Most of the 470-million-year history of plants on land belongs to bryophytes, pteridophytes and gymnosperms, which eventually yielded to the ecological dominance by angiosperms 90 Myr ago. Our knowledge of angiosperm phylogeny, particularly the branching order of the earliest lineages, has recently been increased by the concurrence of multigene sequence analyses. However, reconstructing relationships for all the main lineages of vascular plants that diverged since the Devonian period has remained a challenge. Here we report phylogenetic analyses of combined data--from morphology and from four genes--for 35 representatives from all the main lineages of land plants. We show that there are three monophyletic groups of extant vascular plants: (1) lycophytes, (2) seed plants and (3) a clade including equisetophytes (horsetails), psilotophytes (whisk ferns) and all eusporangiate and leptosporangiate ferns. Our maximum-likelihood analysis shows unambiguously that horsetails and ferns together are the closest relatives to seed plants. This refutes the prevailing view that horsetails and ferns are transitional evolutionary grades between bryophytes and seed plants, and has important implications for our understanding of the development and evolution of plants.

Bees, the largest (>16,000 species) and most important radiation of pollinating insects, originated in early to mid-Cretaceous, roughly in synchrony with the angiosperms (flowering plants). Understanding the diversification of the bees and the coevolutionary history of bees and angiosperms requires a well supported phylogeny of bees (as well as angiosperms). We reconstructed a robust phylogeny of bees at the family and subfamily levels using a data set of five genes (4,299 nucleotide sites) plus morphology (109 characters). The molecular data set included protein coding (elongation factor-1␣, RNA polymerase II, and LW rhodopsin), as well as ribosomal (28S and 18S) nuclear gene data. Analyses of both the DNA data set and the DNA؉morphology data set by parsimony and Bayesian methods yielded a single well supported family-level tree topology that places Melittidae as a paraphyletic group at the base of the phylogeny of bees. This topology (''Melittidae-LT basal'') is significantly better than a previously proposed alternative topology (''Colletidae basal'') based both on likelihood and Bayesian methods. Our results have important implications for understanding the early diversification, historical biogeography, host-plant evolution, and fossil record of bees. The earliest branches of bee phylogeny include lineages that are predominantly host-plant specialists, suggesting that host-plant specificity is an ancestral trait in bees. Our results suggest an African origin for bees, because the earliest branches of the tree include predominantly African lineages. These results also help explain the predominance of Melittidae, Apidae, and Megachilidae among the earliest fossil bees.bee phylogeny ͉ bee evolution ͉ molecular evolution ͉ molecular systematics ͉ coevolution A ngiosperms (flowering plants), with an estimated 250,000-260,000 species (1), represent the largest and most diverse lineage of vascular plants on earth. To Darwin, the rapid emergence and early diversification of the angiosperms was an ''abominable mystery' ' (ref. 2 and refs. therein). Among the most important traits attributable to the explosive radiation of the angiosperms is animal-mediated pollination (3-7). Insects are by far the most important animal pollinators (Ϸ70% of angiosperm species are insect pollinated; ref. 8) and among insects, bees are the most specialized and important pollinator group. All of the Ͼ16,000 species of bees living today (9) rely virtually exclusively on angiosperm products, including pollen and nectar for adult and larval nutrition (10), floral oils for larval nutrition (11, 12), floral waxes and perfumes that serve as sexual attractants (13), and resins for nest construction (14). Bees are morphologically adapted to collecting, manipulating, carrying, and storing pollen and other plant products (15, 16), and many bee species are specialists on one or a few closely related host plants (10).One step toward resolving Darwin's ''abominable mystery'' is to develop a better understanding of the role that bees played in the ev...

We analyzed the higher level phylogeny of the bee family Halictidae based on the coding regions of three single-copy nuclear genes (long-wavelength [LW] opsin, wingless, and elongation factor 1-alpha [EF-1 alpha]). Our combined data set consisted of 2,234 aligned nucleotide sites (702 base pairs [bp] for LW opsin, 405 bp for wingless, and 1,127 bp for EF-1 alpha) and 779 parsimony-informative sites. We included 58 species of halictid bees from 33 genera, representing all subfamilies and tribes, and rooted the trees using seven outgroups from other bee families: Colletidae, Andrenidae, Melittidae, and Apidae. We analyzed the separate and combined data sets by a variety of methods, including equal weights parsimony, maximum likelihood, and Bayesian methods. Analysis of the combined data set produced a strong phylogenetic signal with high bootstrap and Bremer support and high posterior probability well into the base of the tree. The phylogeny recovered the monophyly of the Halictidae and of all four subfamilies and both tribes, recovered relationships among the subfamilies and tribes congruent with morphology, and provided robust support for the relationships among the numerous genera in the tribe Halictini, sensu Michener (2000). Using our combined nucleotide data set, several recently described halictid fossils from the Oligocene and Eocene, and recently developed Bayesian methods, we estimated the antiquity of major clades within the family. Our results indicate that each of the four subfamilies arose well before the Cretaceous-Tertiary boundary and suggest that the early radiation of halictid bees involved substantial African-South American interchange roughly coincident with the separation of these two continents in the late Cretaceous. This combination of single-copy nuclear genes is capable of recovering Cretaceous-age divergences in bees with high levels of support. We propose that LW opsin, wingless, and EF-1 alpha(F2 copy) may be useful in resolving relationships among bee families and other Cretaceous-age insect lineages.

Eusocial organisms are characterized by cooperative brood care, generation overlap and reproductive division of labour. Traits associated with eusociality are most developed in ants, termites, paper wasps and corbiculate bees; the fossil record indicates that each of these advanced eusocial taxa evolved in the Late Cretaceous or earlier (greater than 65 Myr ago). Halictid bees also include a large and diverse number of eusocial members, but, in contrast to advanced eusocial taxa, they are characterized by substantial intraand inter-specific variation in social behaviour, which may be indicative of more recent eusocial evolution. To test this hypothesis, we used over 2400 bp of DNA sequence data gathered from three protein-coding nuclear genes (opsin, wingless and EF-1a) to infer the phylogeny of eusocial halictid lineages and their relatives. Results from relaxed molecular clock dating techniques that utilize a combination of molecular and fossil data indicate that the three independent origins of eusociality in halictid bees occurred within a narrow time frame between approximately 20 and 22 Myr ago. This relatively recent evolution helps to explain the pronounced levels of social variation observed within these bees. The three origins of eusociality appear to be temporally correlated with a period of global warming, suggesting that climate may have had an important role in the evolution and maintenance of eusociality in these bees.

We examined the levels of pollen-host specificity in North American Diadasia (Hymenoptera: Apoidea), a clade of specialist bees. We analysed the scopal pollen loads of 409 individuals representing 25 of the 30 species of Diadasia that occur in North America. Each Diadasia species showed a preference for one of five plant families. However, the 25 species varied in their level of host specificity: the average percentage by volume of the preferred host in pollen loads ranged from > 99% to < 75%. In 17 of the 25 species, all or most individuals examined contained pure loads of one host taxon, while in eight species individuals were less specialized and carried mixtures of several unrelated host taxa. Mapping these host preferences onto a phylogenetic tree indicated that Malvaceae is the most likely ancestral host for the genus, and use of other hosts can be explained by a single switch to each of the other four host-plant families. Thus, most speciation events were not associated with a host switch; this pattern does not support host switching as a niche partitioning strategy to avoid competition. Diadasia species are more likely to use host-plant families that are used by other Diadasia and Emphorine bees; however, there was no evidence of residual adaptation to ancestral hosts. Diet breadth appears to be a labile trait: transitions from narrower to broader host use, as well as vice versa, were observed. The observed patterns of host-use evolution may be driven, in part, by host morphology and/or chemistry.