Phylogenetic systematics and a revised generic classification of anthidiine bees (Hymenoptera: Megachilidae).

The bee tribe Anthidiini (Hymenoptera: Megachilidae) is a large, cosmopolitan group of solitary bees that exhibit intriguing nesting behavior. We present the first molecular-based phylogenetic analysis of relationships within Anthidiini using model-based methods and a large, multi-locus dataset (five nuclear genes, 5081 base pairs), as well as a combined analysis using our molecular dataset in conjunction with a previously published morphological matrix. We discuss the evolution of nesting behavior in Anthidiini and the relationship between nesting material and female mandibular morphology. Following an examination of the morphological characters historically used to recognize anthidiine genera, we recommend the use of a molecular-based phylogenetic backbone to define taxonomic groups prior to the assignment of diagnostic morphological characters for these groups. Finally, our results reveal the paraphyly of numerous genera and have significant consequences for anthidiine classification. In order to promote a classification system based on stable, monophyletic clades, we hereby make the following changes to Michener's (2007) classification: The subgenera Afranthidium (Zosteranthidium) Michener and Griswold, 1994, Afranthidium (Branthidium) Pasteels, 1969 and Afranthidium (Immanthidium) Pasteels, 1969 are moved into the genus Pseudoanthidium, thus forming the new combinations Pseudoanthidium (Zosteranthidium), Pseudoanthidium (Branthidium), and Pseudoanthidium (Immanthidium). The genus Neanthidium Pasteels, 1969 is also moved into the genus Pseudoanthidium, thus forming the new combination Pseudoanthidium (Neanthidium). Based on morphological characters shared with our new definition of the genus Pseudoanthidium, the subgenus Afranthidium (Mesanthidiellum) Pasteels, 1969 and the genus Gnathanthidium Pasteels, 1969 are also moved into the genus Pseudoanthidium, thus forming the new combinations Pseudoanthidium (Mesanthidiellum) and Pseudoanthidium (Gnathanthidium).

Why do leafcutter bees cut leaves? New insights into the early evolution of bees.

Stark contrasts in clade species diversity are reported across the tree of life and are especially conspicuous when observed in closely related lineages. The explanation for such disparity has often been attributed to the evolution of key innovations that facilitate colonization of new ecological niches. The factors underlying diversification in bees remain poorly explored. Bees are thought to have originated from apoid wasps during the Mid-Cretaceous, a period that coincides with the appearance of angiosperm eudicot pollen grains in the fossil record. The reliance of bees on angiosperm pollen and their fundamental role as angiosperm pollinators have contributed to the idea that both groups may have undergone simultaneous radiations. We demonstrate that one key innovation--the inclusion of foreign material in nest construction--underlies both a massive range expansion and a significant increase in the rate of diversification within the second largest bee family, Megachilidae. Basal clades within the family are restricted to deserts and exhibit plesiomorphic features rarely observed among modern bees, but prevalent among apoid wasps. Our results suggest that early bees inherited a suite of behavioural traits that acted as powerful evolutionary constraints. While the transition to pollen as a larval food source opened an enormous ecological niche for the early bees, the exploitation of this niche and the subsequent diversification of bees only became possible after bees had evolved adaptations to overcome these constraints.

Rooting phylogenies using gene duplications: an empirical example from the bees (Apoidea).

The placement of the root node in a phylogeny is fundamental to characterizing evolutionary relationships. The root node of bee phylogeny remains unclear despite considerable previous attention. In order to test alternative hypotheses for the location of the root node in bees, we used the F1 and F2 paralogs of elongation factor 1-alpha (EF-1α) to compare the tree topologies that result when using outgroup versus paralogous rooting. Fifty-two taxa representing each of the seven bee families were sequenced for both copies of EF-1α. Two datasets were analyzed. In the first (the "concatenated" dataset), the F1 and F2 copies for each species were concatenated and the tree was rooted using appropriate outgroups (sphecid and crabronid wasps). In the second dataset (the "duplicated" dataset), the F1 and F2 copies were aligned to each another and each copy for all taxa were treated as separate terminals. In this dataset, the root was placed between the F1 and F2 copies (e.g., paralog rooting). Bayesian analyses demonstrate that the outgroup rooting approach outperforms paralog rooting, recovering deeper clades and showing stronger support for groups well established by both morphological and other molecular data. Sequence characteristics of the two copies were compared at the amino acid level, but little evidence was found to suggest that one copy is more functionally conserved. Although neither approach yields an unambiguous root to the tree, both approaches strongly indicate that the root of bee phylogeny does not fall near Colletidae, as has been previously proposed. We discuss paralog rooting as a general strategy and why this approach performs relatively poorly with our particular dataset.

Under the radar: detection avoidance in brood parasitic bees.

Brood parasitism is a specialized form of parasitism in which the offspring of a parasite develops on the food provisions gathered by a host species for its own young. Obligate brood parasitic lineages have lost the ability to acquire provisions for their young and thus rely entirely on the location of an appropriate host to serve as a food-provider. Solitary bees provide some of the most fascinating examples of brood parasitism in animals. Most solitary bees build and provision their own nests. Some, however, usurp the nests of other species of bees. These brood parasites, or 'cuckoo' bees, deposit their eggs on the pollen provisions collected by a host bee for her own offspring. The provisions stored by the host bee are not sufficient to sustain the development of both the host's larva and that of the brood parasite and the parasite must kill the offspring of its host in order to obtain enough nourishment to complete its development. As a consequence, there is fierce competition between the host bee seeking to protect her nest from attack and the brood parasite seeking to avoid detection by the host in order to successfully deposit her eggs in an appropriate nest. In this paper, I review the behaviours that allow brood parasitic bees to escape detection by their hosts. Identifying these behaviours, and placing them within the general context of strategies employed by brood parasitic bees to parasitize the nests of their hosts, is key to understanding how brood parasitic lineages may have evolved from nest-building ancestors, decrypting the selective pressures that drive evolutionary transitions from one strategy to another and, more broadly, revealing how similar selective pressures in widely divergent lineages of animals have given rise to remarkably convergent behaviours. This article is part of the theme issue 'The coevolutionary biology of brood parasitism: from mechanism to pattern'.

Origins, evolution, and diversification of cleptoparasitic lineages in long-tongued bees.

The evolution of parasitic behavior may catalyze the exploitation of new ecological niches yet also binds the fate of a parasite to that of its host. It is thus not clear whether evolutionary transitions from free-living organism to parasite lead to increased or decreased rates of diversification. We explore the evolution of brood parasitism in long-tongued bees and find decreased rates of diversification in eight of 10 brood parasitic clades. We propose a pathway for the evolution of brood parasitic strategy and find that a strategy in which a closed host nest cell is parasitized and the host offspring is killed by the adult parasite represents an obligate first step in the appearance of a brood parasitic lineage; this ultimately evolves into a strategy in which an open host cell is parasitized and the host offspring is killed by a specialized larval instar. The transition to parasitizing open nest cells expanded the range of potential hosts for brood parasitic bees and played a fundamental role in the patterns of diversification seen in brood parasitic clades. We address the prevalence of brood parasitic lineages in certain families of bees and examine the evolution of brood parasitism in other groups of organisms.