Introduction
Bees are a diverse, fascinating, and important group of insects with an intimate ecological interrelationship with the angiosperm (flowering) plants. The enormous radiation of the flowering plants may be due in part to the nearly simultaneous diversification of the bees. Today bees are one of the most economically and ecologically important insect groups. There are over 16,000 described species of bees and we are just beginning to understand the basal phylogeny of the bees, their historical biogeography, and the antiquity of bees.
This web site is meant to be a primer for anyone interested in understanding more about higher-level bee phylogeny (phylogeny at the level of genus, tribe, subfamily and family). We review previously published studies on the phylogeny of bees, the antiquity of bees, and the historical biogeography of bees. We focus our review primarily on the previously published morphological studies which provide the background and framework for all future studies of bee phylogeny. Finally, we present information on the molecular systematics of bees. We provide background material and literature reviews on eight promising nuclear gene loci which have been used previously in bees. We provide lists of primers and maps of genes (as downloadable pdf files), and we provide recommendations on what genes appear to be useful for reconstructing higher-level relationships. We provide data sets (in the form of Paup NEXUS files) of previous studies of bee phylogeny, as well as Excel spreadsheets listing GenBank numbers in cases where there is a substantial amount of sequence data available.
The genes EF-1alpha (F2 copy), wingless (wnt-1 copy), and LW rhodopsin appear to be the most promising candidates to date and substantial data sets already exist for these genes in various families of bees. We believe that labs interested in reconstructing bee phylogeny (at any level) should collect data from the same set of genes so that these data sets can be combined in the future.
Authors
Bryan N. Danforth (photo) wrote much of the text, developed many of the primers, and prepared some lists of primers and gene maps. Jennifer Fang (photo) developed the web site design, developed many of the primers, and prepared many of the gene summaries, primer lists and gene maps. Sedonia Sipes (photo) wrote the section on dating the antiquity of the bees, developed many primers, collected much of the EF-1alpha and wingless data discussed below. Sean G. Brady
(photo) contributed much information on dating the antiquity of bees using recently developed Bayesian methods, and collected data on arginine kinase, EF-1alpha, and other genes for a broad sample of bees. He, Sedonia Sipes, and Bryan Danforth are collaborating on a higher-level phylogeny of the bees based on single-copy nuclear gene sequences (funded by NSF; DEB-0211701 and DEB-0211837). Eduardo Almeida (photo) wrote the section on colletid bees and provided helpful comments on the text and layout of the website.
Acknowledgements
First, we thank Charles Michener (and his students and associates) for developing a comprehensive understanding of bee taxonomy on a world-wide basis. Dr. Michener’s book "The Bees of the World" (Michener 2000) provides a comprehensive review of bee taxonomy to the level of subgenus and/or genus and provides the background for current research on higher-level bee phylogeny. Second, we thank the authors and contributors to the honey bee genome project which allowed us to develop many new genes for higher-level bee phylogeny. The availability of the honey bee genome now means that one can find the Apis homolog of virtually any gene known in insects. This greatly facilitates primer design and development. Third, we thank many of the authors of the studies we cite below for providing us with information (in some cases unpublished) on primers and data sets. Fourth, we thank the bee collectors who have generously shared their alcohol preseved material with us for examining molecular evolution in bees. Finally, we thank the National Science Foundation for generous financial support for this project.
How to Cite this Web Site
Please cite this web site in publications if you have been able to use the material below in your research:
Danforth, B.N., J. Fang, S. Sipes, S.G. Brady & E. Almeida (2004). Phylogeny and molecular systematics of bees (Hymenoptera: Apoidea). Cornell University, Ithaca, NY http://www.entomology.cornell.edu/BeePhylogeny/