Progress 11/01/01 to 10/31/05
Outputs
During the tenure of this grant we have progressed well beyond expectation with the phylogeny of bumble bees. Our original goals were to collect approximately 35% of the world's species and build a phylogeny based on 3 genes. Instead, we have collected greater than 90% of bumble bee species worldwide and have constructed the phylogeny based on 4 genes. The research from this grant led to a renewal proposal funded last year in July 2004. The overall accomplishments of this project include the development of a new gene PEPCK for phylogenetic analysis of bumble bees, an advanced website for bumble bees, including photographs, distribution data, maps, and the like (built by my collaborator, Paul Williams of the Natural History Museum in London). We completed much of the phylogeny, trained a top student in the field of molecular systematics and have several papers in press. The major goals of the research (1. a well supported phylogeny of the bumble bees, and 2. development
of a useful website for bumble bees) have been accomplished. The phylogeny is ready for publication, and will serve as the basis for understanding important aspects of bumble bee biology, including the cladistic pattern of tongue length traits and aspects of bumble bee behavior that should allow greater productivity in the rearing of bumble bees for greenhouse pollination. The molecular phylogeny will also serve as the basis for an upcoming revision of the subgeneric classification system of bumble bees.
Impacts
The impact of this work is such that it should stand as the definitive phylogeny of the group for some time. Several of the subgeneric designations will require reclassification, although many of the traditional subgeneric groups have been shown to be well supported. Thus, the classification will change but mostly on the scale of closely related species groups within subgenera. We are missing fewer than 10% of the species for the entire genus; there is no analysis of the group that comes even close to this level of completion. From the phylogeny we expect to gain insights into the evolution of traits important in pollination, including tongue length. Patterns of the evolution of coloration should become clear with the phylogeny, a starting point for examining the development and genetics of mimicry evolution in bumble bees. We will also be able to learn more about the evolution of sociality among the other social bees within the corbiculate clade, which, besides bumble
bees, includes the honey bees and stingless bees.
Publications
- Hines, H.M., Cameron, S.A. and Williams, P.H. 2006. Molecular phylogeny of the bumble bee subgenus Pyrobombus (Hymenoptera: Apidae: Bombus) with insights into gene utility for lower level analyses. Invertebrate Systematics.
- Williams, P., Ya, T., Jian, Y., Hines, H. and Cameron, S.A. 2006. The bumble bees of Sichuan (Hymenoptera: Apidae, Bombini). Systematics and Biodiversity (In Press).
- Cameron, S.A. and Williams, P.H. 2003. Phylogeny of the New World bumble bee subgenus Fervidobombus: Congruence from nuclear, mitochondrial, and morphological data. Mol. Phylogenet. Evol. 28: 552-563.
- Dornhaus, A. and Cameron, S.A. 2003. A scientific note on food alert in Bombus transversalis. Apidologie 34: 87-88.
- Cameron, S.A. and Mardulyn, P. 2003. The major opsin gene is useful for inferring higher level phylogenetic relationships of the corbiculate bees. Molecular Phylogenetics and Evolution 28: 610-613.
- Cameron, S.A. and Taylor, O.M. 2003. Nest architecture of the Amazonian bumble bee (Hymenoptera: Apidae). Apidologie 34: 321-331.
- Cameron, S.A. 2004. Phylogeny and biology of the Neotropical orchid bees (Euglossini). Annual Review of Entomology 49: 377-404.
- Williams, P., Ya, T., Jian, Y., Hines, H. and Cameron, S.A. 2004. The bumble bees of Sichuan (Hymenoptera: Apidae, Bombini). Systematics and Biodiversity (In Press).
- Salzat. A. and Cameron, S.A. 2004. Phylogeny of the orchid bees (Hymenoptera: Apinae: Euglossini): DNA and morphology yield equivalent patterns. Mol. Phylogenet. Evol. (In Press).
- Rasmussen, C. and Cameron, S.A. 2004. Conopid fly (Diptera: Conopidae) attacking large orchid bees (Hymenoptera: Apidae: Eulaema spp.). J. Kans. Entomol. Soc. 77 (In Press).
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Progress 01/01/04 to 12/31/04
Outputs
Progress over the last year has taken us near completion of the major goals of this project. We have completed the following: 1. Reconstruction of the phylogeny of the bumble bees worldwide (220 of the 240 species), resulting in a highly supported tree of species groups and subgenera. The monophyly of most of the species groups based on previous taxonomic research is upheld by the molecular data collected for this project. The data include 4 genes (we added the fourth gene this year): Approximately 200 species have been sequenced for three genes (opsin, EF-1a, mt16S and arginine kinase). Tests of utilility of these 4 genes indicate they are useful for lower level analyses of infra-subgeneric relationships, perhaps a surprising result given that they have been applied mostly to higher level analyses. 2. We have made progress on designing a web-based database, which is to be installed on a UIUC server and linked to the Bumble bee database of our collaborator, P.
Williams of the Natural History Museum, London. This will include a complete phylogeny and links for each species to photos and important biological information of use to agriculture.
Impacts
The impact of this work is such that it should stand as the definitive phylogeny of the group for some time. It is unlikely that any of the major subgeneric relationships will change, and even within subgeneric relationships are strongly supported. We are missing only ~25 species for the entire genus; there is no analysis of the group that comes even close to this level of completion. From the phylogeny we expect to gain insights into the evolution of traits important in pollination, including tongue length. Patterns of the evolution of coloration should become clear with the phylogeny, a starting point for examining the development and genetics of mimicry evolution in bumble bees. We will also be able to learn more about the evolution of sociality among the other social bees within the corbiculate clade, which, besides bumble bees, includes the honey bees and stingless bees.
Publications
- Cameron, S.A. 2003. DNA from the elongation factor-1 alpha gene corroborates the phylogenetic pattern from other genes revealing common ancestry of bumble bees and stingless bees (Hymenoptera: Apinae). III Seminario Mesoamericano sobre abejas sin Aguijon, Tapachula, Chiapas, Mexico. pp. 132-136.
- Hines, H.M., Cameron, S.A. and Williams, P.H. 2004. Phylogeny of the subgenus Pyrobombus (Hymenoptera: Apidae: Bombus) with insights into gene utility for lower level analyses. Syst. Entomol. (In Review).
- Williams, P., Ya, T., Jian, Y., Hines, H. and Cameron, S.A. 2004. The bumble bees of Sichuan (Hymenoptera: Apidae, Bombini). Systematics and Biodiversity (In Press).
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Progress 01/01/03 to 12/31/03
Outputs
During this year (2003) we have progressed well beyond expectation with the phylogeny of bumble bees. The field expeditions from last year yielded the material necessary for large-scale DNA sequencing of nearly 200 species within the genus, which contains 247 described species. My graduate student RA on the project (H. Hines) has proven exceptional in both the molecular and theoretical aspects of the research. The following list includes specific details of this year's progress. 1. Approximately 200 species have been sequenced for three genes (opsin, EF-1a, 16S) and a fourth is under exploration (candidates include arginine kinase, wingless and CAD, all of which we have preliminary sequences for). We are also testing utility of these genes for their general level of applicability to resolve closely related species. 2. Multiple phylogenetic analyses of these gene sequences have resolved several controversial questions concerning relationships. In particular, results
show conclusively, even at this incomplete stage of taxon sampling for all intended genes, that the large subgenus Pyrobombus is monophyletic, that the large New World subgenus Fervidobombus is polyphyletic, and that Mendacibombus is sister group to the rest of the bumble bees. We have also worked out the sister group relationships to the Pyrobombus and find that the subgenus Bombus sensu strictu is the sister group. This is important given that the latter subgenera contain some of the most important species used in the pollination industry. 3. Collaborator Williams has organized a large multi-author effort to complete a detailed annotated checklist of the bumble bees of Sichuan Province, China, based on our expedition to that region. This work lays the foundation for additional distributional studies and will be a model for the web-based storage of worldwide distributional data. Moreover, we have consolidated strong collaborations with the Chinese. 4. Additional data have been added
to the Bombus website, and this aspect will continue.
Impacts
The phylogeny is nearing completion and the two publications we will submit over the next few months will be the first notification of its full-scale availability. The expected impact should be considerable based on input from peers at the recent ESA meeting in Cincinnatti. Hines presented a spectacular poster based on 180+ species and 2 genes (plus a portion of a third gene). This poster took the First Place award in the student competion for Section A (Systematics), and there was considerable buzz over the large scope of the work. The fully complete phylogeny will need additional time and funding to maximize the rigor of the end-product and thus its usefulness to the end-users (agricultural community and ecologists and evolutionary biologists). Due to the significance of bumble bees within the agricultural community (worldwide), the complete phylogeny of Bombus may serve to spearhead a sequencing effort at the level of the genome. This is in the early stages of
discussion.
Publications
- Rasmussen, C. and Cameron, S.A. 2004. Conopid fly (Diptera: Conopidae) attacking large orchid bees (Hymenoptera: Apidae: Eulaema spp.). J. Kans. Entomol. Soc. 77 (In Press).
- Cameron, S.A. and Williams, P.H. 2003. Phylogeny of the New World bumble bee subgenus Fervidobombus: Congruence from nuclear, mitochondrial, and morphological data. Mol. Phylogenet. Evol. 28: 552-563.
- Dornhaus, A. and Cameron, S.A. 2003. A scientific note on food alert in Bombus transversalis. Apidologie 34: 87-88.
- Cameron, S.A. and Mardulyn, P. 2003. The major opsin gene is useful for inferring higher level phylogenetic relationships of the corbiculate bees. Molecular Phylogenetics and Evolution 28: 610-613.
- Cameron, S.A. and Taylor, O.M. 2003. Nest architecture of the Amazonian bumble bee (Hymenoptera: Apidae). Apidologie 34: 321-331.
- Cameron, S.A. 2004. Phylogeny and biology of the Neotropical orchid bees (Euglossini). Annual Review of Entomology 49: 377-404.
- Williams, P., Ya, T., Jian, Y., Hines, H. and Cameron, S.A. 2004. The bumble bees of Sichuan (Hymenoptera: Apidae, Bombini). Systematics and Biodiversity (In Press).
- Salzat, A. and Cameron, S.A. 2004. Phylogeny of the orchid bees (Hymenoptera: Apinae: Euglossini): DNA and morphology yield equivalent patterns. Mol. Phylogenet. Evol. (In Press).
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Progress 01/01/02 to 12/31/02
Outputs
During the last year the following has been accomplished in direct association with this grant: 1. A graduate was brought in to the lab in March. 2. Two collecting expeditions were made: one to Turkey by the graduate student, and one to southwest China (Sichuan Province) by Cameron and Williams (collaborator). 3. The Turkey expedition was tremendously successful, with the student bringing back 50 species of Bombus for sequencing. The China trip, likewise was enormously successful as we brought back 37 species. 4. Since the summer expeditions to bring back needed subgenera and species we have sequenced 100+ species for the molecular phylogeny. 5. The collbaorator has identified all China material, and is beginning his assessment of morphological characters for analysis. 6. We have explored the use of the following genes for the molecular phylogeny: 16S (100+ species to date), EF-1a (10-15 species to date). 7. We intend to move on to genes that show sufficient diversity
for the subgeneric relationships to be fully resolved. At this point, we have good indication that many of the subgeneric designations are monophyletic and there appears to be good concordance with the collaborator's morphology-based phylogeny. This is excellent progress for a genus that contains 240 species worldwide. In one semester we have sequenced more than 40% of all known species. The successful sequencing has taken place in spite of the fact the PI moved her lab last year, and the graduate student began her first semester of graduate school only this fall (Sept. 2002).
Impacts
When the project is complete we will have a robust phylogeny of the worldwide taxa of Bombus based on 4 genes and morphology. This will allow us to begin to answer numerous questions that depend on knowledge of relationships of this important native pollinator. When a phylogeny is in place, and the information is placed into a database for ease of information retrieval, the data will be of great use to ecologists, pollination specialists and those planning to rear bumble bees for industrial pollination services.
Publications
- Cameron, S.A. and Whitfield, J.B. 2002. Review of Page, R.D.M and E.C. Holmes, Molecular Evolution: A Phylogenetic Approach, Blackwell Science, Oxford, 1998. Systematic Biology 51: 536-538
- Grissell, E.E. and Cameron, S.A. 2002. A new Leucospis (Hymenoptera: Leucospidae), the first known gregarious species. Journal of Hymenoptera Research 11: 271-278.
- Cameron, S.A. and Taylor, O.M. 2003. Nest architecture of the Amazonian bumble bee (Hymenoptera: Apidae) (Apidologie) (In Press).
- Cameron, S.A. and Mardulyn, P. 2003. The major opsin is still a promising nuclear gene for higher level phylogenetics. (Molecular Phylogenetics and Evolution).
- Cameron, S.A. and Ramirez, S. 2003. Army ant attacks by Eciton hamatum and E. rapax on nests of the Amazonian bumble bee, Bombus transversalis (Hymenoptera: Apidae). (J. Kans. Entomol. Soc.).
- Dornhaus, A. and Cameron, S.A. 2003. Food alert in Bombus transversalis: A scientific note (Apidologie).
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