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Biological Journal of the Linnean Society, 2007, 91, 161C188.

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2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91, 161C188

161 Blackwell Publishing LtdOxford, UKBIJBiological Journal of the Linnean Society0024-4066?

2007 The Linnean Society of London?

2007 91?

161188 Original Articles BUMBLE BEE PHYLOGENY S. A. CAMERON ET AL . *Corresponding author. E-mail: [email protected] A comprehensive phylogeny of the bumble bees (Bombus) S. A. CAMERON1 *, H. M. HINES1 and P. H. WILLIAMS2

1 Department of Entomology,

320 Morrill Hall,

505 S. Goodwin Avenue, University of Illinois, Urbana, IL 61801, USA

2 Department of Entomology, Natural History Museum, South Kensington, London SW7 5BD, UK Received

24 March 2006;

accepted for publication

10 August

2006 Bumble bees (Bombus Latreille) occupy a wide diversity of habitats, from alpine meadows to lowland tropical forest, yet they appear to be similar in morphology throughout their range, suggesting that behavioural adaptations play a more important role in colonizing diverse habitats. Notwithstanding their structural homogeneity, bumble bees exhibit striking inter- and intraspeci?c variation in colour pattern, purportedly the outcome of mimetic evolution. A robust phylogeny of Bombus would provide the framework for elucidating the history of their wide biogeographical distribution and the evolution of behavioural and morphological adaptations, including colour pattern. However, morphological studies of bumble bees have discovered too few phylogenetically informative characters to reconstruct a robust phylogeny. Using DNA sequence data, we report the ?rst nearly complete species phylogeny of bumble bees, including most of the

250 known species from the

38 currently recognized subgenera. Bayesian analysis of nuclear (opsin, EF-1α, arginine kinase, PEPCK) and mitochondrial (16S) sequences results in a highly resolved and strongly supported phylogeny from base to tips, with clear-cut support for monophyly of most of the conventional morphology- based subgenera. Most subgenera fall into two distinct clades (short-faced and long-faced) associated broadly with differences in head morphology. Within the short-faced clade is a diverse New World clade, which includes nearly one- quarter of the currently recognized subgenera, many of which are restricted to higher elevations of Central and South America. The comprehensive phylogeny provides a ?rm foundation for reclassi?cation and for evaluating character evolution in the bumble bees. ?

2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91, 161C188. ADDITIONAL KEYWORDS: classi?cation C corbiculate bees C nuclear genes. INTRODUCTION Bumble bees (Bombus Latreille) are among the more familiar creatures inhabiting meadows, gardens and grasslands of the temperate world (Darwin, 1859: notes 1854C1861, translated by Freeman, 1968;

Sladen, 1912). Robust and vividly coloured, they emerge from hibernation in early Spring in far northerly latitudes, feeding from the earliest willow blossoms before the snows have fully melted. Ther- moregulatory mechanisms maintain their body tem- perature high above ambient in cold to freezing weather (Heinrich, 2004), allowing them full activity under conditions too extreme for other bees. Although they are most abundant in alpine and high-elevation grassland habitats of the northern temperate zone, they range widely from Greenland to the Amazon Basin, from sea level to altitudes of