This site concerns Amelanchier, a member of the Rosaceae (rose) family. The purpose of this site is to present information about the systematics and evolution of these attractive small trees and shrubs, which are commonly called shadbushes, serviceberries, juneberries, sugarplums, and numerous other common names.

Amelanchier grows primarily in early successional habitats of the North Temperate Zone. The genus is most diverse taxomically in North America, especially in the northern United States and southern Canada, and is native to every state of the United States except Hawaii. These plants are valued horticulturally, and their fruits are important to wildlife. The systematics (taxonomy) of shadbushes has long perplexed botanists, horticulturalists, and others, as suggested by the range in number of species recognized in the genus from 6 to 33 in two recent publications (Landry 1975, Phipps et al. 1991; see references on Systematics page). A major source of complexity comes from the occurrence of apomixis (asexual seed production), polyploidy, and hybridization (see Evolution page).

Acknowledgements (1ix13)

We thank the following collaborators, contributors of plant material, students who have contributed to this work, and contacts for localities: Jack Alexander, Larry Alice, Loran Anderson, Tina Ayres, Bruce Baldwin, Mary Barkworth, Scott Bergquist, Luc Brouillet, Larry Brasher, Jan Busco, Pam Camp, Art Campbell, Phil Cantino, David Carmichael, Paul Catling, Ken Chambers, Anita Cholewa, Knub Ib Christensen, Bryan Connolly, Nora Cross, Hubert Daniel, Shawn (Edward) DeKeyser, Jeff Despres, Michelle Dewey, Alvin Diamond, Alison Dibble, Tim Dickinson, Beth Dickson, Michael Donoghue, Frank Drummond, Jim Duncan, Barbara Ertter, Dwayne Estes, James Estes, Rick Gardner, David Giblin, Arthur Gilman, Steve Ginzberg, Gina Gorzejeska, Craig Greene, Christine Guerette, Arthur Haines, Bill Halteman, Curtis Hansen, Ron Hartman, Stuart Hay, Sara Hendershot, Jean Higgins, Garth Holman, Colleen Hyde, Tim Hogan, Steve Jessup, Joachim Kadereit, Irina Kadis, Dean Kelch, Carl Keener, Seung-Chul Kim, Dae-Sin Kim, Walter Kittredge, Ron Lance, Les Landrum, Matt Lavin, Richard LeBlond, Jenny Lellinger, Max Licher, Jeff Lomer, Marcus Lorusso, Rhoda Love, Chuck Lura, Shannon Lyons, Don Mansfield, Brian McGIll, Sue Meades, Wes Messinger, Gerry Moore, David Morgan, Louis Morin, Zack Murrell, Burrell Nelson, Gil Nelson, Ronda Newton, Dick Neubauer, Wes Niles, James Padgett, Bryan & Deta Pearce, Tony Reznicek, Kathryn Richardson, Dick Riding, Anne Rhoads, Helen & Tom Roth, Melanie Schori, Paul Shlicter, Alfred Schotz, Jim Smith, Welby Smith, Dan Spaulding, Malcolm Sproul, Matthew Stoddard, Paul Super, Alfonso Susanna, Nadia Talent, Jerry Tiehm, Rene Vaillancourt, John Vance, Pablo Vargas, Tom Vining, Ed Voss, Richard Ware, Alan Weakley, Jill Weber, Norm Weeden, David Werier, Marty Wojciechowski, Wes Wright, George Yatskievych, and Kay & Jim Young.

We have obtained living plant material from or in the following: Arnold Arboretum, Blue Ridge Parkway National Park, Death Valley National Park, Grand Canyon National Park, Great Smoky Mountain National Park, Howick Arboretum, and Quarry Hill Botanical Garden. We are indebted to the University of Maine DNA Sequencing Facility.

We have studied herbarium specimens in or borrowed specimens from Botanic Garden Edinburgh, Harvard University Herbaria, Humboldt State University, New York Botanical Garden, Philadelphia Academy of Natural Sciences, University of British Columbia, University of Maine, University of New Brunswick, University of North Carolina, University of Utah, University of Vermont, and U.S. National Herbarium.

This work was supported by National Science Foundation grants 9106226, 9806945, and 0743225 and the Maine Agricultural and Forestry Experiment Station.

Publications on Amelanchier

Campbell, C.S. 1999. The evolutionary role of hybridization in agamic complexes, with special emphasis on Amelanchier (Rosaceae). In L.W.D. van Raamsdonk & J.C.M. den Nijs (eds.). Plant evolution in man-made habitats. Proceedings of the VIIth International Symposium of the International Organization of Plant Biosystematists. pdf

Campbell, C.S., L.A. Alice*, and W.A. Wright. 1999. Comparisons of within-population genetic variation in sexual and agamospermous Amelanchier (Rosaceae) using RAPD markers. Plant Systematics and Evolution 215:157-167. pdf

Campbell, C.S., A.C. Dibble*, C.T. Frye, and M.B. Burgess*. 2008 (Accepted for publication). Amelanchier in FNA Editorial Committee, Flora of North America volume 9. Magnoliophyta: Rosidae (in part): Rosales (in part). Oxford University Press, New York.

Campbell, C.S., C.W. Greene, and S.E. Bergquist*. 1987. Apomixis and sexuality in three species of Amelanchier, shadbush (Rosaceae, Maloideae). American Journal of Botany 74:321-328. pdf

Campbell, C.S., C.W. Greene, B.F. Neubauer, & J.M. Higgins*. 1985. Apomixis in Amelanchier laevis, Shadbush (Rosaceae, Maloideae). American Journal of Botany 72:1397-1403. pdf

Campbell, C.S., M.F. Wojciechowski, B G. Baldwin, L.A. Alice*, and M.J. Donoghue. 1997. Persistent nuclear ribosomal DNA sequence polymorphism in the Amelanchier (Rosaceae) agamic complex. Molecular Biology and Evolution 14:81-90. pdf

Campbell, C.S. and W.A. Wright. 1996. Apomixis, hybridization, and taxonomic complexity in northeastern North American Amelanchier (Rosaceae). Folia Geobotancia & Phytotaxonomica 31:345-354. pdf

Campbell, C.S., W.A. Wright, T.F. Vining*, and W. A. Halteman. 1997. Morphological variation in sexual and agamospermous Amelanchier (Rosaceae). Canadian Journal of Botany 75:1166-1173. pdf

Dibble*, A.C. and C. S. Campbell. 1995. Distribution and conservation of nantucket shadbush (Rosaceae). Rhodora 97:339-349.

Dibble*, A. C., F. A. Drummond, and W. E. LaBerge. 1997. Floral syndrome in Amelanchier nantucketensis (Rosaceae) I. Floral density, bee activity, and characterization of andropetaly. Canad. J. Bot. 75: 1851–1859.pdf

Dibble*, A.C. and F.A. Drummond. 1997. Floral syndrome in Amelanchier nantucketensis (Rosaceae). II. Bee preference and diversity associated with andropetaly. Can. J. Bot. 75: 1860–1867.pdf

Dibble*, A.C., W.A. Wright, C.W. Greene, and C.S. Campbell. 1998. Quantitative morphology of the Amelanchier agamic complex (Rosaceae) at a Maine site. Systematic Botany 23:31-40. pdf

Weber*, J.E. and C.S. Campbell. 1989. Breeding systems in a hybrid of a sexual and an apomictic Amelanchier, shadbush (Rosaceae, Maloideae). American Journal of Botany 76:341-347. pdf

Publications on subtribe Pyrineae (formerly subfamily Maloideae)

Campbell, C. S. 2008a (Accepted for publication). Malacomeles in FNA Editorial Committee, Flora of North America volume 9. Magnoliophyta: Rosidae (in part): Rosales (in part). Oxford University Press, New York.

Campbell, C. S. 2008b. (Accepted for publication). Peraphyllum in FNA Editorial Committee, Flora of North America volume 9. Magnoliophyta: Rosidae (in part): Rosales (in part). Oxford University Press, New York.

Campbell, C.S. and T.A. Dickinson. 1990. Apomixis, patterns of morphological variation, and species concepts in subf. Maloideae (Rosaceae). Syst. Bot. 15:124-135. pdf

Campbell, C.S., M.J. Donoghue, B.G. Baldwin and M.F. Wojciechowski. 1995. Phylogenetic relationships in Maloideae (Rosaceae): Evidence from sequences of the internal transcribed spacer of nuclear ribosomal DNA and its congruence with morphology. Amer. J. Bot. 82:903-918. pdf

Campbell, C.S., C.W. Greene and T.A. Dickinson. 1991. Reproductive biology in the Maloideae (Rosaceae). Syst. Bot. 16:333-349. pdf

Campbell, C.S., R.C. Evans*, D.R. Morgan, T.A. Dickinson and M.P. Arsenault*. 2007. Phylogeny of Pyrinae (Rosaceae): Limited resolution of a complex evolutionary history. Pl. Syst. Evol. 266:119-145. pdf

Dickinson, T.A. and C.S. Campbell. 1991. Population structure in the Maloideae (Rosaceae). Systematic Botany 16:350-362. pdf

Evans*, R.C. and C.S. Campbell. 2002. The Origin of the apple subfamily (Rosaceae: Maloideae) is clarified by DNA sequence data from duplicated GBSSI Genes. American Journal of Botany 89:1478-1484. pdf

Publications on groups outside subtribe Pyrineae (formerly subf. Maloideae)

Alice*, L.A. and C.S. Campbell. 1999. Phylogeny of Rubus (Rosaceae) based on nuclear ribosomal DNA internal transcribed spacer sequences. American Journal of Botany 86:81-97. pdf

Alice*, L.A., T. Eriksson, B. Eriksen, and C.S. Campbell. 2001. Hybridization and gene flow between distantly related species of Rubus (Rosaceae): Evidence from nuclear ribosomal DNA internal transcribed spacer region sequences. Systematic Botany 26:769-778. pdf

Evans*, R.C., L.A. Alice*, C. S. Campbell, E.A. Kellogg, and T.A. Dickinson. 2000. The granule-bound starch synthase (GBSSI) gene in Rosaceae: multiple putative loci and phylogenetic utility. Molecular Phylogenetics and Evolution 17:388-400. pdf

Potter, D., T. Eriksson, R. Evans*, S.-H. Oh, and J. Smedmark, D. Morgan, M. Kerr, K. Robertson, M. Arsenault*, and C. Campbell. 2007. Phylogeny and classification of Rosaceae. Plant Systematics and Evolution 266:5-43. pdf

Smedmark, J. E. E., T. Eriksson, R.C. Evans*, and C.S. Campbell. 2003. Ancient Allopolyploid Speciation in Geinae (Rosaceae): Evidence from Nuclear Granule-Bound Starch Synthase (GBSSI) Gene Sequences. Syst. Biol. 52: 374-385. pdf

Wissemann, V. and C.S. Campbell. 2007. Introduction to the 2005 IBC symposium papers on the evolution of Rosaceae. Plant Systematics and Evolution 266:1-3.pdf