Polyploidy: Tricky Taxonomic Delimitation Ben Grady UW-Madison Botany Department 6 March 2007 Polyploidy and Taxonomy: an overview • • • • “Defining” species Polyploid complexes Cardamine pratenesis Lepidium What do we call a species? • Biological Species Concept – Mayr 1942 • Evolutionary Species Concept – Wiley 1978 • Phylogenetic Species Concept – Cracraft 1989 • General consensus: diagnosable units with common evolutionary history From Futuyma 1998 and Judd et al. 2002 The Polyploid Complex • Continued polyploidization in a group or lineage • Much overlap in morphological traits and ecological preferences • Difficult and time-consuming to fully understand relationships within a polyploid complex Grant 1981 The Polyploid Complex • Wow, this is complex! The Polyploid Complex • Count some chromosomes 2n 2n 2n 2n 2n 4n 4n 2n 4n 6n 2n The Polyploid Complex • Determine source of genetic material 2n 2n 2n 2n 2n 4n 4n 2n 4n 6n 2n The Polyploid Complex - Simplified 2n 2n 2n AA AA AA 4n AAAA 4n AAAA 6n AAAAAA The prevalence of Polyploid Complexes • Polyploid complexes are common in certain families – Polypodiaceae – Trilliaceae – Onagraceae – Poaceae – Brassicaceae Grant 1981 Cardamine pratensis complex • Widespread distribution • Morphological variation not discrete • Post-glacial diversification • Hybridization and polyploidy rampant – 2n = 16, 24, 30, 32, 40, 44, 46, 48, 56 • Iberian taxa: – C. pratensis s.s. – C. crassifolia – C. castellana Photos © Carl Farmer Lihova et al. 2003 Lihova et al. 2003 C. pratensis C. pratensis C. castellana C. crassifolia Lihova et al. 2003 C. pratensis in the Iberian Peninsula Lihova et al. 2003 • C. crassifolia and C. castellana appear distinct (both taxa 2n=16) C. castellana C. crassifolia AFLP analysis: C. castellana & C. crassifolia C. castellana 2n = 16 C. crassifolia 2n = 16 Lihova et al. 2003 Many intermediate characters Lihova et al. 2003 Origins of C. castellana? • C. castellana morphologically intermediate • Possible hybrid of C. pratensis and C. crassifolia • What then? – Species? – Subspecies? – No taxonomic recognition? C. pratensis in the Iberian Peninsula Lihova et al. 2003 • C. pratensis s.s. still poses a problem, or does it? – Various polyploids still mostly interfertile (except diploids) – Morphological variation continuous – Multiple origins of polyploids? AFLP analysis: C. pratensis s.s. 30 16 56 56 44 Lihova et al. 2003 Morphological Variation? Figure 5 Lihova et al. 2003 Allopolyploidization – Lepidium Lee et al. 2002 • Lepidium – 175 spp. worldwide – Crazy flowers, well for a mustard… PI gene tree and evolution of stamen arrangement, green-Eurasia, orangeAfrica, blue-S.Amer., black-Austr., purple-N.Amer. lacking lateral stamens, References • Futuyma, D.J. 1998. Evolutionary Biology. Sinauer Associates – Sunderland, Mass. U.S. • Grant, V. 1981. Plant Speciation. Columbia University Press, New York, pp. 324-346. • Judd W.S., C.S. Cambell, E.A. Kellogg, P.F. Stevens, & M.J. Donoghue. 2002. Plant Systematics: An Evolutionary Approach. Sinauer Associates – Sunderland, Mass. U.S. • Lee, J., K. Mummenhoff, & J. Brown. 2002. Allopolyploidization and evolution of species with reduced floral structures in Lepidium L. (Brassicaceae). PNAS 99(26): 16835-16840. • Lihova, J., A. Tribsch, & K. Marhold. 2003. The Cardamine pratensis (Brassicaceae) group in the Iberian Peninsula: taxonomy, polyploidy and distribution. Taxon 52: 783-801. • Marhold, K., & J. Lihova. 2006. Polyploidy, hybridization and reticulate evolution: lessons from the Brassicaceae. Pl. Syst. Evol. 259: 143-174.