrescens Pérez, 1879 INV13248 IBIHM818-21 BOLD:ABZ2397 OR796872 Apidae Eucera nigrescens Pérez, 1879 INV13250 IBIHM820-21 BOLD:ABZ2397 OR796333 Apidae Eucera nigrifacies Lepeletier, 1841 INV12120 IBIHM191-21 BOLD:AER1612'OR795899 Apidae Eucera nigrilabris Lepeletier, 1841 INV12121 IBIHM192-21 BOLD:AEM9739 OR796302 The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 17
Family Species IBI code BOLD code BOLD BIN GenBank Apidae Eucera nigrilabris Lepeletier, 1841 INV13019 IBIHM590-21 BOLD:AEM9739 OR796072 Apidae Eucera nigrilabris Lepeletier, 1841 INV13020 IBIHM591-21 BOLD:AEM9739 OR795907 Apidae Eucera notata Lepeletier, 1841# INV12122 IBIHM193-21 BOLD:AEN4764 OR796709 Apidae Eucera notata Lepeletier, 1841# INV13244 IBIHM814-21 BOLD:AEN4764 OR796384 Apidae Eucera notata Lepeletier, 1841# INV13245 IBIHM815-21 BOLD:AEN4764 OR796357 Apidae Eucera numida Lepeletier, 1841 INV12123 IBIHM194-21 BOLD:AEA1287 OR796729 Apidae Eucera proxima Morawitz, 1875 INV12832 IBIHM1233-22 BOLD:AAI4916 OR796770 Apidae Eucera proxima Morawitz, 1875 INV12833 IBIHM1234-22 BOLD:AAI4916 OR796908 Apidae Eucera punctatissima Pérez, 1895# INV12834 IBIHM1235-22 BOLD:AET2033'OR796940 Apidae Eucera punctatissima Pérez, 1895# INV12835 IBIHM1236-22 BOLD:AET2033'OR795939 Apidae Eucera rufa (Lepeletier, 1841) INV12124 IBIHM195-21 BOLD:ADZ9496 OR796720 Apidae Eucera ruficollis (Brullé, 1832) INV12098 IBIHM169-21 BOLD:AEM7547 OR796148 Apidae Eucera taurica Morawitz, 1870 INV12126 IBIHM197-21 BOLD:AEK9612 OR796182 Apidae Eucera tricincta Erichson, 1835# INV12127 IBIHM198-21 BOLD:AEO1994'OR796080 Apidae Melecta albifrons (Forster, 1771) INV12002 IBIHM073-21 BOLD:ACR2895 OR796340 Apidae Melecta albifrons (Forster, 1771) INV12003 IBIHM074-21 BOLD:ACR2895 OR796141 Apidae Melecta festiva Lieftinck, 1980# INV13336 IBIHM906-21 OR796751 Apidae Melecta sp. INV12001 IBIHM072-21 BOLD:AEO2855 OR796791 Apidae Nomada agrestis Fabricius, 1787 INV12035 IBIHM106-21 BOLD:AEN9589 OR796773 Apidae Nomada agrestis Fabricius, 1787 INV13265 IBIHM835-21 BOLD:AEN9589 OR796836 Apidae Nomada basalis Herrich-Schäffer, 1839 INV13270 IBIHM840-21 BOLD:AEK6178 OR796576 Apidae Nomada basalis Herrich-Schäffer, 1839 INV12036 IBIHM107-21 BOLD:AEN3462 OR796691 Apidae Nomada basalis Herrich-Schäffer, 1839 INV12037 IBIHM108-21 BOLD:AEO4155'OR796398 Apidae Nomada basalis Herrich-Schäffer, 1839 INV12038 IBIHM109-21 BOLD:AEO4155'OR796767 Apidae Nomada basalis Herrich-Schäffer, 1839 INV12039 IBIHM110-21 BOLD:AEO4155'OR795917 Apidae Nomada beaumonti Schwarz, 1967 INV12041 IBIHM112-21 BOLD:AEO4341 OR796899 Apidae Nomada bifasciata Olivier, 1811 INV12040 IBIHM111-21 BOLD:AAE5755 OR795993 Apidae Nomada bifasciata Olivier, 1811 INV12054 IBIHM125-21 BOLD:AAE5755 OR796048 Apidae Nomada bifasciata Olivier, 1811 INV12055 IBIHM126-21 BOLD:AAE5755 OR796033 Apidae Nomada bifasciata Olivier, 1811 INV13025 IBIHM596-21 BOLD:AAE5755 OR796641 18 Wood T et al
Family Species IBI code BOLD code BOLD BIN GenBank Apidae Nomada bluethgeni Stoeckhert, 1943 INV12608 IBIHM1009-22 BOLD:AES2302 OR796419 Apidae Nomada bluethgeni Stoeckhert, 1943 INV12609 IBIHM1010-22 BOLD:AES2302 OR796527 Apidae Nomada carnifex Mocsáry, 1883 INV12042 IBIHM113-21 BOLD:AEO0078 OR796582 Apidae Nomada carnifex Mocsáry, 1883 INV12043 IBIHM114-21 BOLD:AEO0078 OR796575 Apidae Nomada conjungens Herrich-Schäffer, 1839 INV12606 IBIHM1007-22 BOLD:AER6266'OR796630 Apidae Nomada conjungens Herrich-Schäffer, 1839 INV13024 IBIHM595-21 BOLD:AER6266'OR796943 Apidae Nomada corcyraea Schmiedeknecht, 1882# INV12845 IBIHM1246-22 BOLD:AEO6156 OR796137 Apidae Nomada cristata Pérez, 1896# INV12044 IBIHM115-21 BOLD:AEO1919'OR795922 Apidae Nomada dira Schmiedeknecht, 1882# INV12045 IBIHM116-21 BOLD:AEN3179 OR795964 Apidae Nomada discrepans Schmiedeknecht, 1882 INV12048 IBIHM119-21 BOLD:AEN8935'OR796695 Apidae Nomada discrepans Schmiedeknecht, 1882 INV13259 IBIHM829-21 BOLD:AEN8935'OR796009 Apidae Nomada discrepans Schmiedeknecht, 1882 INV13260 IBIHM830-21 BOLD:AEN8935'OR796323 Apidae Nomada distinguenda Morawitz, 1874 INV13023 IBIHM594-21 BOLD:ACY0250 OR796567 Apidae Nomada distinguenda Morawitz, 1874 INV13027 IBIHM598-21 BOLD:ACY0250 OR796090 Apidae Nomada distinguenda Morawitz, 1874 INV12602 IBIHM1003-22 BOLD:AET5764'OR796083 Apidae Nomada duplex Smith, 1854# INV12049 IBIHM120-21 BOLD:AEO0073'OR796114 Apidae Nomada duplex Smith, 1854# INV13264 IBIHM834-21 BOLD:AEO0073'OR796760 Apidae Nomada fallax Pérez, 1913 INV12611 IBIHM1012-22 BOLD:AEW3802'OR796010 Apidae Nomada fallax Pérez, 1913 INV12612 IBIHM1013-22 BOLD:AEW3802'OR796435 Apidae Nomada fallax Pérez, 1913 INV12844 IBIHM1245-22 BOLD:AEW3802'OR796400 Apidae Nomada fallax Pérez, 1913 INV13268 IBIHM838-21 BOLD:AEW3802'OR796892 Apidae Nomada femoralis Morawitz, 1869 INV12052 IBIHM123-21 BOLD:AAI2830 OR796271 Apidae Nomada femoralis Morawitz, 1869 INV12053 IBIHM124-21 BOLD:AEO6156 OR796559 Apidae Nomada femoralis Morawitz, 1869 INV13028 IBIHM599-21 BOLD:AEO6156 OR796372 Apidae Nomada fenestrata Lepeletier, 1841# INV12837 IBIHM1238-22 BOLD:AER8574'OR796254 Apidae Nomada fenestrata Lepeletier, 1841# INV12838 IBIHM1239-22 BOLD:AER8574'OR796509 Apidae Nomada flavoguttata (Kirby, 1802) INV12046 IBIHM117-21 BOLD:AAD4959 OR796305 Apidae Nomada flavoguttata (Kirby, 1802) INV12047 IBIHM118-21 BOLD:AAD4959 OR796404 The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 19
Family Species IBI code BOLD code BOLD BIN GenBank Apidae Nomada flavoguttata (Kirby, 1802) INV13033 IBIHM604-21 BOLD:AAD4959 OR796386 Apidae Nomada flavoguttata (Kirby, 1802) INV13209 IBIHM780-21 BOLD:AAD4959 OR796192 Apidae Nomada fulvicornis Fabricius, 1793 INV12056 IBIHM127-21 BOLD:ACE0147 OR796635 Apidae Nomada fulvicornis Fabricius, 1793 INV12057 IBIHM128-21 BOLD:ACE0147 OR796850 Apidae Nomada fulvicornis Fabricius, 1793 INV13207 IBIHM778-21 BOLD:ACE0147 OR795935 Apidae Nomada fuscicornis Nylander, 1848 INV12601 IBIHM1002-22 BOLD:AAF3571 OR795956 Apidae Nomada glaucopis Pérez, 1890 INV13263 IBIHM833-21 BOLD:AEJ1237 OR795970 Apidae Nomada glaucopis Pérez, 1890 INV12058 IBIHM129-21 BOLD:AEO3423 OR796246 Apidae Nomada glaucopis Pérez, 1890 INV12059 IBIHM130-21 BOLD:AEO3423 OR796282 Apidae Nomada glaucopis Pérez, 1890 INV12078 IBIHM149-21 BOLD:AEO3423 OR795948 Apidae Nomada glaucopis Pérez, 1890 INV12603 IBIHM1004-22 BOLD:AEO3423 OR796184 Apidae Nomada goodeniana (Kirby, 1802) INV13026 IBIHM597-21 BOLD:AEB4149 OR796347 Apidae Nomada hispanica Dusmet y Alonso, 1913# INV12060 IBIHM131-21 BOLD:AAE5753 OR796030 Apidae Nomada hispanica Dusmet y Alonso, 1913# INV13030 IBIHM601-21 BOLD:AAE5753 OR795915 Apidae Nomada illustris Schmiedeknecht, 1882 INV12061 IBIHM132-21 BOLD:AEN8381 OR796081 Apidae Nomada illustris Schmiedeknecht, 1882 INV12062 IBIHM133-21 BOLD:AEN8381 OR796134 Apidae Nomada illustris Schmiedeknecht, 1882 INV12600 IBIHM1001-22 BOLD:AEN8381 OR796058 Apidae Nomada insignipes Schmiedeknecht, 1882# INV12063 IBIHM134-21 BOLD:AEL5320 OR796742 Apidae Nomada insignipes Schmiedeknecht, 1882# INV12064 IBIHM135-21 BOLD:AEL5320 OR796730 Apidae Nomada integra Brullé, 1832 INV12065 IBIHM136-21 BOLD:ABZ1320 OR796129 Apidae Nomada integra Brullé, 1832 INV12095 IBIHM166-21 BOLD:AEN8926 OR796937 Apidae Nomada integra Brullé, 1832 INV12096 IBIHM167-21 BOLD:AEN8926 OR796270 Apidae Nomada integra Brullé, 1832 INV12097 IBIHM168-21 BOLD:AEN8926 OR796201 Apidae Nomada integra Brullé, 1832 INV13266 IBIHM836-21 BOLD:AEN8926 OR796275 Apidae Nomada integra Brullé, 1832 INV12066 IBIHM137-21 BOLD:AEO3904'OR796826 Apidae Nomada kohli Schmiedeknecht, 1882 INV12067 IBIHM138-21 BOLD:AAX4976 OR796380 Apidae Nomada kohli Schmiedeknecht, 1882 INV12090 IBIHM161-21 BOLD:AAX4976 OR796272 Apidae Nomada linsenmaieri Schwarz, 1974 INV12068 IBIHM139-21 BOLD:AAF3571 OR796021 Apidae Nomada linsenmaieri Schwarz, 1974 INV12069 IBIHM140-21 BOLD:AAF3571 OR796221 20 Wood T et al
Family Species IBI code BOLD code BOLD BIN GenBank Apidae Nomada linsenmaieri Schwarz, 1974 INV13211 IBIHM782-21 BOLD:AAF3571 OR796156 Apidae Nomada linsenmaieri Schwarz, 1974 INV13267 IBIHM837-21 BOLD:AAF3571 OR796673 Apidae Nomada maculicornis Pérez, 1884 INV12070 IBIHM141-21 BOLD:AEO4739 OR796126 Apidae Nomada marshamella (Kirby, 1802) INV09254 IBIHM067-20 BOLD:AAC4884 OR796390 Apidae Nomada melathoracica Imhoff, 1834 INV12071 IBIHM142-21 BOLD:AAX4983 OR796895 Apidae Nomada melathoracica Imhoff, 1834 INV12072 IBIHM143-21 BOLD:AAX4983 OR796185 Apidae Nomada merceti Alfken, 1909# INV12073 IBIHM144-21 BOLD:AEO1963'OR796423 Apidae Nomada merceti Alfken, 1909# INV12074 IBIHM145-21 BOLD:AEO1963'OR796917 Apidae Nomada merceti Alfken, 1909# INV12084 IBIHM155-21 BOLD:AEO1963'OR796138 Apidae Nomada merceti Alfken, 1909# INV13210 IBIHM781-21 BOLD:AEO1963'OR796735 Apidae Nomada merceti Alfken, 1909# INV13212 IBIHM783-21 BOLD:AEO1963'OR796586 Apidae Nomada merceti Alfken, 1909# INV13213 IBIHM784-21 BOLD:AEO1963'OR795932 Apidae Nomada minuscula Noskiewicz, 1930 INV12075 IBIHM146-21 BOLD:AAP1578 OR796161 Apidae Nomada mutabilis Morawitz, 1870 INV12076 IBIHM147-21 BOLD:AAX4985 OR796609 Apidae Nomada nobilis Herrich-Schäffer, 1839 INV12077 IBIHM148-21 BOLD:AAU8975 OR796447 Apidae Nomada nobilis Herrich-Schäffer, 1839 INV13262 IBIHM832-21 BOLD:AAU8975 OR796171 Apidae Nomada pectoralis Morawitz, 1877 INV12079 IBIHM150-21 BOLD:AEO0458 OR796426 Apidae Nomada piccioliana Magretti, 1883# INV12080 IBIHM151-21 BOLD:AAF3496 OR796703 Apidae Nomada piccioliana Magretti, 1883# INV12081 IBIHM152-21 BOLD:AAF3496 OR795947 Apidae Nomada rubricoxa Schwarz, 1977# INV12082 IBIHM153-21 BOLD:AEO1962'OR796120 Apidae Nomada rubricoxa Schwarz, 1977# INV12083 IBIHM154-21 BOLD:AEO1962'OR796308 Apidae Nomada rubricoxa Schwarz, 1977# INV12085 IBIHM156-21 BOLD:AEO1962'OR796117 Apidae Nomada sanguinea Smith, 1854 INV12086 IBIHM157-21 BOLD:AEN8255 OR796588 Apidae Nomada sanguinea Smith, 1854 INV12087 IBIHM158-21 BOLD:AEN8255 OR796950 Apidae Nomada sanguinea Smith, 1854 INV13271 IBIHM841-21 BOLD:AEN8255 OR796187 Apidae Nomada sexfasciata Panzer, 1799 INV12088 IBIHM159-21 BOLD:AAI2916 OR796031 Apidae Nomada sexfasciata Panzer, 1799 INV13269 IBIHM839-21 BOLD:AAI2916 OR795973 Apidae Nomada sheppardana (Kirby, 1802) INV12089 IBIHM160-21 BOLD:AAP1578 OR796595 Apidae Nomada similis Morawitz, 1872 INV12091 IBIHM162-21 BOLD:AEX4551'OR796147 Apidae Nomada stigma Fabricius, 1804 INV12092 IBIHM163-21 BOLD:ABA8671 OR796450 The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 21
Family Species IBI code BOLD code BOLD BIN GenBank Apidae Nomada striata Fabricius, 1793 INV12093 IBIHM164-21 BOLD:ABY7961 OR796506 Apidae Nomada striata Fabricius, 1793 INV12605 IBIHM1006-22 BOLD:ABY7961 OR796348 Apidae Nomada striata Fabricius, 1793 INV13206 IBIHM777-21 BOLD:ABY7961 OR795994 Apidae Nomada striata Fabricius, 1793 INV13208 IBIHM779-21 BOLD:ABY7961 OR796502 Apidae Nomada succincta Panzer, 1798 INV12094 IBIHM165-21 BOLD:ABX5010 OR796256 Apidae Nomada succincta Panzer, 1798 INV13261 IBIHM831-21 BOLD:ABX5010 OR796153 Apidae Pasites maculatus Jurine, 1807 INV12027 IBIHM098-21 BOLD:AAL3976 OR796569 Apidae Tetralonia cinctella (Saunders, 1908) INV12102 IBIHM173-21 BOLD:AEN9450'OR796857 Apidae Tetralonia dentata (Germar, 1839) INV12107 IBIHM178-21 BOLD:AEO2705'OR796915 Apidae Tetralonia fulvescens Giraud, 1863 INV12113 IBIHM184-21 BOLD:ADJ0585 OR796497 Apidae Tetralonia fulvescens Giraud, 1863 INV12112 IBIHM183-21 OR796536 Apidae Tetralonia iberica Dusmet y Alonso 1926 INV12117 IBIHM188-21 BOLD:AER1385 OR796061 Apidae Tetralonia nana Morawitz, 1874 INV12119 IBIHM190-21 BOLD:ABW6400 OR796841 Apidae Tetralonia strigata (Lepeletier, 1841) INV12125 IBIHM196-21 BOLD:AEN8168 OR796240 Apidae Thyreus hirtus (de Beaumont, 1940) INV12004 IBIHM075-21 BOLD:AEO3985 OR796533 Apidae Thyreus hirtus (de Beaumont, 1940) INV12005 IBIHM076-21 BOLD:AEO3985 OR796549 Apidae Thyreus hirtus (de Beaumont, 1940) INV12007 IBIHM078-21 BOLD:AEO3985 OR796598 Apidae Thyreus histrionicus (Illiger, 1806) INV12457 IBIHM1310-22 BOLD:AAO3396 OR796245 Apidae Thyreus histrionicus (Illiger, 1806) INV12458 IBIHM1311-22 BOLD:AAO3396 OR796238 Apidae Thyreus histrionicus (Illiger, 1806) INV12460 IBIHM1313-22 BOLD:AAO3396 OR796771 Apidae Thyreus orbatus (Lepeletier, 1841) INV12006 IBIHM077-21 BOLD:AAI0456 OR796313 Apidae Thyreus orbatus (Lepeletier, 1841) INV12010 IBIHM081-21 BOLD:AAI0456 OR796110 Apidae Thyreus orbatus (Lepeletier, 1841) INV12011 IBIHM082-21 BOLD:AAI0456 OR796498 Apidae Thyreus picaron Lieftinck, 1968 INV12461 IBIHM1314-22 BOLD:AED2570 OR796358 Apidae Thyreus ramosus (Lepeletier, 1841) INV12008 IBIHM079-21 BOLD:AAK1852 OR796224 Apidae Thyreus ramosus (Lepeletier, 1841) INV12009 IBIHM080-21 BOLD:AAK1852 OR796037 Apidae Thyreus truncatus (Pérez, 1883) INV12459 IBIHM1312-22 BOLD:AES2873 OR795902 Apidae Xylocopa cantabrita Lepeletier, 1841# INV12173 IBIHM244-21 BOLD:AEO2264 OR796867 Apidae Xylocopa cantabrita Lepeletier, 1841# INV13040 IBIHM611-21 BOLD:AEO2264 OR796858 Apidae Xylocopa iris (Christ, 1791) INV12172 IBIHM243-21 BOLD:AEO4657'OR796217 22 Wood T et al
Family Species IBI code BOLD code BOLD BIN GenBank Apidae Xylocopa iris (Christ, 1791) INV13231 IBIHM801-21 BOLD:AEO4657'OR796736 Apidae Xylocopa valga Gerstäcker, 1872 INV12171 IBIHM242-21 BOLD:AEN0425 OR796293 Apidae Xylocopa violacea (Linnaeus, 1758) INV12174 IBIHM245-21 BOLD:AAJ9209 OR796614 Apidae Xylocopa violacea (Linnaeus, 1758) INV13232 IBIHM802-21 BOLD:AAJ9209 OR796465 Colletidae Colletes abeillei Pérez, 1903 INV12347 IBIHM418-21 BOLD:ACP8014 OR796715 Colletidae Colletes abeillei Pérez, 1903 INV13153 IBIHM724-21 OR796563 Colletidae Colletes acutus Pérez, 1903 INV12348 IBIHM419-21 BOLD:AAI9265 OR796731 Colletidae Colletes albomaculatus (Lucas, 1849) INV12349 IBIHM420-21 BOLD:ABU9046 OR796089 Colletidae Colletes albomaculatus (Lucas, 1849) INV13308 IBIHM878-21 BOLD:ABU9046 OR795920 Colletidae Colletes dinizi Kuhlmann, Ortiz & Ornosa, 2001# INV12490 IBIHM1343-22 BOLD:AEU9447'OR796281 Colletidae Colletes eous Morice, 1904 INV12353 IBIHM424-21 BOLD:ADW0513 OR796015 Colletidae Colletes fodiens (Fourcroy, 1785) INV12354 IBIHM425-21 BOLD:ABZ6809 OR795931 Colletidae Colletes foveolaris Pérez, 1903 INV12484 IBIHM1337-22 BOLD:AEU9446 OR796674 Colletidae Colletes hylaeiformis Eversmann, 1852 INV13309 IBIHM879-21 BOLD:AEU8542'OR796326 Colletidae Colletes hylaeiformis Eversmann, 1852 INV12355 IBIHM426-21 BOLD:AEW0951'OR796829 Colletidae Colletes marginatus Smith, 1846 INV12358 IBIHM429-21 BOLD:AAN3911 OR796699 Colletidae Colletes mlokossewiczi Radoszkowski, 1891 INV12487 IBIHM1340-22 BOLD:ABA8594 OR796561 Colletidae Colletes nigricans Gistel, 1857 INV12357 IBIHM428-21 BOLD:AEZ9412 OR796581 Colletidae Colletes nigricans Gistel, 1857 INV12485 IBIHM1338-22 BOLD:AEZ9412 OR796377 Colletidae Colletes nigricans Gistel, 1857 INV13310 IBIHM880-21 BOLD:AEZ9412 OR796103 Colletidae Colletes pulchellus Pérez, 1903 INV12488 IBIHM1341-22 BOLD:AAN3911 OR796951 Colletidae Colletes pulchellus Pérez, 1903 INV12489 IBIHM1342-22 BOLD:AAN3911 OR796173 Colletidae Hylaeus angustatus (Schenck, 1861) INV12178 IBIHM249-21 BOLD:AAK3477 OR796750 Colletidae Hylaeus angustatus (Schenck, 1861) INV12194 IBIHM265-21 BOLD:AAK3477 OR796433 Colletidae Hylaeus angustatus (Schenck, 1861) INV13224 IBIHM795-21 BOLD:AEN9677'OR796842 Colletidae Hylaeus angustatus (Schenck, 1861) INV13316 IBIHM886-21 BOLD:AEN9677'OR796848 Colletidae Hylaeus annularis (Kirby, 1802) INV12179 IBIHM250-21 BOLD:AEK2943 OR796470 Colletidae Hylaeus annularis (Kirby, 1802) INV12180 IBIHM251-21 BOLD:AEK2943 OR796346 Colletidae Hylaeus annularis (Kirby, 1802) INV13320 IBIHM890-21 BOLD:AEK2943 OR796738 Colletidae Hylaeus clypearis (Schenck, 1853) INV12183 IBIHM254-21 BOLD:AAK3480 OR796853 The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 23
Family Species IBI code BOLD code BOLD BIN GenBank Colletidae Hylaeus communis Nylander, 1852 INV13222 IBIHM793-21 BOLD:AAE5080 OR796880 Colletidae Hylaeus confusus Nylander, 1852 INV12184 IBIHM255-21 BOLD:AAD9315 OR796100 Colletidae Hylaeus coriaceus (Pérez, 1895) INV12195 IBIHM266-21 BOLD:AEO4115'OR796508 Colletidae Hylaeus coriaceus (Pérez, 1895) INV12196 IBIHM267-21 BOLD:AEO4115'OR796623 Colletidae Hylaeus cornutus Curtis, 1831 INV12185 IBIHM256-21 BOLD:AAF4031 OR796243 Colletidae Hylaeus cornutus Curtis, 1831 INV13315 IBIHM885-21 BOLD:AAF4031 OR796231 Colletidae Hylaeus crassanus (Warncke, 1972) INV12187 IBIHM258-21 BOLD:AEN8873'OR796190 Colletidae Hylaeus difformis (Eversmann, 1852) INV12186 IBIHM257-21 BOLD:AAK3468 OR796480 Colletidae Hylaeus dilatatus (Kirby, 1802) INV12498 IBIHM1351-22 BOLD:AEU1509'OR796942 Colletidae Hylaeus dilatatus (Kirby, 1802) INV12188 IBIHM259-21 OR796510 Colletidae Hylaeus gredleri Förster, 1871 INV12190 IBIHM261-21 BOLD:AEO3886'OR796225 Colletidae Hylaeus hyalinatus Smith, 1842 INV12191 IBIHM262-21 BOLD:AAE1166 OR796593 Colletidae Hylaeus hyalinatus Smith, 1842 INV13156 IBIHM727-21 BOLD:AAE1166 OR796740 Colletidae Hylaeus imparilis Förster, 1871 INV12192 IBIHM263-21 BOLD:AAE5079 OR796253 Colletidae Hylaeus incongruus Förster, 1871 INV01584 IBIHM038-19 BOLD:ABU9205 OR796047 Colletidae Hylaeus incongruus Förster, 1871 INV12189 IBIHM260-21 BOLD:ABU9205 OR795925 Colletidae Hylaeus incongruus Förster, 1871 INV12200 IBIHM271-21 BOLD:ABU9205 OR796457 Colletidae Hylaeus incongruus Förster, 1871 INV12201 IBIHM272-21 BOLD:ABU9205 OR796935 Colletidae Hylaeus incongruus Förster, 1871 INV13154 IBIHM725-21 BOLD:ABU9205 OR796429 Colletidae Hylaeus incongruus Förster, 1871 INV13155 IBIHM726-21 BOLD:ABU9205 OR796655 Colletidae Hylaeus incongruus Förster, 1871 INV13157 IBIHM728-21 BOLD:ABU9205 OR795953 Colletidae Hylaeus incongruus Förster, 1871 INV13223 IBIHM794-21 BOLD:ABU9205 OR796339 Colletidae Hylaeus kahri Förster, 1871 INV12193 IBIHM264-21 BOLD:AAN3379 OR796084 Colletidae Hylaeus lineolatus (Schenck, 1861) INV12496 IBIHM1349-22 BOLD:AAI8781 OR796924 Colletidae Hylaeus lineolatus (Schenck, 1861) INV12497 IBIHM1350-22 BOLD:AAI8781 OR796005 Colletidae Hylaeus penalaris Dathe, 1979# INV12197 IBIHM268-21 BOLD:AEO2266'OR796414 Colletidae Hylaeus pictipes Nylander, 1852 INV12198 IBIHM269-21 BOLD:AAK3488 OR796642 Colletidae Hylaeus pictipes Nylander, 1852 INV13314 IBIHM884-21 BOLD:AAK3488 OR796490 Colletidae Hylaeus pictipes Nylander, 1852 INV13319 IBIHM889-21 BOLD:AAK3488 OR796261 Colletidae Hylaeus pictus (Smith, 1853) INV12202 IBIHM273-21 BOLD:AEO2267 OR796295 24 Wood T et al
Family Species IBI code BOLD code BOLD BIN GenBank Colletidae Hylaeus pilosulus (Pérez, 1903) INV12203 IBIHM274-21 BOLD:AAX2603 OR796362 Colletidae Hylaeus pilosulus (Pérez, 1903) INV12204 IBIHM275-21 BOLD:AAX2603 OR796658 Colletidae Hylaeus punctatus (Brullé, 1832) INV12205 IBIHM276-21 BOLD:AAE5060 OR796056 Colletidae Hylaeus punctulatissimus Smith, 1842 INV12206 IBIHM277-21 BOLD:AAK3323 OR796210 Colletidae Hylaeus signatus (Panzer, 1798) INV12207 IBIHM278-21 BOLD:AAI8577 OR796312 Colletidae Hylaeus soror (Pérez, 1903) INV12499 IBIHM1352-22 BOLD:AET8250'OR796434 Colletidae Hylaeus sulphuripes (Gribodo, 1894) INV12208 IBIHM279-21 BOLD:AEJ3211 OR796043 Colletidae Hylaeus taeniolatus Förster, 1871 INV12199 IBIHM270-21 BOLD:AEO5452'OR796300 Colletidae Hylaeus variegatus (Fabricius, 1798) INV12209 IBIHM280-21 BOLD:AAK3338 OR796896 Colletidae Hylaeus variegatus (Fabricius, 1798) INV12210 IBIHM281-21 BOLD:AAK3338 OR796050 Colletidae Hylaeus variegatus (Fabricius, 1798) INV13317 IBIHM887-21 BOLD:AAK3338 OR795903 Halictidae Ceylalictus variegatus (Olivier, 1789) INV12279 IBIHM350-21 BOLD:AAV7281 OR796006 Halictidae Ceylalictus variegatus (Olivier, 1789) INV12281 IBIHM352-21 BOLD:AAV7281 OR796925 Halictidae Dufourea gaullei Vachal, 1897# INV12275 IBIHM346-21 BOLD:AEN9216'OR796099 Halictidae Dufourea gaullei Vachal, 1897# INV12276 IBIHM347-21 BOLD:AEN9216'OR796808 Halictidae Dufourea gaullei Vachal, 1897# INV13293 IBIHM863-21 BOLD:AEN9216'OR795909 Halictidae Dufourea halictula (Nylander, 1852) INV13217 IBIHM788-21 BOLD:AAI9074 OR796349 Halictidae Dufourea trautmanni Dusmet y Alonso, 1935# INV12273 IBIHM344-21 BOLD:AEN3645 OR796622 Halictidae Dufourea trautmanni Dusmet y Alonso, 1935# INV12274 IBIHM345-21 BOLD:AEN3645 OR796922 Halictidae Halictus brunnescens (Eversmann, 1852) INV12259 IBIHM330-21 BOLD:AAE3033 OR796366 Halictidae Halictus brunnescens (Eversmann, 1852) INV12263 IBIHM334-21 BOLD:AAE3033 OR796704 Halictidae Halictus brunnescens (Eversmann, 1852) INV13054 IBIHM625-21 BOLD:AAE3033 OR796014 Halictidae Halictus confusus Smith, 1853 INV12257 IBIHM328-21 BOLD:AEN9591'OR795965 Halictidae Halictus crenicornis Blüthgen, 1923 INV12258 IBIHM329-21 BOLD:AAD5869 OR796521 Halictidae Halictus crenicornis Blüthgen, 1923 INV13055 IBIHM626-21 BOLD:AAD5869 OR796393 Halictidae Halictus fulvipes (Klug, 1817) INV02289 IBIHM013-19 BOLD:ADS5152 OR796212 Halictidae Halictus gemmeus Dours, 1872 INV01552 IBIHM032-19 BOLD:ABV4897 OR796916 Halictidae Halictus gemmeus Dours, 1872 INV01553 IBIHM005-19 BOLD:ABV4897 OR796430 The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 25
Family Species IBI code BOLD code BOLD BIN GenBank Halictidae Nomiapis diversipes (Latreille, 1806) INV13219 IBIHM790-21 BOLD:AAF6016 OR796329 Halictidae Nomiapis diversipes (Latreille, 1806) INV13273 IBIHM843-21 BOLD:AAF6016 OR796328 Halictidae Nomiapis paulyi Wood & Le Divelec, 2022 INV13272 IBIHM842-21 BOLD:ADD4589 OR796401 Halictidae Nomioides facilis (Smith, 1853) INV12280 IBIHM351-21 BOLD:AEJ1330 OR796135 Halictidae Nomioides minutissimus (Rossi, 1790) INV12282 IBIHM353-21 BOLD:AEO2657'OR796520 Halictidae Sphecodes albilabris (Fabricius, 1793) INV12283 IBIHM354-21 BOLD:AAI0130 OR796694 Halictidae Sphecodes alternatus Smith, 1853 INV12284 IBIHM355-21 BOLD:ACD7802 OR795918 Halictidae Sphecodes alternatus Smith, 1853 INV13303 IBIHM873-21 BOLD:AEO6580 OR795940 Halictidae Sphecodes crassanus Warncke, 1992 INV12285 IBIHM356-21 BOLD:ACD7802 OR796474 Halictidae Sphecodes crassanus Warncke, 1992 INV13294 IBIHM864-21 BOLD:ACD7802 OR796115 Halictidae Sphecodes croaticus Meyer, 1922 INV12286 IBIHM357-21 BOLD:ACP5320 OR796439 Halictidae Sphecodes ephippius (Linnaeus, 1767) INV12287 IBIHM358-21 BOLD:AAD0332 OR795936 Halictidae Sphecodes ephippius (Linnaeus, 1767) INV13298 IBIHM868-21 BOLD:AAD0332 OR796660 Halictidae Sphecodes gibbus (Linnaeus, 1758) INV12289 IBIHM360-21 BOLD:AAD5232 OR796455 Halictidae Sphecodes hirtellus Blüthgen, 1923# INV13117 IBIHM688-21 BOLD:AEO0791'OR796859 Halictidae Sphecodes longulus Hagens, 1882 INV13116 IBIHM687-21 BOLD:AFM8564 OR796939 Halictidae Sphecodes marginatus Hagens, 1882 INV12308 IBIHM379-21 BOLD:AAY9961 OR796732 Halictidae Sphecodes marginatus Hagens, 1882 INV13085 IBIHM656-21 BOLD:AAY9961 OR796758 Halictidae Sphecodes marginatus Hagens, 1882 INV13295 IBIHM865-21 BOLD:AAY9961 OR796911 Halictidae Sphecodes monilicornis (Kirby, 1802) INV12290 IBIHM361-21 BOLD:AAI0259 OR796749 Halictidae Sphecodes monilicornis (Kirby, 1802) INV13121 IBIHM692-21 BOLD:AAI0259 OR796438 Halictidae Sphecodes monilicornis (Kirby, 1802) INV13297 IBIHM867-21 BOLD:AAI0259 OR796947 Halictidae Sphecodes monilicornis (Kirby, 1802) INV13304 IBIHM874-21 BOLD:AAI0259 OR796125 Halictidae Sphecodes monilicornis (Kirby, 1802) INV13305 IBIHM875-21 BOLD:AAI0259 OR796204 Halictidae Sphecodes niger Hagens, 1874 INV13115 IBIHM686-21 BOLD:AAN0084 OR796179 Halictidae Sphecodes olivieri Lepeletier, 1825 INV12291 IBIHM362-21 BOLD:ACR8452 OR796589 Halictidae Sphecodes pellucidus Smith, 1845 INV12848 IBIHM1249-22 BOLD:AAI0260 OR796409 Halictidae Sphecodes pseudofasciatus Blüthgen, 1925 INV13289 IBIHM859-21 BOLD:AEO2633'OR796725 Halictidae Sphecodes puncticeps Thomson, 1870 INV12288 IBIHM359-21 BOLD:AAY9709 OR796664 Halictidae Sphecodes puncticeps Thomson, 1870 INV12293 IBIHM364-21 BOLD:AAY9709 OR796418 32 Wood T et al
Family Species IBI code BOLD code BOLD BIN GenBank Halictidae Sphecodes puncticeps Thomson, 1870 INV13299 IBIHM869-21 BOLD:AAY9709 OR796341 Halictidae Sphecodes puncticeps Thomson, 1870 INV13300 IBIHM870-21 BOLD:AAY9709 OR796556 Halictidae Sphecodes puncticeps Thomson, 1870 INV13306 IBIHM876-21 BOLD:AAY9709 OR796011 Halictidae Sphecodes reticulatus Thomson, 1870 INV12309 IBIHM380-21 BOLD:ABV3743 OR796220 Halictidae Sphecodes reticulatus Thomson, 1870 INV13296 IBIHM866-21 BOLD:ABV3743 OR796903 Halictidae Sphecodes rubicundus Hagens, 1875 INV13119 IBIHM690-21 BOLD:AAJ3388 OR796026 Halictidae Sphecodes rubicundus Hagens, 1875 INV13122 IBIHM693-21 BOLD:AAJ3388 OR796827 Halictidae Sphecodes rubicundus Hagens, 1875 INV13302 IBIHM872-21 BOLD:AAJ3388 OR796605 Halictidae Sphecodes rubripes Spinola, 1838# INV12294 IBIHM365-21 BOLD:AAM4113 OR796687 Halictidae Sphecodes ruficrus (Erichson, 1835) INV12296 IBIHM367-21 BOLD:AAF2925 OR795901 Halictidae Sphecodes ruficrus (Erichson, 1835) INV13120 IBIHM691-21 BOLD:AAF2925 OR796013 Halictidae Sphecodes ruficrus (Erichson, 1835) INV13123 IBIHM694-21 BOLD:AAF2925 OR796553 Halictidae Sphecodes ruficrus (Erichson, 1835) INV13124 IBIHM695-21 BOLD:AAF2925 OR796373 Halictidae Sphecodes ruficrus (Erichson, 1835) INV13301 IBIHM871-21 BOLD:AAF2925 OR796906 Halictidae Sphecodes rufiventris (Panzer, 1798) INV12295 IBIHM366-21 BOLD:ABA8649 OR796177 Halictidae Sphecodes spinulosus Hagens, 1875 INV12297 IBIHM368-21 BOLD:ACQ2281 OR796564 Halictidae Systropha grandimargo Pérez, 1905 INV12272 IBIHM343-21 BOLD:AEO6068'OR796633 Megachilidae Afranthidium carduele malacopygum (Gribodo, 1894) INV12447 IBIHM1300-22 BOLD:AEO1769 OR796464 Megachilidae Afranthidium carduele malacopygum (Gribodo, 1894) INV12448 IBIHM1301-22 BOLD:AEO1769 OR796832 Megachilidae Afranthidium carduele malacopygum (Gribodo, 1894) INV13349 IBIHM919-21 BOLD:AEO1769 OR796188 Megachilidae Afranthidium schulthessii (Friese, 1897) INV12211 IBIHM282-21 BOLD:AEO4048 OR796024 Megachilidae Aglaoapis tridentata (Nylander, 1848) INV12022 IBIHM093-21 BOLD:AAO2962 OR796801 Megachilidae Anthidiellum breviusculum (Pérez, 1890) INV12212 IBIHM283-21 BOLD:AEN7284 OR796051 Megachilidae Anthidiellum strigatum (Panzer, 1804) INV02280 IBIHM010-19 BOLD:AAY8402 OR796788 Megachilidae Anthidiellum strigatum (Panzer, 1804) INV12213 IBIHM284-21 BOLD:AAY8402 OR796748 Megachilidae Anthidiellum strigatum (Panzer, 1804) INV13350 IBIHM920-21 BOLD:AAY8402 OR796325 Megachilidae Anthidium cingulatum Latreille, 1809 INV12215 IBIHM286-21 BOLD:AAN9423 OR796467 Megachilidae Anthidium diadema Latreille, 1809 INV12445 IBIHM1298-22 BOLD:AER0584 OR796499 Megachilidae Anthidium diadema Latreille, 1809 INV12446 IBIHM1299-22 BOLD:AET3713 OR796149 The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 33
Family Species IBI code BOLD code BOLD BIN GenBank Megachilidae Anthidium florentinum (Fabricius, 1775) INV12214 IBIHM285-21 BOLD:ABW0328 OR796484 Megachilidae Anthidium florentinum (Fabricius, 1775) INV15674 IBIHY017-22 BOLD:ABW0328 OR795941 Megachilidae Anthidium loti Perris, 1852 INV12216 IBIHM287-21 BOLD:ACG1108 OR796007 Megachilidae Anthidium manicatum (Linnaeus, 1758) INV12217 IBIHM288-21 BOLD:AAA9946 OR796247 Megachilidae Anthidium manicatum (Linnaeus, 1758) INV13142 IBIHM713-21 BOLD:AAA9946 OR796417 Megachilidae Anthidium manicatum (Linnaeus, 1758) INV13143 IBIHM714-21 BOLD:AAA9946 OR796645 Megachilidae Anthidium oblongatum (Illiger, 1806) INV12219 IBIHM290-21 BOLD:AAC5344 OR795959 Megachilidae Anthidium punctatum Latreille, 1809 INV12218 IBIHM289-21 BOLD:AEN2704 OR796680 Megachilidae Anthidium taeniatum Latreille, 1809 INV12220 IBIHM291-21 BOLD:AEO3965 OR796160 Megachilidae Anthidium taeniatum Latreille, 1809 INV13341 IBIHM911-21 BOLD:AEO3965 OR796399 Megachilidae Chelostoma campanularum (Kirby, 1802) INV13220 IBIHM791-21 BOLD:AAB1123 OR796035 Megachilidae Chelostoma campanularum (Kirby, 1802) INV13221 IBIHM792-21 BOLD:AAB1123 OR796600 Megachilidae Chelostoma campanularum (Kirby, 1802) INV12443 IBIHM1296-22 BOLD:AET7155'OR796495 Megachilidae Chelostoma campanularum (Kirby, 1802) INV12444 IBIHM1297-22 BOLD:AET7155'OR796053 Megachilidae Chelostoma florisomne (Linnaeus, 1758) INV12442 IBIHM1295-22 BOLD:AEU4316'OR796494 Megachilidae Chelostoma rapunculi (Lepeletier, 1841) INV12400 IBIHM1253-22 BOLD:AAA7122 OR796768 Megachilidae Chelostoma rapunculi (Lepeletier, 1841) INV12401 IBIHM1254-22 BOLD:AAA7122 OR796371 Megachilidae Coelioxys acanthurus (Illiger, 1806) INV12449 IBIHM1302-22 BOLD:AET0371 OR796065 Megachilidae Coelioxys afer Lepeletier, 1841 INV12450 IBIHM1303-22 BOLD:AAV8048 OR796344 Megachilidae Coelioxys afer Lepeletier, 1841 INV13347 IBIHM917-21 BOLD:AAV8048 OR796385 Megachilidae Coelioxys argenteus Lepeletier, 1841 INV01263 IBIHM023-19 BOLD:AAV9350 OR796944 Megachilidae Coelioxys argenteus Lepeletier, 1841 INV12031 IBIHM102-21 BOLD:AAV9350 OR796524 Megachilidae Coelioxys aurolimbatus Förster, 1853 INV12452 IBIHM1305-22 BOLD:AES4301'OR796710 Megachilidae Coelioxys brevis Eversmann, 1852 INV12032 IBIHM103-21 BOLD:ADD7530 OR796712 Megachilidae Coelioxys echinatus Förster, 1853 INV12034 IBIHM105-21 BOLD:AET4410'OR796408 Megachilidae Coelioxys echinatus Förster, 1853 INV12451 IBIHM1304-22 BOLD:AET4410'OR796886 Megachilidae Coelioxys haemorrhoa Förster, 1853 INV12033 IBIHM104-21 BOLD:ADC9939 OR796175 Megachilidae Coelioxys haemorrhoa Förster, 1853 INV12453 IBIHM1306-22 BOLD:ADC9939 OR796411 Megachilidae Coelioxys haemorrhoa Förster, 1853 INV12454 IBIHM1307-22 BOLD:ADC9939 OR796918 34 Wood T et al
Family Species IBI code BOLD code BOLD BIN GenBank Megachilidae Coelioxys obtusus Pérez, 1884 INV12455 IBIHM1308-22 BOLD:AEO4483'OR795979 Megachilidae Coelioxys obtusus Pérez, 1884 INV13345 IBIHM915-21 BOLD:AEO4483'OR796752 Megachilidae Coelioxys obtusus Pérez, 1884 INV13346 IBIHM916-21 BOLD:AEO4483'OR796683 Megachilidae Dioxys cinctus (Jurine, 1807) INV12017 IBIHM088-21 BOLD:AAY8179 OR795977 Megachilidae Dioxys cinctus (Jurine, 1807) INV12018 IBIHM089-21 BOLD:AAY8179 OR796094 Megachilidae Dioxys pumilus Gerstäcker, 1869 INV12019 IBIHM090-21 BOLD:AEO4333'OR796459 Megachilidae Dioxys pumilus Gerstäcker, 1869 INV12020 IBIHM091-21 BOLD:AEO4333'OR796017 Megachilidae Dioxys pumilus Gerstäcker, 1869 INV12021 IBIHM092-21 BOLD:AEO4333'OR796503 Megachilidae Ensliniana bidentata (Friese, 1899)# INV12016 IBIHM087-21 BOLD:AEN9289'OR796930 Megachilidae Heriades crenulata Nylander, 1856 INV12579 IBIHM550-21 BOLD:AAO0527 OR796208 Megachilidae Heriades crenulata Nylander, 1856 INV13355 IBIHM925-21 BOLD:AAO0527 OR795921 Megachilidae Heriades rubicola Pérez, 1890 INV12578 IBIHM549-21 BOLD:ABV4003 OR796690 Megachilidae Heriades rubicola Pérez, 1890 INV13354 IBIHM924-21 BOLD:ABV4003 OR796343 Megachilidae Heriades truncorum (Linnaeus, 1758) INV12596 IBIHM567-21 BOLD:AAI8422 OR796706 Megachilidae Heriades truncorum (Linnaeus, 1758) INV13125 IBIHM696-21 BOLD:AAI8422 OR796003 Megachilidae Heriades truncorum (Linnaeus, 1758) INV13356 IBIHM926-21 BOLD:AAI8422 OR796811 Megachilidae Hoplitis acuticornis (Dufour & Perris, 1840) INV12386 IBIHM457-21 BOLD:AAK6030 OR796711 Megachilidae Hoplitis adunca (Panzer, 1798) INV12421 IBIHM1274-22 BOLD:AAI1794 OR796834 Megachilidae Hoplitis albiscopa (Friese, 1899)# INV12411 IBIHM1264-22 BOLD:AEO0400'OR796864 Megachilidae Hoplitis albiscopa (Friese, 1899)# INV12412 IBIHM1265-22 BOLD:AEO0400'OR796350 Megachilidae Hoplitis albiscopa (Friese, 1899)# INV13368 IBIHM938-21 BOLD:AEO0400'OR796606 Megachilidae Hoplitis annulata (Latreille, 1811) INV12383 IBIHM454-21 BOLD:ADD6906 OR796334 Megachilidae Hoplitis annulata (Latreille, 1811) INV13366 IBIHM936-21 BOLD:ADD6906 OR796278 Megachilidae Hoplitis anthocopoides (Schenck, 1853) INV12422 IBIHM1275-22 BOLD:AAC3950 OR796540 Megachilidae Hoplitis antigae (Pérez, 1895) INV12416 IBIHM1269-22 BOLD:AEN2131 OR796920 Megachilidae Hoplitis antigae (Pérez, 1895) INV12599 IBIHM570-21 BOLD:AEN2131 OR796927 Megachilidae Hoplitis antigae (Pérez, 1895) INV13369 IBIHM939-21 BOLD:AEN2131 OR796276 Megachilidae Hoplitis benoisti (Alfken, 1935) INV12423 IBIHM1276-22 BOLD:AEV6823'OR796146 Megachilidae Hoplitis bisulca (Gerstäcker, 1869) INV12588 IBIHM559-21 BOLD:AEO0529 OR796955 Megachilidae Hoplitis brachypogon (Pérez, 1879)# INV12393 IBIHM464-21 BOLD:AEP1421 OR796219 The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 35
Family Species IBI code BOLD code BOLD BIN GenBank Megachilidae Hoplitis brachypogon (Pérez, 1879)# INV13367 IBIHM937-21 BOLD:AEP1421 OR795944 Megachilidae Hoplitis cadiza (Warncke, 1991)# INV12419 IBIHM1272-22 BOLD:AER8784'OR796351 Megachilidae Hoplitis cadiza (Warncke, 1991)# INV12420 IBIHM1273-22 BOLD:AER8784'OR796941 Megachilidae Hoplitis campanularis (Morawitz, 1879)# INV12413 IBIHM1266-22 BOLD:AES6061'OR796849 Megachilidae Hoplitis campanularis (Morawitz, 1879)# INV12414 IBIHM1267-22 BOLD:AES6061'OR796517 Megachilidae Hoplitis claviventris (Thomson, 1872) INV12402 IBIHM1255-22 BOLD:AAE5480 OR796781 Megachilidae Hoplitis cristatula (van der Zanden, 1990) INV12593 IBIHM564-21 BOLD:AEL6177 OR796087 Megachilidae Hoplitis grumi (Morawitz, 1894)# INV12403 IBIHM1256-22 BOLD:AET3653'OR796482 Megachilidae Hoplitis leucomelana (Kirby, 1802) INV12398 IBIHM469-21 BOLD:AEL2158 OR796262 Megachilidae Hoplitis marchali (Pérez, 1902) INV12404 IBIHM1257-22 BOLD:AET3654'OR796375 Megachilidae Hoplitis ochraceicornis (Ferton, 1902)# INV12424 IBIHM1277-22 BOLD:AER7475'OR796745 Megachilidae Hoplitis ochraceicornis (Ferton, 1902)# INV12425 IBIHM1278-22 BOLD:AER7475'OR796551 Megachilidae Hoplitis praestans (Morawitz, 1894) INV12415 IBIHM1268-22 BOLD:AER7476'OR796883 Megachilidae Hoplitis ravouxi (Pérez, 1902) INV12417 IBIHM1270-22 BOLD:AAI1840 OR796297 Megachilidae Hoplitis ravouxi (Pérez, 1902) INV12418 IBIHM1271-22 BOLD:AET2330'OR796284 Megachilidae Hoplitis stecki (Frey-Gessner, 1908) INV02288 IBIHM012-19 BOLD:ACF9884 OR796747 Megachilidae Hoplitis stecki (Frey-Gessner, 1908) INV12577 IBIHM548-21 BOLD:ACF9884 OR796934 Megachilidae Hoplitis tridentata (Dufour & Perris, 1840) INV12592 IBIHM563-21 BOLD:AAF2246 OR796545 Megachilidae Icteranthidium grohmanni (Spinola, 1838) INV12221 IBIHM292-21 BOLD:AEH0395 OR796554 Megachilidae Icteranthidium grohmanni (Spinola, 1838) INV13353 IBIHM923-21 BOLD:AEH0395 OR796612 Megachilidae Lithurgus chrysurus Fonscolombe, 1834 INV12234 IBIHM305-21 BOLD:AAY7690 OR796128 Megachilidae Lithurgus tibialis Morawitz, 1875 INV12235 IBIHM306-21 BOLD:ABW1174 OR796131 Megachilidae Megachile albisecta (Klug, 1817) INV12237 IBIHM308-21 BOLD:AEO5749 OR796529 Megachilidae Megachile albisecta (Klug, 1817) INV02293 IBIHM043-19 BOLD:AEO5750 OR796820 Megachilidae Megachile albonotata Radoszkowski, 1886# INV12236 IBIHM307-21 BOLD:AAY7863 OR796644 Megachilidae Megachile apicalis Spinola, 1808 INV12238 IBIHM309-21 BOLD:AAF4159 OR796843 Megachilidae Megachile apicalis Spinola, 1808 INV13127 IBIHM698-21 BOLD:AAF4159 OR796073 Megachilidae Megachile apicalis Spinola, 1808 INV13342 IBIHM912-21 BOLD:AAF4159 OR796453 36 Wood T et al
Family Species IBI code BOLD code BOLD BIN GenBank Megachilidae Megachile argentata (Fabricius, 1793) INV03735 IBIHM048-19 BOLD:AAD7803 OR796463 Megachilidae Megachile argentata (Fabricius, 1793) INV12437 IBIHM1290-22 BOLD:AAD7803 OR796168 Megachilidae Megachile argentata (Fabricius, 1793) INV12438 IBIHM1291-22 BOLD:AAD7803 OR796062 Megachilidae Megachile argentata (Fabricius, 1793) INV13129 IBIHM700-21 BOLD:AAD7803 OR796759 Megachilidae Megachile argentata (Fabricius, 1793) INV13343 IBIHM913-21 BOLD:AAD7803 OR796717 Megachilidae Megachile argentata (Fabricius, 1793) INV13348 IBIHM918-21 BOLD:AAD7803 OR795928 Megachilidae Megachile argentata (Fabricius, 1793) INV15662 IBIHY006-22 BOLD:AAD7803 OR796548 Megachilidae Megachile argentata (Fabricius, 1793) INV15675 IBIHY018-22 BOLD:AAD7803 OR796040 Megachilidae Megachile centuncularis (Linnaeus, 1758) INV12239 IBIHM310-21 BOLD:AAA8356 OR796591 Megachilidae Megachile centuncularis (Linnaeus, 1758) INV13130 IBIHM701-21 BOLD:AAA8356 OR796214 Megachilidae Megachile centuncularis (Linnaeus, 1758) INV15682 IBIHY024-22 BOLD:AAA8356 OR795976 Megachilidae Megachile circumcincta (Kirby, 1802) INV12240 IBIHM311-21 BOLD:AAC0049 OR796424 Megachilidae Megachile ericetorum Lepeletier, 1841 INV12241 IBIHM312-21 BOLD:AEA9090 OR796315 Megachilidae Megachile giraudi Gerstäcker, 1869 INV12242 IBIHM313-21 BOLD:AAY7848 OR796814 Megachilidae Megachile giraudi Gerstäcker, 1869 INV12250 IBIHM321-21 BOLD:AAY7848 OR795983 Megachilidae Megachile lagopoda (Linnaeus, 1761) INV12243 IBIHM314-21 BOLD:AAE8874 OR796800 Megachilidae Megachile leachella Curtis, 1828 INV12244 IBIHM315-21 BOLD:AAD2767 OR796799 Megachilidae Megachile leachella Curtis, 1828 INV13126 IBIHM697-21 BOLD:AAD2767 OR796560 Megachilidae Megachile lefebvrei (Lepeletier, 1841) INV12245 IBIHM316-21 BOLD:ABW1753 OR796585 Megachilidae Megachile marginata Smith, 1853 INV12256 IBIHM327-21 BOLD:AAK6960 OR796666 Megachilidae Megachile maritima (Kirby, 1802) INV12246 IBIHM317-21 BOLD:AAD7888 OR796336 Megachilidae Megachile melanopyga Costa, 1863 INV12247 IBIHM318-21 BOLD:AAE8870 OR796688 Megachilidae Megachile melanopyga Costa, 1863 INV13128 IBIHM699-21 BOLD:AAE8870 OR796692 Megachilidae Megachile octosignata Nylander, 1852 INV12248 IBIHM319-21 BOLD:ADM5545 OR796519 Megachilidae Megachile opacifrons Pérez, 1897 INV12249 IBIHM320-21 BOLD:ACF9800 OR796342 Megachilidae Megachile pusilla Pérez, 1884 INV12254 IBIHM325-21 BOLD:AAQ0516 OR796440 Megachilidae Megachile pusilla Pérez, 1884 INV15709 IBIHY032-22 BOLD:AAQ0516 OR796806 Megachilidae Megachile pyrenaica Lepeletier, 1841 INV12251 IBIHM322-21 BOLD:AAO0637 OR795950 Megachilidae Megachile rotundata (Fabricius, 1787) INV12252 IBIHM323-21 BOLD:AEO2625'OR796888 The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 37
Family Species IBI code BOLD code BOLD BIN GenBank Megachilidae Megachile sicula (Rossi, 1792) INV12253 IBIHM324-21 BOLD:AEO4733 OR796310 Megachilidae Megachile thevestensis Ferton, 1909# INV12439 IBIHM1292-22 BOLD:AEU0213'OR796095 Megachilidae Megachile versicolor Smith, 1844 INV12440 IBIHM1293-22 BOLD:AAD5414 OR796286 Megachilidae Megachile versicolor Smith, 1844 INV12441 IBIHM1294-22 BOLD:AAD5414 OR796813 Megachilidae Megachile willughbiella (Kirby, 1802) INV12255 IBIHM326-21 BOLD:ACE6545 OR796784 Megachilidae Osmia anceyi Pérez, 1879# INV12435 IBIHM1288-22 BOLD:AEO4345'OR795934 Megachilidae Osmia anceyi Pérez, 1879# INV12436 IBIHM1289-22 BOLD:AEO4345'OR796078 Megachilidae Osmia anceyi Pérez, 1879# INV13361 IBIHM931-21 BOLD:AEO4345'OR796194 Megachilidae Osmia andrenoides Spinola, 1808 INV12585 IBIHM556-21 BOLD:AEO0032'OR796613 Megachilidae Osmia argyropyga Pérez, 1879# INV12582 IBIHM553-21 BOLD:AEO4346'OR796318 Megachilidae Osmia aurulenta (Panzer, 1799) INV12580 IBIHM551-21 BOLD:AAE5409 OR796218 Megachilidae Osmia aurulenta (Panzer, 1799) INV13131 IBIHM702-21 BOLD:AAE5409 OR796819 Megachilidae Osmia bicornis (Linnaeus, 1758) INV12434 IBIHM1287-22 BOLD:AAD6282 OR796570 Megachilidae Osmia bicornis (Linnaeus, 1758) INV12590 IBIHM561-21 BOLD:AAD6282 OR796571 Megachilidae Osmia bicornis (Linnaeus, 1758) INV13140 IBIHM711-21 BOLD:AAD6282 OR796206 Megachilidae Osmia bicornis (Linnaeus, 1758) INV13141 IBIHM712-21 BOLD:AAD6282 OR796069 Megachilidae Osmia brevicornis (Fabricius, 1798) INV12586 IBIHM557-21 BOLD:AEF6206 OR796722 Megachilidae Osmia caerulescens (Linnaeus, 1758) INV01272 IBIHM024-19 BOLD:AAD0313 OR796932 Megachilidae Osmia caerulescens (Linnaeus, 1758) INV12389 IBIHM460-21 BOLD:AAD0313 OR796122 Megachilidae Osmia caerulescens (Linnaeus, 1758) INV13132 IBIHM703-21 BOLD:AAD0313 OR796264 Megachilidae Osmia caerulescens (Linnaeus, 1758) INV13133 IBIHM704-21 BOLD:AAD0313 OR796121 Megachilidae Osmia caerulescens (Linnaeus, 1758) INV13136 IBIHM707-21 BOLD:AAD0313 OR796831 Megachilidae Osmia caerulescens (Linnaeus, 1758) INV13362 IBIHM932-21 BOLD:AAD0313 OR796700 Megachilidae Osmia caerulescens (Linnaeus, 1758) INV13364 IBIHM934-21 BOLD:AAD0313 OR796822 Megachilidae Osmia cephalotes Morawitz, 1870 INV12584 IBIHM555-21 BOLD:AEN9982'OR796902 Megachilidae Osmia cyanoxantha Pérez, 1879# INV12387 IBIHM458-21 BOLD:AEO1610'OR796769 Megachilidae Osmia dimidiata Morawitz, 1870 INV12583 IBIHM554-21 BOLD:AEO3684 OR796840 Megachilidae Osmia emarginata Lepeletier, 1841 INV12589 IBIHM560-21 BOLD:AAE4126 OR796550 Megachilidae Osmia ferruginea Latreille, 1811 INV12390 IBIHM461-21 BOLD:AEN8200'OR796956 Megachilidae Osmia gallarum Spinola, 1808 INV12388 IBIHM459-21 BOLD:AAO8736 OR796919 38 Wood T et al
Family Species IBI code BOLD code BOLD BIN GenBank Megachilidae Osmia iberica van der Zanden, 1987# INV12581 IBIHM552-21 BOLD:AEN8862'OR796166 Megachilidae Osmia labialis Pérez, 1879 INV12591 IBIHM562-21 BOLD:ADY8979 OR796634 Megachilidae Osmia labialis Pérez, 1879 INV13360 IBIHM930-21 BOLD:ADY8979 OR796812 Megachilidae Osmia latreillei (Spinola, 1806) INV12397 IBIHM468-21 BOLD:AAZ7870 OR796226 Megachilidae Osmia leaiana (Kirby, 1802) INV12433 IBIHM1286-22 BOLD:AAI1846 OR795900 Megachilidae Osmia ligurica Morawitz, 1868 INV12391 IBIHM462-21 BOLD:ADH8891 OR796157 Megachilidae Osmia ligurica Morawitz, 1868 INV13134 IBIHM705-21 BOLD:ADH8891 OR796542 Megachilidae Osmia lunata Benoist, 1928# INV12594 IBIHM565-21 BOLD:AEO3079'OR796181 Megachilidae Osmia melanogaster Spinola, 1808 INV12392 IBIHM463-21 BOLD:AEN4264 OR796787 Megachilidae Osmia niveata (Fabricius, 1804) INV12399 IBIHM470-21 BOLD:AAP2416 OR796451 Megachilidae Osmia niveata (Fabricius, 1804) INV13137 IBIHM708-21 BOLD:AAP2416 OR796054 Megachilidae Osmia niveata (Fabricius, 1804) INV13138 IBIHM709-21 BOLD:AAP2416 OR796289 Megachilidae Osmia niveata (Fabricius, 1804) INV13363 IBIHM933-21 BOLD:AAP2416 OR796743 Megachilidae Osmia niveocincta Pérez, 1879# INV12394 IBIHM465-21 BOLD:AEO5031'OR796088 Megachilidae Osmia niveocincta Pérez, 1879# INV12432 IBIHM1285-22 BOLD:AEO5031'OR796215 Megachilidae Osmia rufohirta Latreille, 1811 INV12396 IBIHM467-21 BOLD:AEO0004'OR796661 Megachilidae Osmia scutellaris Morawitz, 1868 INV12428 IBIHM1281-22 BOLD:AEJ4270 OR795981 Megachilidae Osmia scutellaris Morawitz, 1868 INV12429 IBIHM1282-22 BOLD:AEJ4270 OR796066 Megachilidae Osmia signata Erichson, 1839 INV12395 IBIHM466-21 BOLD:AEA2838 OR796369 Megachilidae Osmia signata Erichson, 1839 INV13139 IBIHM710-21 BOLD:AEA2838 OR796012 Megachilidae Osmia submicans Morawitz, 1870 INV12385 IBIHM456-21 BOLD:AAK5820 OR796167 Megachilidae Osmia submicans Morawitz, 1870 INV13135 IBIHM706-21 BOLD:AAK5820 OR796821 Megachilidae Osmia tergestensis Ducke, 1897 INV12426 IBIHM1279-22 BOLD:AES2567 OR796151 Megachilidae Osmia tergestensis Ducke, 1897 INV12427 IBIHM1280-22 BOLD:AES2567 OR796693 Megachilidae Osmia tricornis Latreille, 1811 INV12587 IBIHM558-21 BOLD:AEN7876 OR796603 Megachilidae Osmia uncicornis Pérez, 1895# INV12430 IBIHM1283-22 BOLD:AET6037'OR796119 Megachilidae Osmia uncicornis Pérez, 1895# INV12431 IBIHM1284-22 BOLD:AET6037'OR796632 Megachilidae Osmia versicolor Latreille, 1811 INV12384 IBIHM455-21 BOLD:AAZ7638 OR796491 Megachilidae Protosmia asensioi Griswold & Parker, 1987# INV12409 IBIHM1262-22 BOLD:AET0069'OR796232 Megachilidae Protosmia capitata (Schletterer, 1889)# INV12595 IBIHM566-21 BOLD:AEO6065'OR796234 The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 39
Family Species IBI code BOLD code BOLD BIN GenBank Megachilidae Protosmia exenterata (Pérez, 1895)# INV12407 IBIHM1260-22 BOLD:ACL7801 OR796481 Megachilidae Protosmia exenterata (Pérez, 1895)# INV12408 IBIHM1261-22 BOLD:ACL7801 OR796034 Megachilidae Protosmia glutinosa (Giraud, 1871)# INV12405 IBIHM1258-22 BOLD:AET0068'OR796626 Megachilidae Protosmia glutinosa (Giraud, 1871)# INV12406 IBIHM1259-22 BOLD:AET0068'OR796923 Megachilidae Pseudoanthidium eximium (Giraud, 1863) INV12222 IBIHM293-21 BOLD:AEO0467 OR796544 Megachilidae Pseudoanthidium melanurum (Klug, 1832) INV12223 IBIHM294-21 BOLD:AEN8472'OR796397 Megachilidae Pseudoanthidium reticulatum (Mocsáry, 1884) INV12226 IBIHM297-21 BOLD:AEO1983 OR796518 Megachilidae Pseudoanthidium scapulare (Latreille, 1809) INV12224 IBIHM295-21 BOLD:AAK2762 OR796572 Megachilidae Pseudoanthidium scapulare (Latreille, 1809) INV13351 IBIHM921-21 BOLD:AAK2762 OR796139 Megachilidae Pseudoanthidium scapulare (Latreille, 1809) INV13352 IBIHM922-21 BOLD:AAK2762 OR796446 Megachilidae Pseudoanthidium stigmaticorne (Dours, 1873) INV12225 IBIHM296-21 BOLD:ABV1020 OR795943 Megachilidae Rhodanthidium infuscatum (Erichson, 1839) INV12233 IBIHM304-21 BOLD:AEO5272 OR796410 Megachilidae Rhodanthidium septemdentatum (Latreille, 1809) INV12232 IBIHM303-21 BOLD:AAY5674 OR796686 Megachilidae Rhodanthidium siculum (Spinola, 1838) INV12230 IBIHM301-21 BOLD:AEA0375 OR796863 Megachilidae Rhodanthidium sticticum (Fabricius, 1787) INV12231 IBIHM302-21 BOLD:AEH2459 OR796679 Megachilidae Stelis breviuscula (Nylander, 1848) INV12026 IBIHM097-21 BOLD:ACE4655 OR796532 Megachilidae Stelis breviuscula (Nylander, 1848) INV13357 IBIHM927-21 BOLD:ACE4655 OR796105 Megachilidae Stelis nasuta (Latreille, 1809) INV12000 IBIHM071-21 BOLD:AEC4732 OR796907 Megachilidae Stelis ortizi M.Schwarz & Gusenleitner, 2010# INV12023 IBIHM094-21 BOLD:AEO2242'OR796249 Megachilidae Stelis phaeoptera (Kirby, 1802) INV12024 IBIHM095-21 BOLD:AAY9796 OR796535 Megachilidae Stelis punctulatissima (Kirby, 1802) INV12025 IBIHM096-21 BOLD:AAJ4972 OR795972 Megachilidae Trachusa byssina (Panzer, 1798) INV12228 IBIHM299-21 BOLD:AAF1587 OR796792 Megachilidae Trachusa interrupta (Fabricius, 1781) INV12229 IBIHM300-21 BOLD:ACW1366 OR796154 Megachilidae Trachusa laeviventris (Dours, 1873) INV12227 IBIHM298-21 BOLD:AEN7816 OR796437 Melittidae Dasypoda cingulata Erichson, 1835 INV12480 IBIHM1333-22 BOLD:ACK7071 OR796071 Melittidae Dasypoda cingulata Erichson, 1835 INV12481 IBIHM1334-22 BOLD:ACK7071 OR796577 40 Wood T et al
Family Species IBI code BOLD code BOLD BIN GenBank Melittidae Dasypoda cingulata Erichson, 1835 INV13148 IBIHM719-21 BOLD:ACK7071 OR796207 Melittidae Dasypoda crassicornis Friese, 1896 INV12366 IBIHM437-21 BOLD:AEO3711 OR795961 Melittidae Dasypoda crassicornis Friese, 1896 INV13311 IBIHM881-21 BOLD:AEO3711 OR796874 Melittidae Dasypoda crassicornis Friese, 1896 INV13313 IBIHM883-21 BOLD:AEO3711 OR796331 Melittidae Dasypoda dusmeti Quilis, 1928 INV12363 IBIHM434-21 BOLD:ACK5966 OR796505 Melittidae Dasypoda dusmeti Quilis, 1928 INV13144 IBIHM715-21 BOLD:ACK5966 OR796584 Melittidae Dasypoda dusmeti Quilis, 1928 INV13152 IBIHM723-21 BOLD:ACK5966 OR796682 Melittidae Dasypoda hirtipes (Fabricius, 1793) INV12365 IBIHM436-21 BOLD:AEO2254'OR796810 Melittidae Dasypoda hirtipes (Fabricius, 1793) INV13145 IBIHM716-21 BOLD:AEO2254'OR796001 Melittidae Dasypoda hirtipes (Fabricius, 1793) INV13151 IBIHM722-21 BOLD:AEO2254'OR796845 Melittidae Dasypoda morotei Quilis, 1928 INV12362 IBIHM433-21 BOLD:AEN9179 OR796897 Melittidae Dasypoda morotei Quilis, 1928 INV12478 IBIHM1331-22 BOLD:AEN9179 OR796496 Melittidae Dasypoda morotei Quilis, 1928 INV12479 IBIHM1332-22 BOLD:AEN9179 OR795926 Melittidae Dasypoda morotei Quilis, 1928 INV13312 IBIHM882-21 BOLD:AEN9179 OR796109 Melittidae Dasypoda pyrotrichia Förster, 1855 INV12476 IBIHM1329-22 BOLD:AES3843'OR796042 Melittidae Dasypoda radchenkoi Ghisbain & Wood, 2023 INV12361 IBIHM432-21 BOLD:AEO4511'OR796794 Melittidae Dasypoda radchenkoi Ghisbain & Wood, 2023 INV12477 IBIHM1330-22 BOLD:AEO4511'OR796019 Melittidae Dasypoda visnaga (Rossi, 1790) INV12367 IBIHM438-21 BOLD:AEO0075 OR796650 Melittidae Macropis fulvipes (Fabricius, 1804) INV12360 IBIHM431-21 BOLD:AAJ1462 OR795995 Melittidae Macropis fulvipes (Fabricius, 1804) INV13321 IBIHM891-21 BOLD:AAJ1462 OR796427 Melittidae Melitta leporina (Panzer, 1799) INV12368 IBIHM439-21 BOLD:AAF4411 OR796838 The BOLD BIN system uses algorithms to cluster sequences into operational taxonomic units (OTUs) that closely correspond to species (Ratnasingham and Hebert 2013). A total of 535 BINs were retrieved by BOLD (Ratnasingham and Hebert 2007) and 112 BINs are unique to our dataset (Table 1, Suppl. material 1). Sequences from two specimens were not attributed to BINs as their sequences were only 418 or 326 base pairs in length (Suppl. material 1). To our knowledge, this is the first study to systematically focus on DNA barcoding bees in the Iberian Peninsula. This study shows that DNA barcode sequences, based on the COI mitochondrial gene fragment, can be highly useful in identifying Iberian bee samples to species level with 486 of the 514 species (94.9%) assigned to unique BINs. However, 27 species were assigned to at least two BINS (Table 2); these cases are discussed below. The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 41
central (Castelo Branco) Portugal which fell into BOLD:AEO5653, with this BIN also containing three sequences from Spain (Granada, Málaga, Toledo) generated by Wood (2023). Barcoding members of the subgenus Chrysandrena Hedicke, 1933, to which Andrena hesperia belongs, does not appear to be straightforward, with multiple mitochondrial lineages present in addition to those presented here (Wood, unpublished data). The three specimens from southern Portugal (Beja) were collected from the same site on the same day, with two mitochondrial lineages present and no apparent morphological differences. Further study is required, but based on morphology, the judgement is made that, at the present time, there is no justification to split up A. hesperia. Andrena limata Smith, 1853 and A. thoracica (Kirby, 1802) Andrena (Melandrena) species often present complex COI data (Wood 2023). Specimens identified in our study as Andrena limata and Andrena thoracica were found to share a BIN (BOLD:AAE1815) and A. limata sequences were also placed in an additional BIN (BOLD:AEX3903). These two BINs were separated by a maximum genetic distance of 3.77%. Though we did not sequence any Iberian specimens of A. nitida (Müller, 1766), these three species combined present serious taxonomic problems regarding their consistent recognition across their range due to variable expression of morphological characters. Analysis of additional sequences from inside and outside of Iberia (Wood 2023) identified five mitochondrial clades covering these three nominal species, one for A. thoracica, one for A. nitida and three for A. limata. From the data we present here, the A. limata/A. thoracica BIN BOLD:AAE1815 contains sequences from Portugal, Spain, Italy, Germany, Austria, Poland, Romania, Turkey, Lebanon, Israel and Kazakhstan and corresponds to the limata #1 + thoracica lineages of Wood (2023). The A. limata BIN BOLD:AEX3903 contains sequences from Morocco, Portugal, Spain and France and corresponds to the limata #2 lineage of Wood (2023). Though none of the data we presented fell into this BIN, an additional BIN BOLD: AEW6788 is available containing A. limata/A. nitida sequences from Spain, France, Switzerland, United Kingdom, Belgium, the Netherlands, Germany, Austria, Finland and Russia and corresponding to the limata #3 + nitida lineages of Wood (2023). The five clades of Wood (2023), therefore, fall into three BINs, but without agreement with the existing morphological concepts. In this context, the results presented here fit into this broader pattern of complex COI data. It is clear that more powerful genetic techniques are required to resolve species boundaries in this difficult group of species and, thus, no action is taken here. Andrena morio Brullé, 1832 Sequences of Andrena morio produced two BINs, with specimens from central Portugal (Castelo Branco) falling into both BINs, which were minimally separated by a maximum genetic distance of 2.64%. Andrena morio produces a range of barcodes, with a broad species concept followed by Wood (2023) who synonymised the predominantly Iberian A. hispania Warncke, 1967 with it. The genetic distances reported here are considered to 48 Wood T et al
represent variation when considered across the entire range of A. morio and the genetic differentiation that it presents (Wood 2023). Andrena propinqua Schenck, 1853 Andrena propinqua has been a somewhat controversial taxon and has been placed either in synonymy with or as a subspecies of A. dorsata (Kirby, 1802) in the past. The current evidence supports its species status, based on analysis of Ultra-conserved Elements (Gueuning et al. 2020), as barcodes alone are insufficient to allow for consistent separation. Sequences from central Portugal (Castelo Branco and Coimbra) and southern Spain (Málaga) fell into BOLD:AAJ2115 which contains a mixture of A. dorsata and A. propinqua sequences from across Europe, whereas sequences from central and northern Portugal (Aveiro, Castelo Branco, Coimbra) fell into BOLD:AEN9473 which currently contains five sequences from Iberia only. The two BINs were separated by a maximum genetic distance of 5.41%. Based on morphological analysis, A. dorsata is restricted in Iberia to northern Spain, from Galicia across the Cantabrian mountains to the Pyrenees (Wood 2023). Given the complexity present, no taxonomic action is taken, as these variable barcodes are considered to represent only variation, following the study of Gueuning et al. (2020). Andrena russulaLepeletier, 1841 Praz et al. (2022) recently resolved many classification issues within the subgenus Taeniandrena Hedicke, 1933, including for Andrena russula. Sequences of the true A. russula were generated from central Portugal (Castelo Branco, Santarém), falling into BOLD:AAZ1205, along with sequences from Portugal, Spain, Morocco and Italy. More broadly, these sequences correspond to the A. russula of Praz et al. (2022) and the A. russula #1 of Wood (2023). However, in southern Portugal (Faro, Setubal), this mitochondrial lineage is replaced by BOLD:AEN8931 which is strongly separated from the previous BIN (maximum genetic distance of 5.60%) and which conforms to the "sp. nov. 2" lineage of Praz et al. (2022) and the A. russula #2 lineage of Wood (2023). Morphologically, there are no apparent differences between the two lineages. Interestingly, BOLD:AEN8931 contains four currently unpublished sequences from France. Taxonomic revision is required to determine how to recognise this divergent mitochondrial lineage morphologically, as it likely represents yet another cryptic and undescribed Taeniandrena species in the west Mediterranean (Wood et al. 2021, Wood 2022, Praz et al. 2022). Given the occurrence in France, this lineage should be present also in southern and eastern Spain. No concrete action is taken here. Anthidium diademaLatreille, 1809 Two specimens of Anthidium diadema, both from Madrid, were separated by a genetic distance of 2.81% and consequently fell into separate BINs. The two specimens were male and female, so a comparison of morphological differences is not possible. No further action is taken here without further genetic sampling. The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 49
Anthophora plumipes (Pallas, 1772) Anthophora plumipes is the most abundant anthophorine species in the West Palaearctic. Specimens from central Portugal (Coimbra) and southern Spain (Cádiz) fell into BOLD:AEN8725 which so far contains sequences from Iberia only. A specimen from northern Portugal (Braga) produced a sequence which fell into BOLD:AEO1358 which contains sequences from across the West Palaearctic. Whilst the sequences were moderately separated (maximum genetic distance of 5.86%), in a broader analysis of samples of A. plumipes from across the West Palaearctic (Wood, Boustani & Rasmont, unpublished data), no clear geographically consistent clades could be observed. Moreover, the morphology of specimens was consistent across this range, with variation predominantly comprising hair colour. A broad species concept is, therefore, retained for A. plumipes. Chelostoma campanularum (Kirby, 1802) Two specimens from northern Portugal (Viseu) fell into BOLD:AAB1123 which contains typical Chelostoma campanularum sequences from across Europe (and Canada where the species is introduced). However, two specimens from southern Spain (Granada, Málaga) fell into BOLD:AET7155 which was separated by a maximum genetic distance of 4.25%. These divergent sequences merit further study, as the taxonomy of the small-bodied Chelostoma species around C. campanularum (subgenus Foveosmia) is challenging; no action is taken at the present time. Colletes hylaeiformis Eversmann, 1852 Specimens from central Portugal (Castelo Branco, BOLD:AEU8542) and central Spain (Ávila, BOLD:AEW0951) were assigned to different BINS (maximum genetic distance 4.70%). Both BINs contained only a single sequence and neither BIN matched BOLD:AAJ7533 which is the BIN for Colletes hylaeiformis sequences from Austria, France, Germany and Hungary. Further study is required to establish if Iberian material is simply genetically differentiated due to isolation by distance or whether these differences are more substantive. Flavipanurgus granadensis(Warncke, 1987) Flavipanurgus granadensis was described with a locus typicus in the Sierra Nevada (Granada) in southern Spain. Sequences from south-western Spain (Cádiz, BOLD:AEO48 46) and south-eastern Spain (Murcia, BOLD:AEO4847) fell into distinct BINs and were separated by a maximum genetic distance of 5.93%. The BOLD:AEO4846 BIN additionally contains three sequences from Málaga and Cuenca (Wood & Boustani, unpublished data). Given the taxonomic complexity within Flavipanurgus (Cross and Wood 2018), it is not impossible that the specimen from Murcia represents an additional cryptic species, but dedicated study is necessary, as well as genotyping the population present on the Sierra Nevada. 50 Wood T et al
Halictus maculatus Smith, 1848 Halictus maculatus is a widespread species and a sequence from northern Portugal (Porto) fell into BOLD:ACH4344 which contains sequences from Austria, France, Italy and Russia. However, a sequence from southern Spain (Granada) fell into BOLD:AEO0160 which is a unique BIN, separated by a maximum genetic distance of 3.6%. The specimen is a female and it is not impossible that it represents the Spanish endemic species H. toparensis OrtizSánchez & Pauly, 2017 which was described from a single male from southern Spain (Almería) and which is morphologically close to H. maculatus (Ortiz-Sánchez and Pauly 2017). This species has recently been found to be more widespread in central Spain (Álvarez Fidalgo et al. 2023) and so is presence also in Granada is possible. Further barcoding of H. toparensis males is required in order to definitively associate the female sex for this species. It should also be noted that none of our material fell into BOLD:AAY53 83 which is the principal European BIN for H. maculatus (by number of sequences and geographical distribution), containing sequences from Austria, France, Italy, Germany and Switzerland. Further study is required. Hoplitis ravouxi(Pérez, 1902) Hoplitis ravouxi is a widespread European species. A sequence from eastern Spain (Teruel, BOLD:AAI1840) fell into a BIN with sequences from France and Germany. A sequence from southern Spain (Málaga, BOLD:AET2330) was separated by a maximum genetic distance of 2.64%, with the resulting BIN being unique. Further study is required, as this southern sequence may simply represent separation by distance. Hylaeus angustatus(Schenck, 1861) Four specimens of Hylaeus angustatus were separated in two BINs, one that we consider to be the true H. angustatus (BOLD:AAK3477) containing specimens from northern and eastern Spain (León, Teruel, as well as other sequences from France, Italy, Germany and Austria) and the other containing specimens from Portugal (Castelo Branco, Viseu; BIN known from Iberia only) considered to be what was reported from Iberia as H. angustatus punctifrons (Pérez, 1903) (BOLD:AEN9677) that is not currently recognised at the species level (Ghisbain & Rosa et al. 2023). However, this BIN was strongly separated genetically from Hylaeus angustatus by a high genetic distance of 6.84%. Hylaeus angustatus is a morphologically and genetically very variable species that most probably represents a species complex currently including two described species, H. angustatus and H. mariannae Theunert, 2013, this latter species being represented by an additional highly divergent BIN (Le Divelec, unpublished data) and which cannot be associated with Iberian material morphologically or genetically. The morphological boundaries of the two Iberian forms of H. angustatus are still unclear and further work is needed to clarify the taxonomy and nomenclature of this species group. Lasioglossum calceatum (Scopoli, 1763) For this extremely common and widespread species, sequences from northern Portugal (Aveiro, Viana do Castelo, BOLD:AAB0353) fell into the typical BIN for this species with The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 51
hundreds of sequences from across the Palaearctic. However, a sequence from southern Spain (Málaga, BOLD:AEO1453) was placed in a unique BIN which was actually genetically closer to L. albipes (Fabricius, 1781). This BIN was separated from our typical L. calceatum BIN by a maximum genetic distance of 6.55%. Both L. calceatum and L. albipes are very rare in southern Iberia (Ortiz-Sánchez and Pauly 2017) and are more or less restricted to mountains. The specimen producing the unique BIN was collected at high altitude in the Sierra de las Nieves (between 1200 and 1600 m) and may represent an isolated population of L. calceatum or perhaps an undescribed species. Further study is required. Lasioglossum leucozonium (Schrank, 1781) and Lasioglossum leucozonium cedri Ebmer, 1976 Lasioglossum leucozonium is a widespread Palaearctic species and a sequence from northern Portugal (Aveiro, BOLD:AAA2322) fell into the typical L. leucozonium BIN with hundreds of sequences from Europe to Central Asia, as well as North America where it is introduced. However, sequences from northern and central Portugal (Aveiro, Leiria) and southern Spain (Granada) fell into BOLD:AEN9620 which contains sequences from Portugal, Spain and Morocco. This BIN was separated by a maximum genetic distance of 6.42% and represents the taxon ssp. cedri which is not currently recognised at species level. A revision of the subgenus Leuchalictus Warncke, 1975 to which these taxa belong is currently in preparation (S. Flaminio, pers. comm.) and so no taxonomic action is taken here. Lasioglossum malachurum (Kirby, 1802) A sequence from southern Spain (Málaga, BOLD:AAE5496) fell into the typical L. malachurum BIN with hundreds of sequences from the West Palaearctic. Surprisingly, a specimen from northern Portugal (Aveiro, BOLD:AEO8786) was strongly separated by a maximum genetic distance of 13.66%, though it was closest to the typical L. malachurum BIN. Further study is required. Lasioglossum transitorium(Schenck, 1868) and L. transitorium planulum (Pérez, 1903) Lasioglossum transitorium planulum is the western subspecies of L. transitorium and is found across Iberia (Ortiz-Sánchez and Pauly 2017). Sequences from northern Portugal (Aveiro) and eastern and southern Spain (Cuenca, Granada) fell into BOLD:AEM8844 that contains sequences from only Spain and Portugal. However, a specimen from northeastern Spain (Lleida) fell into BOLD:AER6557 (containing additional sequences from France and Switzerland) which was separated by a maximum genetic distance of 2.01%. Detailed study is required to establish whether these represent the taxa L. transitorium planulum and L. transitorium s. str., respectively and whether they should be treated as distinct species or not. 52 Wood T et al
Lasioglossum xanthopus (Kirby, 1802) and L. xanthopus soreli (Dours, 1872) A sequence from southern Spain (Granada, BOLD:AAE1789) fell into the typical L. xanthopus BIN with sequences from Belgium, France, Germany, Kyrgyzstan, the Netherlands, Switzerland, Turkey and the United Kingdom. A sequence from central Portugal (Castelo Branco) fell into BOLD:AEW1421 which contains one other sequence from Morocco (Wood, unpublished data) and which was separated by a maximum genetic distance of 2.01%. Further study is required to establish whether this material represents a distinct species and whether it could be referrable to the concept of L. xanthopus soreli (for which the type material is lost). Lasioglossum soreli was described from Algeria and, although the type material is lost, if further genetic analysis of Iberian and North African xanthopus sensu lato is conducted, the taxon soreli can be newly delineated. Megachile albisecta (Klug, 1817) Sequences from southern Spain (Granada, BOLD:AEO5749) and northern Portugal (Bragança, BOLD:AEO5750) were separated by a maximum genetic distance of 2.17%. Both BINs also contained sequences from France and Italy in addition to the Spanish sequences. Given the low genetic distance, this is considered to simply represent variation. Nomada basalisHerrich-Schäffer, 1839 This morphologically highly variable species was placed in three BINs, specifically specimens from central Portugal (Castelo Branco, BOLD:AEK6178, sequences also from France and Italy), eastern Spain (Cuenca, BOLD:AEN3462, Iberia only) and central Portugal (Lisbon) and central and southern Spain (Madrid, Málaga, BOLD:AEO4155, Iberia only). The maximum gentic distance between members of these BINs was 5.71%. BOLD:AEK6178 also contains sequences from France and Italy, but the two other BINs are currently known only from Iberia. Nomada basalis is the brood-parasite of Eucera (Hetereucera) species, such as E. elongatula Vachal, 1907 (Baldock et al. 2018); this subgenus is diversified and species-rich in Iberia, offering many potential hosts for N. basalis and possibly unknown related taxa. A large number of names are available for N. basalis as, due to its morphological variation, it was described many times (Smit 2018). Taxonomic revision is required to establish whether N. basalis represents a species complex or whether it is simply a highly variable species. Nomada distinguendaMorawitz, 1874 A sequence from northern Portugal (Aveiro, BOLD:ACY0250) clustered with N. distinguenda from France, Italy and the Netherlands. However, an additional sequence from northern Portugal (Bragança, BOLD:AET5764) formed a unique BIN. The maximum genetic distance between these BINs was 3.6%. Importantly, none of the Iberian material fell into BOLD:AAO6546 which contains N. distinguenda sequences from Bulgaria, France, Germany and Hungary or BOLD:AEA0257 which contains N. distinguenda sequences from Austria, Israel and Italy. Detailed taxonomic revision is clearly required to resolve species concepts within N. distinguenda given the genetic variation visible here. The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 53
Nomada femoralis Morawitz, 1869 A sequence from a female specimen from southern Spain (Granada, BOLD:AAI2830) clustered with N. femoralis sequences from Austria, France, Germany, Italy and Morocco. However, specimens from northern and southern Portugal (Aveiro, Faro) and central Spain (Ávila, BOLD:AEO6156) formed a distinct BIN, with an additional specimen from Morocco (Wood, unpublished data). The maximum genetic distance between the two BINs was 7.05%. This result is surprising, as N. femoralis is nominally one of the easiest European Nomada species to recognise in the male sex due to the form of shape of the anterior femorae which are produced ventrally into long triangular shapes (Smit 2018). The male specimen IBIHM124-21 from Ávila morphologically appears inseparable from typical N. femoralis from Central Europe, presenting these modified anterior femorae. Adding complexity, the two female specimens that fell into this BIN were originally identified as N. femoralis (IBIHM599-21, northern Portugal, Aveiro) and N. corcyraea Schmiedeknecht, 1882 (IBIHM1246-22, southern Portugal, Faro). The morphological characters allowing recognition of the females are therefore unclear and a revision of this group of species is required. The aberrant "femoralis" BIN (BOLD:AEO6156) is genetically closest to N. armata Herrich-Schäffer, 1839 and is strongly differentiated from typical N. femoralis, at least based on these barcoding results. Nomada glaucopis Pérez, 1890 Most of the sequences we generated from across Spain (Granada, Guadalajara, Madrid, Málaga, BOLD:AEO3423) formed a BIN with additional sequences from France. However, a single sequence from central Portugal (Castelo Branco, BOLD:AEJ1237) was separated by a maximum genetic distance of 3.11%. Further study is required. Nomada integra Brullé, 1832 Nomada integra sequences were placed into three BINs, specifically with sequences from central Spain (Ávila, BOLD:ABZ1320), central Portugal (Castelo Branco) and southern Spain (Málaga, BOLD:AEN8926) and central Spain (Segovia, BOLD:AEO3904). The maximum genetic distance between these BINs was 3.43%. The sequence from Ávila (BOLD:ABZ1320) probably represents the true N. integra, with additional sequences from Austria, Belgium, Germany and Greece. The sequences from Portugal and southern Spain form a BIN (BOLD:AEN8926) with additional sequences from France and the sequence from Segovia (BOLD:AEO3904) forms a unique BIN. Nomada integra is a brood-parasite of members of the Andrena subgenus Chlorandrena Pérez, 1890 (Smit 2018), a subgenus that is highly diversified in Iberia (Wood 2023). Several lineages of N. integra may exist, possibly representing distinct species, all of which attack different Chlorandrena species. Detailed study is required to demonstrate this; for now, our data are presented simply as N. integra. Sphecodes alternatus Smith, 1853 and S. crassanus Warncke, 1992 Warncke (1992) described S. crassanus as distinct from S. alternatus, a position that has been largely followed by subsequent authors (Amiet et al. 2007, Bogusch and Straka 2012, 54 Wood T et al
Ghisbain et al. 2023b). In the female sex, S. crassanus can be separated from S. alternatus due to the denser punctation anterior to the ocellar triangle, which is slightly convex and more sparsely punctate in S. alternatus (Bogusch and Straka 2012). However, two specimens of S. crassanus showed minimal genetic differentiation from a specimen of S. alternatus despite this clear morphological difference and were placed in the same BIN (BOLD:ACD7802). Furthermore, an additional specimen of "S. alternatus" from central Portugal was strongly separated genetically from both S. crassanus + S. alternatus. This latter Portuguese specimen represents S. algeriensis Alfken, 1914 (considered as a subspecies of S. alternatus by Warncke (1992)), based on its morphology with a more densely punctate frons than typical S. alternatus and also because the BIN ( BOLD: AEO6580) includes an unpublished sequence from Egypt; S. algeriensis is not currently recognised as a good species and focused taxonomic work is required to formally establish its status. The maximum genetic distance between the two BINs was 7.97%. Species that cannot currently be identified A single Melecta specimen from the Sierra Nevada (Granada, IBIHM072-21) fell into a unique BIN (BOLD:AEO2855) containing another unpublished sequence from Spain. The taxonomy of melectine bees is complicated and a full molecular revision is required to ensure that morphological concepts (Lieftinck 1980) actually work consistently given the extreme difficulty in species identification in this group. Confirmed or suspected synonymies Eucera longicornis (Linnaeus, 1758) and Eucera hispaliensis Pérez, 1902 Dorchin (2023) concluded that E. hispaliensis was a junior synonym of the widespread E. longicornis. In line with this, sequences of "E. hispaliensis" generated from Portugal (Castelo Branco, Vila Real) and Spain (Granada, Málaga) fell into BOLD:ABZ4790, which contains E. longicornis sequences from across Europe. The synonymy of Dorchin (2023) is, therefore, further supported by this genetic evidence. Eucera nigrescens Pérez, 1879 and Eucera codinai Dusmet y Alonso, 1926 It is suspected that E. codinai is a junior synonym of the widespread E. nigrescens (E. Dorchin, pers. comm.). The Iberian distribution appears to be that E. nigrescens is present in the north, being replaced in central and southern Iberia by E. codinai (Baldock et al. 2018). Morphological differences are minor and are predominantly based on changes in hair colour. Analysis of "E. codinai" sequences from southern Spain (Cádiz) and central Portugal (Castelo Branco) showed that they fell into BOLD:ABZ2397 which contains many sequences of E. nigrescens from across Europe eastwards to Japan. All our data are, therefore, presented as E. nigrescens and the formal synonymy of E. codinai with E. nigrescens is likely to be made in the future following type revision. The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 55
Different species assigned to the same BIN Finally, in addition to the nominal species which were assigned to multiple BINs, there are a few species that unexpectedly fell into the same BIN despite some morphological differences. These are highlighted here. Colletes marginatusSmith, 1846 and Colletes pulchellus Pérez, 1903 These two species are challenging to separate morphologically and old records of C. marginatus from Iberia can be impossible to assign to either C. marginatus or C. pulchellus without a morphological re-examination of the specimens in question (Baldock et al. 2018, M. Kuhlmann, pers. comm.). Three specimens nominally belonging to C. marginatus and C. pulchellus produced sequences that fell into the same BIN (BOLD:AAN3911). The specimens comprised two specimens identified as C. pulchellus (IBIHM1342-22; one male from Granada, IBIHM1341-22; one female from Girona; both det. M. Kuhlmann) and one specimen identified as C. marginatus (IBIHM429-21; one female from Segovia, det. M. Kuhlmann). This BIN contains specimens from Germany, the Netherlands, Finland, Norway, Spain and France and, thus, would certainly include sequences of the true C. marginatus which was described from Britain. Further study is required, as several Colletes species are very difficult to recognise through DNA barcoding alone, such as the succinctus-group (Kuhlmann et al. 2007, Zenz et al. 2021). Nomada sheppardana (Kirby, 1802) and Nomada minuscula Noskiewicz, 1930 The species status of N. minuscula has proven controversial, more and more authors have recognised N. minuscula as distinct (e.g. Scheuchl 2000, Amiet et al. 2007, Schmidt et al. 2015, Scheuchl and Willner 2016, Smit 2018, Ghisbain et al. 2023b) . Schmidt et al. (2015) found that specimens belonging to these two species were assigned to two BINs - BOLD:AAP1578 for N. minuscula and BOLD:ABA2390 for N. sheppardana - and, hence, the authors supported a position of two distinct species. Analysis of one female of "N. sheppardana" from central Spain (IBIMH160-21; Ávila) and one female of "N. minuscula" from southern Spain (IBIHM146-21; Málaga) produced sequences that fell into the same BIN (BOLD:AAP1578), i.e. the BIN of N. minuscula only. The two specimens clearly differ morphologically and conform to the morphological concepts of N. sheppardana and N. minuscula. The same problem occurs in other parts of Europe, with all Swiss members of this species pair producing the same barcode and falling into the N. minuscula BIN (Praz et al. 2023: Figure 7). It is not clear why this is occurring or why specimens of "N. sheppardana" from Germany represent a distinct BIN. Dedicated taxonomic work is required, including generating sequences from the United Kingdom, from which N. sheppardana was originally described. Hylaeus incongruus Förster, 1871 The taxonomy of the Hylaeus gibbus-group has a long and confused history (Dathe 1980, Straka and Bogusch 2011, Le Divelec 2022). Barcoding specimens of this group from Iberia produced sequences that fell into two BINs corresponding to Hylaeus confusus 56 Wood T et al
Nylander, 1852 (BOLD:AAD9315) and H. incongruus (BOLD:ABU9205) in Central Europe. This caveat is important, as the scale of the analysis can impact interpretation, for example, when including data from the east Mediterranean (Le Divelec, unpublished data). We sequenced one specimen morphologically identified as H. confusus from northern Spain (León) and this sequence clearly clusters with H. confusus sequences from across Europe (France, Switzerland, Belgium, the Netherlands, Ireland, United Kingdom, Germany, Austria, Denmark, Norway and Finland). In this context, the identification and genetic identity of H. confusus in Iberia does not appear to be problematic. However, the situation with the remaining sequences generated from members of the gibbus-group is more challenging. A total of eight sequences were generated from across central and northern Iberia. Two sequences from north-western Iberia (INV12201; Bragança and INV12200; Ourense) were generated from specimens displaying redmarkings on tergum 1 and were originally identified as H. praenotatus Förster, 1871. However, H. praenotatus as reported from Iberia by Ortiz-Sánchez et al. (2002) and Baldock et al. (2018) are H. praenotatus auctorum. True H. praenotatus is a synonym of H. gibbus Saunders, 1850 (Le Divelec 2022, see also Ghisbain et al. 2023b) which does not appear to occur in Iberia. The two sequences fell into the H. incongruus BIN BOLD: ABU9205 (additional sequences from Morocco, France, Switzerland, Italy, Belgium, the Netherlands, Germany, Austria, Czechia, Norway, Finland, Greece, Cyprus, Lebanon and Egypt). Morphologically, these H. praenotatus auctorum specimens do not meaningfully differ from H. incongruus morphologically, except in the colour of tergum 1. Sequencing of six additional specimens with entirely dark terga from Portugal (Aveiro, Bragança, Viana do Castelo and Viseu) and Spain (Lleida) that were originally identified as H. gibbus produced sequences that also fell into the same H. incongruus BIN BOLD: ABU9205. Therefore, none of the eight specimens originally identified as either H. praenotatus or H. gibbus genetically matched with H. gibbus as currently delineated genetically (BIN BOLD:AAE1200, sequences from France, Switerland, Belgium, Italy, Germany, Austria and Hungary). There was a slight, but consistent genetic difference between the red-marked and dark specimens. This dark clade represents H. purpurissatus (Vachal, 1895), a taxon which was recently recognised at a specific level (Straka and Bogusch 2011, Le Divelec 2022). However, the morphological differences are small and inconsistent and may not represent good species (Le Divelec, unpublished data). Given these challenges associated with morphological delineation with species within the gibbusgroup and these molecular results, we present all eight sequences from Iberia as H. incongruus; dedicated study of the gibbus-group across the Mediterranean Basin is required. Project description Title: The InBIO Barcoding Initiative Database: DNA Barcodes of Iberian bees The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 57
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Suppl. material 3: DS-IBIHY01 IBI Bees of Iberia library - Darwin Core Standard Authors: Thomas Wood, Hugo Gaspar, Romain Le Divelec, Andreia Penado, Teresa Luisa Silva, Vanessa Mata, Joana Veríssimo, Deniz Michez, Silvia Castro, João Loureiro, Pedro Beja, Sónia Ferreira Data type: Record information - specimen data in Darwin Core Standard format Brief description: The file includes information about all records in BOLD for the DS-IBIHY01 IBI Bees of Iberia library. It contains collection and identification data. Download file (413.16 kb) Suppl. material 4: NJ tree of bee DNA barcodes Authors: Thomas Wood, Hugo Gaspar, Romain Le Divelec, Andreia Penado, Teresa Luisa Silva, Vanessa Mata, Joana Veríssimo, Deniz Michez, Silvia Castro, João Loureiro, Pedro Beja, Sónia Ferreira Data type: Phylogenetic tree Brief description: NJ tree of all Bee DNA barcodes generated in the InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees. Download file (44.67 kb) Suppl. material 5: BOLD Systems Hitlist of Portuguese Bees Authors: Thomas Wood, Sónia Ferreira Data type: Species list Brief description: List of bee species with occurrence known in Portugal, but lacking specimens and/or sequences and/or DNA barcodes in BOLD database on 8 February 2024. Download file (11.51 kb) The InBIO Barcoding Initiative Database: DNA barcodes of Iberian Bees 71