William (Ned) Friedman


Department of Ecology and Evolutionary Biology

University of Colorado


Plant Development and Evolutionary History

My research program focuses on the organismic interfaces between developmental, phylogenetic and evolutionary biology. Within the past fifteen years, remarkable advances in the study of the phylogenetic relationships of organisms have provided the raw materials for critical studies of character evolution in plants, animals, fungi, and all other forms of life. Armed with hypotheses of relationships among organisms, I seek to explore how patterns of morphology, anatomy and cell biology have evolved through the modification of developmental processes. My goal, with each study I initiate, is to examine the origin and subsequent radiation of a major group of photosynthetic organisms, and to reconstruct the evolutionary events that led to the establishment of defining structural and developmental features of the lineage.

Along with the origins of vascular plants and seed plants, the origin of flowering plants represents one of the three most significant evolutionary radiations of land plants during the last 475 million years. With over 250,000 extant species, angiosperms are the largest and most diverse group of plants ever to have evolved. Paradoxically, we know less about the origin and early evolutionary history of angiosperms than we do about many considerably older groups of land plants. Indeed, Darwin's "abominable mystery" continues to challenge evolutionary biologists.

Current research is devoted to understanding the early evolution of flowering plants, particularly the developmental basis for the diversification of the angiosperm female gametophyte, double fertilization, and endosperm, three of the most important and defining reproductive features of flowering plants. Our analyses draw upon a variety of approaches from cell biology to developmental biology (heterochrony and modularity) to the integration of inclusive fitness theory.

Representative Publications

Friedman, W.E. 2009. The meaning of Darwin's "abominable mystery." American Journal of Botany 96: 5-21. | pdf

Friedman, W.E. 2008. Hydatellaceae are water lilies with gymnospermous tendencies. Nature 453: 94-97. | pdf

Winther J.L. and W.E. Friedman. 2008. Arbuscular mycorrhizal associations in Lycopodiaceae. New Phytologist 177: 790-801. | pdf | commentary

Friedman, W.E. 2006. Embryological evidence for developmental lability during early angiosperm evolution. Nature 441: 337-340. | pdf

Friedman, W.E., R.C. Moore and M.D. Purugganan. 2004. The evolution of plant development. American Journal of Botany 91: 1726-1741. | pdf

Williams, J.H. and W.E. Friedman. 2002. Identification of diploid endosperm in an early angiosperm lineage. Nature 415: 522-526. | pdf