Hybridization is an important part of the evolutionary history of flowering plants. If hybridization has occurred among the species of a taxon under cladistic analysis the results are varied but always present additional difficulties. Hybridization results in incongruent intersecting data that obscure the underlying hierarchy. Guidelines and methods are examined for their usefulness in identifying possible hybrids in a cladistic study. Seven genera are analyzed cladistically and the resulting cladograms examined for possible hybrids. These hypotheses of hybridization are then compared to other data, such as distribution and cytology, to see if the hypotheses of hybridization are supported or rejected. The more hybrids an analysis contains and the more complex the interactions, the more difficult it becomes to identify possible hybrids and their parents. It is difficult to overemphasize the importance of hybridization and polyploidy in evolution because they are outstanding features of many plant groups. According to some authorities 30-80% of the species of angiosperms are polyploids (Goldblatt, 1979; Lewis, 1979; Stebbins, 1974), which allows for the possibility of a tremendous amount of hybridization. Of course, these figures do not include diploid hybrids. Despite its importance, hybridization has been virtually ignored by those who have dominated the discussion of evolutionary theory. This is a result, no doubt, of evolutionary theory being largely in the hands of scientists who work on groups in which such phenomena as polyploidy and hybridization have a strong relationship with unisexuality and are not considered important in evolution. There are different types of hybridization. Figure 1 summarizes some of the possibilities (a more detailed explanation is found in Funk, 1981) but does not include introgression. For the purpose of this paper it is important to note that many hybrids are sexually reproducing individuals and are morphologically distinct and in some manner reproductively isolated from both parents. Thus, they behave as species no matter whose definition you chose to adopt. THE STUDY OF HYBRIDIZATION The basic concept of phylogenetic systematics (sensu Hennig, 1966) is an ever branching pattern or hierarchy. The method of cladistics (phylogenetic systematics) seeks to discover these patterns by grouping together taxa that share apomorphies (evolutionarily novel, unique, or derived characters). Hybridization, or reticulate evolution, is inconsistent with a method designed to depict hierarchies. Hybridization is, therefore, a cause of incongruent, intersecting data that obscure phylogenetic information. Cladists have been concerned with this problem for several years. Most realize that any method that seeks to identify patterns of relationship must be able to accommodate hybridization because of its frequency. Workers in the problems of hybridization and phylogenetic systematics include Bremer (1983), Bremer and Wanntorp (1977), Clark (1982), Funk (1981), Humphries (1983), Humphries and Funk (1984), Nelson (1983), Nelson and Platnick (1980), Rosen (1979), Wagner (1969, 1983), Wanntorp (1983), and Wiley and Brooks (1982). One favorite method of botanists in estimating the closeness of relationships among taxa is the percentage of hybridization in crossing studies. An important point about such hybridization studies was made by Rosen (1979: 277): reproductive compatibility is a primitive attribute for the members of a lineage and has, therefore, no power to specify relationships within a genealogical framework. We cannot use the ability of two or more species to hybridize as an indication of close relationship because the ability is relaI A number of people, who do not necessarily agree with everything that I have said, have kindly provided me with data and comments on various drafts of the manuscript and this paper would not have been possible without their assistance. They include: R. Jansen, J. Semple, C. Clark, R. Sanders, C. Humphries, H.-E. Wanntorp, K. Bremer, G. Nelson, N. Platnick, D. Rosen, and P. Weston. I appreciate the assistance of B. Kahn in helping prepare the illustrations. 2 Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. ANN. MISSOURI BOT. GARD. 72: 681-715. 1985. This content downloaded from 207.46.13.148 on Sun, 11 Sep 2016 04:15:12 UTC All use subject to http://about.jstor.org/terms 682 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 72