[en] Although bees are one of the major lineages of pollinators and are today quite diverse, few well-preserved fossils are available from which to establish the tempo of their diversification/extinction since the Early Cretaceous. Here we present a reassessment of the taxonomic affinities of Melitta willardi Cockerell 1909, preserved as a compression fossil from the Florissant shales of Colorado, USA. Based on geometric morphometric
wing shape analyses M. willardi cannot be confidently assigned to the genus Melitta Kirby (Anthophila, Melittidae). Instead, the species exhibits phenotypic affinity with the subfamily Andreninae (Anthophila, Andrenidae), but does not appear to belong to any of the known genera therein. Accordingly, we describe a new genus, Andrenopteryx gen. n., based on wing shape as well as additional morphological features and to accommodate M. willardi. The new combination Andrenopteryx willardi (Cockerell) is established.
Adams DC, Rohlf FJ, Slice DE (2004) Geometric morphometrics: ten years of progress following the 'revolution'. Italian Journal of Zoology 71: 5-16. doi: 10.1080/11250000409356545
Adams DC, Otárola-Castillo E (2013) Geomorph: An R package for the collection and analysis of geometric morphometric shape data. Methods in Ecology and Evolution 4(4): 393-399. doi: 10.1111/2041-210X.12035
Bookstein FL (1991) Morphometric tools for landmark data: Geometry and biology. Cambridge University Press, Cambridge, 435.
Cardinal S, Danforth BN (2013) Bees diversified in the age of eudicots. Proceedings of the Royal Society B-Biological Sciences 280(1755): 1-9.
Cockerell TDA (1909) New North American bees. The Canadian Entomologist 41: 393-395. doi: 10.4039/Ent41393-11
Cockerell TDA (1927) Zoology of Colorado. University Colorado, Denver, 262 pp.
Danforth BN, Cardinal S, Praz C, Almeida EAB, Michez D (2013) The impact of molecular data on our understanding of bee phylogeny and evolution. Annual Review of Entomology 58: 57-78. doi: 10.1146/annurev-ento-120811-153633
De Meulemeester T, Michez D, Aytekin AM, Danforth BN (2012) Taxonomic affinity of halictid bee fossils (Hymenoptera: Anthophila) based on geometric morphometrics analyses of wing shape. Journal of Systematic Palaeontology 10(4): 755-764. doi: 10.1080/14772019.2011.628701
Dellicour S, Lecocq T, Kuhlmann M, Mardulyn P, Michez D (2014) Molecular phylogeny, biogeography, and host plant shifts in the bee genus Melitta (Hymenoptera: Anthophila). Molecular Phylogenetics and Evolution 70: 412-419. doi: 10.1016/j.ympev.2013.08.013
Engel MS (2001a) A monograph of the Baltic Amber bees and evolution of the Apoidea (Hymenoptera). Bulletin of the American Museum of Natural History 259: 1-192. doi: 10.1206/0003-0090(2001)259<0001:AMOTBA>2.0.CO;2
Engel MS (2001b) The first large carpenter bee from the Tertiary of North America, with a consideration of the geological history of Xylocopinae (Hymenoptera: Apidae). Transactions of the American Entomological Society 127(2): 245-254.
Engel MS (2002) Halictine bees from the Eocene-Oligocene boundary of Florissant, Colorado (Hymenoptera: Halictidae). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 225(2): 251-273.
Engel MS (2008) A new species of Ctenoplectrella in baltic amber (Hymenoptera: Megachilidae). Acta Zoologica Academiae Scientiarum Hungaricae 54(4): 319-324.
Engel MS (2011) Systematic melittology: where to from here? Systematic Entomology 36: 2-15. doi: 10.1111/j.1365-3113.2010.00544.x
Engel MS, Michener CD (2013) A minute stingless bee in Eocene Fushan amber from northeastern China (Hymenoptera: Apidae). Journal of Melittology 14: 1-10.
Engel MS, Ortega-Blanco J, Nascimbene PC, Singh H (2013) The bees of Early Eocene Cambay amber (Hymenoptera: Apidae). Journal of Melittology 25: 1-12.
Engel MS, Breitkreuz LCV (2013) A male of the bee genus Agapostemon in Dominican amber (Hymenoptera: Halictidae). Journal of Melittology 16: 1-9.
Gonzalez VH, Engel MS (2011) A new species of the bee genus Ctenoplectrella in middle Eocene Baltic amber (Hymenoptera, Megachilidae). Zookeys 111: 41-49. doi: 10.3897/ zookeys.111.1593
Gumiel M, Catala S, Noireau F, Rojas de Arias A, Garcia A, Dujardin JP (2003) Wing geometry in Triatoma infestans (Klug) and T. melanosoma Martinez, Olmedo & Carcavallo (Hemiptera: Reduviidae). Systematic Entomology 28(2): 173-179. doi: 10.1046/j.1365-3113.2003.00206.x
Huberty CJ, Olejnik S (2006) Applied MANOVA and Discriminant Analysis. Second Edition. New Jersey, 488 pp. doi: 10.1002/047178947X
Kandemir I, Ö-zkan A, Fuchs S (2011) Reevaluation of honeybee (Apis mellifera) microtaxonomy: A geometric morphometric approach. Apidologie 42(5): 618-627. doi: 10.1007/ s13592-011-0063-3
Michener CD (1979) Biogeography of the bees. Annals of the Missouri Botanical Garden 66: 277-342.
Michener CD, Poinar G (1996) The known bee fauna of the Dominican amber. Journal of the Kansas Entomological Society 69(4): 353-361.
Michener CD (2000) The bees of the world. The Johns Hopkins University Press, Baltimore, xiv+[i]+913 pp., + 16 pls.
Michener CD (2007) The bees of the world, second edition. The Johns Hopkins University Press, Baltimore, xvi+[i]+953 pp., +20 pls. doi: 10.1080/00379271.2007.10697535
Michez D, Eardley C (2007) Monographic revision of the bee genus Melitta Kirby 1802 (Hymenoptera: Apoidea: Melittidae). Annales de la Société entomologique de France 43 (4): 379-440. doi: 10.2307/2398833
Michez D, De Meulemeester T, Nel A, Rasmont P, Patiny S (2009) New fossil evidence of the early diversification of bees: Paleohabropoda oudardi from the French Paleocene (Hymenoptera, Apidae, Anthophorini). Zoologica Scripta 38(2): 171-181. doi: 10.1111/j.1463-6409.2008.00362.x
Michez D, Vanderplanck M, Engel MS (2012) Fossil bees and their plant associates. In: Patiny S (Ed) Evolution of plant-pollinator relationships. Cambridge University Press, Cambridge, 103-164.
Pretorius E (2005) Using geometric morphometrics to investigate wing dimorphism in males and females of Hymenoptera-a case study based on the genus Tachysphex Kohl (Hymenoptera: Sphecidae: Larrinae). Australian Journal of Entomology 44: 113-121. doi: 10.1111/j.1440-6055.2005.00464.x
R Development Core Team (2013) A language and environment for statistical computing, version 3.0.2, ISBN 3-900051-07-0, R Foundation for Statistical Computing. Vienna, Austria.
Roberts DW (2008) Statistical analysis of multidimensional fuzzy set ordinations. Ecology 89 (5): 1246-1260. doi: 10.1890/07-0136.1
Rohlf FJ, Marcus LF (1993) A revolution in Morphometrics. Trends in Ecology & Evolution 8(4): 129-132. doi: 10.1016/0169-5347(93)90024-J
Rohlf FJ (1999) Shape Statistics: Procrustes Superimpositions and Tangent Spaces. Journal of Classification 16: 197-223. doi: 10.1007/s003579900054
Rohlf FJ (2013a) tpsUTIL Version 1.56. Department of Ecology and Evolution, State University of New York at Stony Brook, New-York.
Rohlf FJ (2013b) tpsDIG Version 2.17. Department of Ecology and Evolution, State University of New York at Stony Brook, New-York.
Rohlf FJ (2013c) tpsSMALL Version 1.25. Department of Ecology and Evolution, State University of New York at Stony Brook, New-York.
Wappler T, Engel MS (2003) The middle Eocene bee faunas of Eckfeld and Messel, Germany (Hymenoptera: Apoidea). Journal of Paleontology 77(5): 908-921. doi: 10.1666/0022-3360(2003)077<0908:TMEBFO>2.0.CO;2
Wappler T, De Meulemeester T, Aytekin AM, Michez D, Engel MS (2012) Geometric morphometric analysis of a new Miocene bumble bee from the Randeck Maar of southwestern Germany (Hymenoptera: Apidae). Systematic Entomology 37(4): 784-792. doi: 10.1111/j.1365-3113.2012.00642.x
Wedmann S, Wappler T, Engel MS (2009) Direct and indirect fossil records of megachilid bees from the Paleogene of Central Europe (Hymenoptera: Megachilidae). Naturwissenschaften 96(6): 703-712. doi: 10.1007/s00114-009-0525-x
Zeuner FE, Manning FJ (1976) A monograph on fossil bees (Hymenoptera: Apoidea). Bulletin of the British Museum (Natural History), Geology 27(3): 149-268.