Transfer of Dodecatheon to Primula Primulaceae Author s Austin R Mast and James L Reveal Source Brittonia 59 1 79 82 2007 Published By The New York Botanical Garden DOI 10 1663 0007 196X 2007 59 79 TODTPP 2 0 CO 2 URL http www bioone org doi full 10 1663 0007 196X 282007 2959 5B79 3ATODTPP 5D2 0 CO 3B2 BioOne www bioone org is a a nonprofit online aggregation of core research in the biological ecological and environmental sciences BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies associations museums institutions and presses Your use of this PDF the BioOne Web site and all posted and associated content indicates your acceptance of BioOne s Terms of Use available at www bioone org page terms of use Usage of BioOne content is strictly limited to personal educational and non commercial use Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors nonprofit publishers academic institutions research libraries and research funders in the common goal of maximizing access to critical research Transfer of Dodecatheon to Primula Primulaceae Austin R Mast1 and James L Reveal2 Department of Biological Sciences Florida State University Tallahassee FL 32306 U S A e mail amast bio fsu edu 2 University of Maryland College Park Maryland 20742 5815 U S A and The New York Botanical Garden Bronx New York 10458 5126 U S A mailing address 18625 Spring Canyon Road Montrose Colorado 81401 7906 U S A e mail jreveal umd edu 1 Abstract Phylogenies inferred from both chloroplast and nuclear DNA regions have placed the small genus Dodecatheon the shooting stars 17 spp among the descendants of the most recent common ancestor of Primula the primroses ca 430 spp This is congruent with conclusions previously derived from morphology and cytology We illustrate how failure to formalize this information in the circumscription of Primula might have impeded understanding of the evolution of distyly as seen in Primula and buzz pollination as seen in Dodecatheon To avoid future confusion we make new combinations for Dodecatheon in Primula and erect Primula sect Dodecatheon in Primula subgenus Auriculastrum for the taxa from Dodecatheon Key words laceae Buzz pollination distyly Dodecatheon heterostyly Primula Primu Recent molecular evidence from both the chloroplast cpDNA K llersj et al 2000 Mast et al 2001 2004 2006 Trift et al 2002 and nuclear Martins et al 2003 genomes is congruent with prior hypotheses based on morphological and cytological evidence Thompson 1953 Wendelbo 1961 Richards 1993 Holmgren 1994 that Dodecatheon L 17 spp is most closely related to one of the seven subgenera of Primula L ca 430 spp Primulaceae The study by Mast et al 2004 represents the most extensive sampling of cpDNA characters seven regions and taxa within Dodecatheon 16 of the 17 spp recognized here and the closely related Primula subgen Auriculastrum Schott all four sections sampled three of them exhaustively to date In the cpDNA phylogeny inferred in that study sampled members of Dodecatheon form a monophyletic group 97 bootstrap frequency that is sister to the only species of Primula sect Suffrutescens A J Richards 100 bootstrap frequency of P subgen Auriculastrum The study by Martins et al 2003 represents the most complete sampling of nuclear DNA in this case the two internal transcribed spacers of the nuclear ribosomal DNA and subgenera of Primula five of seven to date In the nuclear DNA phylogeny inferred in that study the single sampled species of Dodecatheon is sister to the single sampled species of P subgen Auriculastrum 100 bootstrap frequency Thus DNA sampling of two independent linkage partitions sensu Slowinski Page 1999 is congruent in support of the conclusion that Primula is paraphyletic with respect to Dodecatheon While the buzz pollinated flowers of Dodecatheon distinguish the lineage from others in Primula individuals of Dodecatheon in sterile pre anthetic and fruiting stages are strikingly similar to members of P subgen Auriculastrum Both groups have members that grow in moist alpine meadows and along streamsides have chromosome numbers of 2n 44 and produce valvate capsules on long scapes arising from a rosette of fleshy lance Brittonia 59 1 2007 pp 79 82 2007 by The New York Botanical Garden Press Bronx NY 10458 5126 U S A ISSUED 30 March 2007 80 brittonia shaped leaves with involute vernation Thompson 1953 Furthermore Thompson p 75 noted that D jeffreyi Van Houtte and P parryi A Gray a member of P subgen Auriculastrum are virtually indistinguishable when the corollas and inserted anthers are removed Both Dodecatheon and P subgen Auriculastrum occur in western North America though each is also found elsewhere one sp of Dodecatheon extends across the Bering Strait P subgen Auriculastrum is also found in Pacific coastal Asia and in Europe Some previous authors e g Pax 1889 grouped Dodecatheon at the tribal rank as Cyclameae Dumort with another buzz pollinated genus the Mediterranean genus Cyclamen L which was still then considered part of the family c f K llersj et al 2000 However recent authors Thompson 1953 Wendelbo 1961 Richards 1993 2002 Holmgren 1994 have taken the geographical morphological and cytological similarities as evidence for a close phylogenetic relationship between Dodecatheon and Primula and interpreted floral similarities between Dodecatheon and Cyclamen as due to convergence on buzz pollination Dodecatheon is one of four genera composed of species that have been segregated from Primula but that are inferred to be descended from the most recent common ancestor MRCA of Primula using morphological and cytological Wendelbo 1961 cpDNA K llersj et al 2000 Mast et al 2001 2004 2006 Trift et al 2002 and nuclear DNA Martins et al 2003 evidence Like 92 of species in Primula one of the four segregated genera Dionysia Fenzl ca 50 spp of mostly cushion plants from the Middle East Grey Wilson 1989 Trift et al 2004 is distylous In distylous plants a genetic polymorphism produces two floral types the pin morph has stigmas high and anthers low in the corolla tube and small pollen the thrum morph has stigmas low and anthers high in the corolla tube and large pollen Darwin 1877 Ganders 1979 Barrett 1992 2002 Richards 2002 Like the remaining 8 of species in Primula the three other
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