Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2017-02-26
Page range: 41–77
Abstract views: 244
PDF downloaded: 3

Updated list of bumblebees (Hymenoptera: Apidae) from the Spanish Pyrenees with notes on their decline and conservation status 

Departamento de Zoología y Antropología Física. Facultad de Biología. Universidad Complutense. c/ José Antonio Nováis, 12. 28040 Madrid, Spain.
Departamento de Biología Animal, Ecología, Parasitología y Edafología. Universidad de Salamanca. Campus Miguel de Unamuno s/n. 37071 Salamanca, Spain.
Departamento Zoología y Antropología Física, Facultad de Veterinaria, Campus de Espinardo, Universidad de Murcia, 30100 Murcia, Spain.
Hymenoptera Biodiversity Bombus decline habitat loss pollinators Spain

Abstract

The Pyrenees, where Euro-Siberian, Mediterranean and alpine faunas join together, have a large biodiversity of bumblebees. We compiled historical literature records of bumblebee species from the Spanish Pyrenees, and then compared these to contemporary surveys to assess trends in elevational distribution. Twenty-eight species (including thirty-five subspecies) were found in the contemporary survey. Nine species and two subspecies previously present were not detected, some included on the Spanish Red List. With the exception of a few species, a reduction of the altitudinal range and an orophilous tendency was observed at both upper and lower elevational levels, suggesting an upward trend towards better-preserved high areas. Our results reinforce the need to develop new protection programs and more restrictive conservation measures for bumblebee populations, species and their habitats.

 

References

  1. Ascher, J.S. & Pickering, J. (2015) Discover Life bee species guide and world checklist (Hymenoptera: Apoidea: Anthophila). Available from: http://wwworkersdiscoverlife.org/mp/20q?guide=Apoidea species (accessed 6 June 2016)

    Barea-Azcón, J.M., Ballesteros, E. & Moreno, D. (2008) Libro Rojo de Invertebrados de Andalucía. Consejería de Medio Ambiente (Junta de Andalucía), Sevilla, 1430 pp.

    Bofill, J.M. (1905) Catàlech de Insectes de Catalunya. Hymenòpteres. Apidae. Butlletí de la Institusio Catalana de Historia Natural, 1905, 1–72.

    Bommarco, R., Lundin, O., Smith, H.G. & Rundlöf, M. (2012) Drastic historic shifts in bumblebee community composition in Sweden. Proceedings of the Royal Society B, 279, 309–315.
    https://doi.org/10.1098/rspb.2011.0647

    Bosch, M. (1999) Biologia de la reproducció de la tribu Delphinieae a la Mediterrània occidental. Institut d’Estudis Catalans, Barcelona, 376 pp.

    Brown, M.J.F. & Paxton, R.J. (2009) The conservation of bees: a global perspective. Apidologie, 40, 410–416.
    https://doi.org/10.1051/apido/2009019

    Cameron, S.A., Hines, H.M. & Williams, P.H. (2007) A comprehensive phylogeny of the bumblebees (Bombus). Biological Journal of the Linnean Society, 91, 161–188.
    https://doi.org/10.1111/j.1095-8312.2007.00784.x

    Cameron, S.A., Lozier, J.D., Strange, J.P., Koch, J.B, Cordes, N., Leellen, C., Solter, L.F. & Griswold, T.L. (2011) Patterns of widespread decline in North American bumble bees. Proceedings of the National Academy of Sciences, 108 (2), 662–667.
    https://doi.org/10.1073/pnas.1014743108

    Cariveau, D.P., Powell, J.E., Koch, H., Winfree, R. & Moran, N.A. (2014) Variation in gut microbial communities and its association with pathogen infection in wild bumble bees (Bombus). The ISME Journal, 8, 2369–2379.
    https://doi.org/10.1038/ismej.2014.68

    Castro, L. (1988) Sobre los Bombinae (Hymenoptera: Apidae) de las Sierras de Cazorla, Segura y Alcaraz (Spain) (I). Anales de Biología, 15, 95–100.

    Castro, L. (1993) Sobre los Bombinae (Hymenoptera, Apidae) de la Sierra de Albarracín (I). Zapateri Revista aragonesa de Entomología, 3, 5–14.

    Castro, L. (1994) Sobre los Bombinae (Hymenoptera, Apidae) de la Sierra de Albarracín (II). Zapateri Revista aragonesa de entomología, 4, 137–144.

    Castro, L. (1996) Insecta, Hymenoptera, 1: Apidae s.s. Catálogos de la entomofauna Aragonesa, 9, 3–6.

    Castro, L. (2006) Nuevos datos sobre la distribución ibérica de Bombus gerstaeckeri Morawitz, 1881 (Hymenoptera: Apoidea: Apidae) y primera cita para Castilla y León (Spain). Boletín de la Sociedad Entomológica Aragonesa, 38, 106.

    Castro, L. & Ornosa, C. (1985) Nuevos datos sobre Bombus (Pyrobombus) pratorum (Linneo, 1761) (Hym., Apidae). Boletín de la Asociación Española de Entomología, 9, 389.

    Cepero, A., Ravoet, J., Gómez-Moracho, T., Bernal, J.L., Del Nozal, M.J., Bartolomé, C., Maside, X., Meana, A., González-Porto, A.V., de Graaf, D.C., Martín-Hernández, R. & Higes, M. (2014) Holistic screening of collapsing honey bee colonies in Spain: a case study. BMC Research Notes, 7, 649.
    https://doi.org/10.1186/1756-0500-7-649

    Clough,Y., Ekroos, J., Báldi, A., Batáry, P., Bommarco, R., Gross, N., Holzschuh, A., Hopfenmüller, S., Knop, E., Kuussaari, M., Lindborg, R., Marini, L.,Öckinger, E., Potts, S.G., Pöyry, J., Roberts, S.P.M., Steffan-Dewenter, I. & Smith, H.G. (2014) Density of insect-pollinated grassland plants decreases with increasing surrounding land-use intensity. Ecology Letters, 17, 1168–1177.
    https://doi.org/10.1111/ele.12325

    Colla, S.R., Gadallah, F., Richardson, L., Wagner, D. & Gall, L. (2012) Assessing declines of North American bumble bees (Bombus spp.) using museum specimens. Biodiversity and Conservation, 21, 3585–3595.
    https://doi.org/10.1007/s10531-012-0383-2

    Condamine, F.L. & Hines, H.M. (2015) Historical species losses in bumblebee evolution. Biology Letters, 11, 20141049.
    https://doi.org/10.1098/rsbl.2014.1049

    CORINE Biotopes Project (1989) An Inventory of Sites of Importance for Nature Conservation in the European Community. Corine Land Cover, 1989, 1–2000.

    Delmas, R. (1976) Contribution à l´etude de la fauna française des Bombidae (Hymenoptera, Apoidea, Bombidae). Annales de la Société entomologique de France, New Series, 12, 247–290.

    Dillon, M.E. & Dudley, R. (2014) Surpassing Mt. Everest: extreme flight performance of alpine bumble-bees. Biology Letters, 10, 20130922.
    https://doi.org/10.1098/rsbl.2013.0922

    Dupont, Y.L., Damgaard, C. & Simonsen, V. (2011) Quantitative Historical Change in Bumblebee (Bombus spp.) Assemblages of Red Clover Fields. PLoS ONE, 6, e25172.
    https://doi.org/10.1371/journal.pone.0025172

    Edwards, M. & Williams, P.H. (2004) Where have all the bumblebees gone, and could they ever return? British Wildlife, 15, 305–312.

    Fillat, F., Aguirre, A.J., Pauné, F. & Fondevilla, C. (2012) La conservación de la montaña alpina y el bienestar humano. In: Evaluación de los Ecosistemas del Milenio en España EME 2012. Conservación de los servicios de los ecosistemas y la biodiversidad para el bienestar humano. Fundación Biodiversidad, Ministerio de Agricultura, Alimentación y Medio Ambiente, pp. 116–132. Available from: http://www.ecomilenio.es/wp-content/uploads/2012/04/AMBIENTA_98web.pdf (accessed 11 January 2017)

    Frankham, R. (2010) Challenges and opportunities of genetic approaches to biological conservation. Biological Conservation, 143, 1919–1927.
    https://doi.org/10.1016/j.biocon.2010.05.011

    Fürst, M.A., McMahon, D.P., Osborne, J.L., Paxton, R.J. & Brown, M.J.F. (2014) Disease associations between honeybees and bumblebees as a threat to wild pollinators. Nature, 506, 364–373.
    https://doi.org/10.1038/nature12977

    García, D., Quevedo, M., Obeso, J.R. & Abajo, A. (2004) Fragmentation patterns and protection of montane forest in the Cantabrian Range (NW Spain). Forest Ecology and Management, 208, 29–43.
    https://doi.org/10.1016/j.foreco.2004.10.071

    Goulson, D. (2009) Bumblebees. Oxford University Press, Croydon, 317 pp.

    Goulson, D. (2010) Impacts of non-native bumblebees in Western Europe and North. Applied Entomology and Zoology, 45, 7–12.
    https://doi.org/10.1303/aez.2010.7

    Goulson, D., Hanley, M.E., Darvill, B., Ellis, J.S. & Knight, M.E. (2005) Causes of rarity in bumblebees. Biological Conservation, 122, 1–8.
    https://doi.org/10.1016/j.biocon.2004.06.017

    Goulson, D., Lye, G.C. & Darvill, B. (2008a) Diet breadth, coexistence and rarity in bumblebees. Biodiversity Conservation, 17, 3269–3288.
    https://doi.org/10.1007/s10531-008-9428-y

    Goulson, D., Lye, G.C. & Darvill, B. (2008b) Decline and conservation of bumble bees. Annual Review of Entomology, 53, 191–208.
    https://doi.org/10.1146/annurev.ento.53.103106.093454

    Goulson, D., Nicholls, E., Botías, C. & Rotheray, E.L. (2015) Bee declines driven by combined stressfrom parasites, pesticides, and lack of flowers. Sciencexpress, 26, 1–16.

    Greystock, P., Blaneb, E.J., McFrederick, E.J.Q., Goulson, D. & Hughes, W.H. (2015) Do managed bees drive parasite spread and emergence in wild bees? International Journal for Parasitology: Parasites and Wildlife, 5 (1), 64–75.
    https://doi.org/10.1016/j.ijppaw.2015.10.001

    Groom, S., Stevens, M. & Schwarz, M.P. (2014) Parallel responses of bees to Pleistocene climate change in three isolated archipelagos of the southwestern Pacific. Proceedings of the Royal Society of London B: Biological Sciences, 281, 20133293.
    https://doi.org/10.1098/rspb.2013.3293

    Herrera, J.M., Ploquin, E., Rodríguez-Pérez, J. & Obeso, J.R. (2014) Determining habitat suitability for bumblebees in a mountain system: a baseline approach for testing the impact of climate change on species presence and abundance. Journal of Biogeography, 41, 700–712.
    https://doi.org/10.1111/jbi.12236

    Iserbyt, S., Durieux, E.A. & Rasmont, P. (2008) The remarkable diversity of bumblebees (Hymenoptera: Apidae: Bombus) in the Eyne Valley (France, Pyrénées–Orientales). Annales de la Société Entomologique de France, 44, 211–241.
    https://doi.org/10.1080/00379271.2008.10697558

    IUCN (2001) IUCN Red List Categories and Criteria. Version 3.1. IUCN, Gland, ii + 30 pp.

    IUCN (2013) The IUCN Red list of Threatened Species. Available from: http://www.iucnredlist.org/details/13341899/0 (accessed 16 January 2016)

    Kerr, J.T., Pindar, A., Galpern, P., Packer, L., Potts, S.G., Roberts, S.M., Rasmont, P., Schweiger, O., Colla, S.R., Richardson, L.L., Wagner, D.L., Gall, L.F., Sikes, D.S. & Pantoja, A. (2015) Climate change impacts on bumblebees converge across continents. Science, 349 (6244), 177–180.
    https://doi.org/10.1126/science.aaa7031

    La Greca, M. (1990) The Insect Biogeography of West Mediterranean Islands. In: International Symposium on Biogeographical aspects of insularity. Accademia Nazionale dei Lincei, Roma, pp. 469–491.

    Lecocq, T., Lhomme, P., Michez, D., Dellicour, S., Valterova, I. & Rasmont, P. (2011) Molecular and chemical characters to evaluate species status of two cuckoo bumblebees: Bombus barbutellus and Bombus maxillosus (Hymenoptera, Apidae, Bombini). Systematic Entomology, 36, 453–469.
    https://doi.org/10.1111/j.1365-3113.2011.00576.x

    Løken, A. (1984) Scandinavian species of the genus Psithyrus Lepeletier (Hymenoptera: Apidae). Entomologica Scandinavica Supplement, 23, 16438,

    Manino, A., Patetta, A., Porporato, M., Quaranta, M., Intoppa, F., Piazza, M.G. & Frilli, F. (2007) Bumblebee (Bombus Latreille, 1802) distribution in high mountains and global warming. Redia, 90, 125–129.

    Matthew, P.R, Tinsley, C., Brown, M.J.F. & Goulson, D. (2013) Investigating the impact of deploying commercial Bombus terrestris for crop pollination on pathogen dynamics in wild bumble bees. Journal of Apicultural Research, 52, 149–157.
    https://doi.org/10.3896/IBRA.1.52.3.06

    Meeus, I., Brown, M.J.F. & De Graaf, D.C. (2011) Effects of invasive parasites on Bumble Bee declines. Conservation Biology, 25, 662–671.
    https://doi.org/10.1111/j.1523-1739.2011.01707.x

    Miller-Struttmann, N.E., Geib, J.C., Franklin, J.D., Kevan, P.G., Holdo, R.M., Ebert-May, D., Lynn, A.M., Kettenbach, J.A., Hedrick, E. & Galen, C. (2015) Functional mismatch in a bumble bee pollination mutualism under climate change. Science, 349 (6255), 1541–1544.
    https://doi.org/10.1126/science.aab0868

    Obeso, J.R. (1992) Geographic distribution and community structure of bumblebees in the Northern Iberian Peninsula. Oecologia, 89, 244–252.
    https://doi.org/10.1007/BF00317224

    Ornosa, C. (1984) La subfamilia Bombinae (Hym., Apidae) de la fauna española. Colección Tesis Doctorales, nº 203/84, Ed. Universidad Complutense de Madrid, Madrid, 334 pp.

    Ornosa, C. (1986a) Especies de Bombinae de la Sierra de los Ancares (Hymenoptera, Apidae). Actas de las VIII Jornadas de la Asociación Española de Entomología, Sevilla, 1986, 1042–1050.

    Ornosa, C. (1986b) Revisión de los Bombinae estudiados por D. Modesto Quilis (Hymenoptera, Apidae). Boletín de la Asociación Española de Entomología, 10, 359–368.

    Ornosa, C. (1991) Consideraciones taxonómicas sobre el subgénero Megabombus Dalla Torre, 1880, en la Peninsula Ibérica (Hym., Apidae, Bombinae). Anales de Biología, 17 (Biología Animal 6), 33–41.

    Ornosa, C. (2006) Bombus (Confusibombus) confusus Schenck, 1861. In: Verdú, J.R. y Galante, E. (Eds.), Libro Rojo de los Invertebrados de España. Dirección General para la Biodiversidad, Ministerio de Medio Ambiente, Madrid, pp. 203.

    Ornosa, C. (2011) Bombus (Confusibombus) confusus Schenck, 1861. In: Verdú, J.R., Numa, C. & y Galante, E. (Eds.), Atlas y Libro Rojo de los Invertebrados amenazados de España (Especies Vulnerables). Dirección General de Medio Natural y Política Forestal, Ministerio de Medio Ambiente, Medio Rural y Marino, Madrid, pp. 1174–1178.

    Ornosa, C. & Gayubo, S.F. (1989) Notas sobre los Bombinae de la Sierra de Béjar (Hym., Apidae). Boletín de la Asociación española de Entomología, 13, 173–182.

    Ornosa, C. & Ortiz-Sánchez, F.J. (2004) Hymenoptera: Apoidea I. In: Consejo Superior de Investigaciones Cientificas (Ed.), Fauna Ibérica. Vol. 23. Museo Nacional de Ciencias Naturales, CSIC, Madrid, pp. 1–556.

    Ornosa C. & Ortiz-Sánchez, F.J. (2008a) Bombus (Melanobombus) lapidarius decipiens (Pérez, 1890). In: Barea-Azcón, J.M., Ballesteros, E. & Moreno, D. (Coords.), Libro Rojo de Invertebrados de Andalucía. Consejería de Medio Ambiente (Junta de Andalucía), Tomo III, 1190–1193.

    Ornosa, C. & Ortiz-Sánchez, F.J. (2008b) Bombus pratorum santonae Tkalců, 1985. In: Barea-Azcón, J.M., Ballesteros, E. & Moreno, D. (Coords.), Libro Rojo de Invertebrados de Andalucía. Conserjería de Medio Ambiente (Junta de Andalucía), Tomo III, pp. 1351.

    Ornosa, C. & Ortiz-Sánchez, F.J. (2008c) Psithyrus (Fernaldaepsithyrus) sylvestris Lepeletier, 1833. In: Barea-Azcón, J.M., Ballesteros, E. & Moreno, D. (Coords.), Libro Rojo de Invertebrados de Andalucía. Consejería de Medio Ambiente (Junta de Andalucía), Tomo IV, pp. 1352.

    Ornosa, C. & Torres, F. (2009a) Mendacibombus (Mendacibombus) mendax (Gerstaecker, 1869) En Peligro. In: Verdú, J.R. & Galante, E. (Eds.), Atlas de Invertebrados de España. Especies En Peligro y en Peligro Crítico. Dirección general para la Biodiversidad, Ministerio de Medio Ambiente y Medio Rural y Marino, Madrid, pp. 1–340.

    Ornosa, C. & Torres, F. (2009b) Psithyrus (Fernaldaepsithyrus) flavidus (Eversmann, 1852). En Peligro. In: Verdú, J.R. & Galante, E. (Eds.), Atlas de Invertebrados de España. Especies En Peligro y en Peligro Crítico. Dirección general para la Biodiversidad, Ministerio de Medio Ambiente y Medio Rural y Marino, Madrid, pp. 1–340.

    Ornosa, C. & Castro, L. (2011) Bombus (Megabombus) gerstaeckeri Morawitz, 1881. In: Verdú, J.R., Numa, C. & y Galante, E. (Eds.), Atlas y Libro Rojo de los Invertebrados amenazados de España (Especies Vulnerables). Dirección General de Medio Natural y Política Forestal, Ministerio de Medio Ambiente, Medio Rural y Marino, Madrid, pp. 412–416.

    Ornosa, C. & Ortiz-Sánchez, F.J. (2011) Bombus (Cullumanobombus) cullumanus (Kirby, 1802). In: Verdú, J.R., Numa, C. & y Galante, E. (Eds.), Atlas y Libro Rojo de los Invertebrados amenazados de España (Especies Vulnerables). Dirección General de Medio Natural y Política Forestal, Ministerio de Medio Ambiente, Medio Rural y Marino, Madrid, pp. 408–411

    Ornosa, C. & Torres, F. (2011) Bombus (Thoracobombus) inexspectatus (Tkalcu, 1963). In: Verdú, J.R., Numa, C. & Galante, E. (Eds.), Atlas y Libro Rojo de los Invertebrados amenazados de España (Especies Vulnerables). Dirección General de Medio Natural y Política Forestal, Ministerio de Medio Ambiente, Medio Rural y Marino, Madrid, pp. 417–421.

    Patiny, S., Michez, D. & Rasmont, P. (2009) A survey and review of the status of wild bees in the West Palaearctic region. Apidologie, 40, 313–331.
    https://doi.org/10.1051/apido/2009028

    Pyke, G.H., Thomson, J.D., Inouye, D.W. & Miller, T.J. (2016) Effects of climate change on phenologies and distributions of bumble bees and the plants they visit. Ecosphere, 7 (3), 1–19.
    https://doi.org/10.1002/ecs2.1267

    Ploquin, E.F., Herrera, J.M. & Obeso, J.R. (2013) Bumblebee community homogenization after uphill shifts in montane areas of northern Spain. Oecologia, 173, 1649–1660.
    https://doi.org/10.1007/s00442-013-2731-7

    Ponchau, O., Iserbyt S., Verhaeghe, J.C. & Rasmont, P. (2006) Is the caste-ratio of the oligolectic bumblebee Bombus gerstaeckeri Morawitz (Hymenoptera: Apidae) biased to queens? Annals de la Socièté entomologique de France, 42, 207–214.

    Potts, S.G., Biesmeijer, J.C., Kremen, C., Neumann, P., Schweiger, O. & Kunin, W.E. (2010) Global pollinator declines: trends, impacts and drivers. Trends in Ecology & Evolution, 25, 345–353.
    https://doi.org/10.1016/j.tree.2010.01.007

    Potts, S.G., Imperatriz-Fonseca, V., Ngo, H.T., Aizen, M.A., Biesmeijer, J.C., Breeze, T.D., Dicks, L.V., Garibaldi, L.A., Hill, R., Steele, J. & Vanbergen, A.J. (2016) Safeguarding pollinators and their values to human well-being. Nature, 540, 220–229.
    https://doi.org/10.1038/nature20588

    Quilis, M. (1927) Los Ápidos de España. Genus Bombus Latr. Anales del Instituto Nacional de 2ª Enseñanza de Valencia, Trabajo del Laboratorio de Historia Natural, 16: 1–121.

    Quilis, M. (1932) Los Psithyrus españoles (Hym. Apid.). Eos, 8, 185–222.

    Quintero, C., Morales, C.L. & Aizen, M.A. (2009) Effects of anthropogenic habitat disturbance on local pollinator diversity and species turnover across a precipitation gradient. Biodiversity and Conservation, 19, 257–274.
    https://doi.org/10.1007/s10531-009-9720-5

    Rasmont, P. (1986) Pyrobombus jonellus (Kirby) dans la péninsule ibérique et les Pyrenées (Hymenoptera: Apidae, Bombinae). Entomologische Berichten, 46, 185–189.

    Rasmont, P. (1988) Monographie écologique et zoogéographique des Bourdons de France et de Belgique (Hymenoptera, Apidae, Bombinae). Faculté des Sciences agronomiques de L´Etat, Gembloux, 310 pp.

    Rasmont, P., Pauly, A., Terzo, M., Patiny, S., Michez, D., Iserbyt, S., Barbier, Y. & Haubruge, E. (2006) The survey of wild bees (Hymenoptera, Apoidea) in Belgium and France. Status of the World's Pollinators. Food and Agriculture. FAO, Rome, 18 pp.

    Rasmont, P. & Iserbyt, I. (2010–2013) Atlas of the European Bees: genus Bombus. 3rd Edition. STEP Project, Atlas Hymenoptera, Mons, Gembloux. Available from: http://www.zoologie.umh.ac.be//hymenoptera/page.asp?ID=169 (accessed 7 October 2015)

    Rasmont, P., Franzén, M., Lecocq. T., Harpke, A., Roberts, S.P.M., Biesmeijer, K., Castro, L., Cederberg, B., Dvořák, L., Fitzpatrick, Ú., Gonseth, Y., Haubruge, E., Mahé, G., Manino, A., Michez, D., Neumayer, J., Ødegaard, F., Paukkunen, J., Pawlikowski, T., Potts, S.G., Reemer, M., Settele, J., Straka, J. & Schweiger, O. (2015) Climatic risk and distribution Atlas of European bumblebees. Biorisk, 10, 1–246.
    https://doi.org/10.3897/biorisk.10.4749

    Roura-Pascual, N., Pons, P., Etienne, M. & Lambert, B. (2005) Transformation of a rural landscape in the Eastern Pyrenees between 1953 and 2000. Mountain Research and Development, 25, 252–261.
    https://doi.org/10.1659/0276-4741(2005)025[0252:TOARLI]2.0.CO;2

    Santamaría, S., Castro L., García-Camacho, R., Giménez, L., Méndez, M., Milla, R., Teixido, A.L. & Torices, R. (2011) Abejorros (Bombus spp.: Hymenoptera, Apidae) del Jou de Los Cabrones (Parque Nacional Picos De Europa) y confirmación de la Presencia de Bombus mendax Gerstaecker, 1869 en la Cordillera Cantábrica (España). Boletín de la Sociedad Entomológica Aragonesa, 48, 143–146.

    Suhonen, J., Rannikko, J. & Sorvari, J. (2015) The rarity of host species affects the co-extinction risk in socially parasitic bumblebee Bombus (Psithyrus) species. Annales Zoologici Fennici, 52, 236–242.
    https://doi.org/10.5735/086.052.0402

    Tkalců, B. (1962) Sur la faunistique des Bourdons d´Espagne. Bulletin de la Société Entomologique de Mulhouse, May–June, 44–48.

    Vargas, P., Ornosa, C., Blanco, J.L., Romero, D., Fernández-Mazuecos, M. & Rodríguez-Gironés, M.A. (2013) En búsqueda de áreas de diversidad genética en Sierra Nevada. In: Ramírez, L. & Asensio, B. (Eds.), Proyectos de investigación en parques nacionales. MAGRAMA, Madrid, 223–242.

    Vargas, P., Ornosa, C., Ortiz-Sánchez, F.J. & Arroyo, J. (2010) Is the occluded corolla of Antirrhinum bee-specialized?. Journal of Natural History, 44, 1427-1443.

    https://doi.org/10.1080/00222930903383552

    Verdú, J.R. & Galante, E. (2006) Libro Rojo de Invertebrados de España. Dirección General para la Biodiversidad. Ministerio de Medio Ambiente. Madrid, 414 pp.

    Verdú, J.R. & Galante, E. (2009) Atlas de Invertebrados de España. Especies En Peligro y en Peligro Crítico. Dirección General para la Biodiversidad, Ministerio de Medio Ambiente y Medio Rural y Marino, Madrid, 342 pp.

    Verdú, J.R., Numa, C. & Galante, E. (2011) Atlas y Libro Rojo de los Invertebrados amenazados de España (Especies Vulnerables). I y II. Dirección General de Medio Natural y Política Forestal, Ministerio de Medio Ambiente, Medio Rural y Marino, Madrid, 1316 pp.

    Vigna-Taglianti, A., Audisio, P.A., Brondi., Bologna, M.A., Carpaneto, G. M., De Biasse, A., Farttoni, S., Piatella, E., Sindaco, R., Venchi, A. & Zapparoli., M. (1999) A proposal for a chorotype classification of the Near East fauna, in the framework of the Westen Palearctic region. Biogeographia, 20, 31–59.

    Wallberg, A., Han, F., Wellhagen, G., Dahle, B., Kawata, M., Haddad, N., Simões, Z.L.P., Allsopp, M.H., Kandemir, I., De la Rúa, P., Pirk, C.W.W. & Webster, M.T. (2014) A worldwide survey of genome sequence variation provides insight into the evolutionary history of the honeybee Apis mellifera. Nature Genetics, 46, 1081–1088.
    https://doi.org/10.1038/ng.3077

    Williams, P.H. (1998) An annotated checklist of bumblebees with an analysis of patterns of descriptions (Hymenoptera: Apidae, Bombini). Bulletin of the Natural History of the Museum of London (Entomology), 67, 79–152.

    Williams, P. (2005) Does specialization explain rarity and decline among British bumblebees? A response to Goulson et al. Biological Conservation, 122 (1), 33–43.
    https://doi.org/10.1016/j.biocon.2004.06.019

    Williams, P.H. & Osborne, J.L. (2009) Bumblebee vulnerability and conservation world–wide. Apidologie, 40, 367–387.
    https://doi.org/10.1051/apido/2009025

    Williams, P.H., Cameron, S., Hines, H., Cederberg, B. & Rasmont, P. (2008) A simplified subgeneric classification of the bumblebees (Genus Bombus). Apidologie, 39, 46–74.
    https://doi.org/10.1051/apido:2007052

    Williams, P.H., Brown, M.J.F., Carolan, J.C., An, J., Goulson, D., Aytekin, A.M., Best, L.R., Byvaltsev, A.M., Cederberg, B., Dawson, R., Huang, J., Ito, M., Monfared, A., Raina, R.H., Schmid-Hempel, P., Sheffield, C.S., Šima, P. & Xie, Z. (2012) Unveiling cryptic species of the bumblebee subgenus Bombus s. str. worldwide with COI barcodes (Hymenoptera: Apidae). Systematics and Biodiversity, 10, 21–56.
    https://doi.org/10.1080/14772000.2012.664574

    Williams, P.H., Thorp, R.W., Richardson, L.L. & Colla, S.R. (2014) Bumble Bees of North America: An Identification Guide. Princeton University Press, Princeton, NJ, 208 pp.