Chapter 44 Compositae metatrees: the next generation Vicki A. Funk, Arne A. Anderberg, Bruce G. Baldwin, Randall J. Bayer, J. Mauricio Bonifacino, Use Breitwieser, Luc Brouillet, Rodrigo Carhajal, Raymund Chan, Antonio X.P. Coutinho, Daniel J. Crawford, Jorge V. Crisci, Michael O. Dillon, Susana E. Freire, Merce Galhany-Casals, Nuria Garcia-Jacas, Birgit Gemeinholzer, Michael Gruenstaeudl, Hans V. Hansen, Sven Himmelreich, Joachim W. Kadereit, Mari Kallersjo, Vesna Karaman-Castro, Per Ola Karis, Liliana Katinas, Sterling C. Keeley, Norhert Kilian, Rebecca T. Kimball, Timothy K. Lowrey, Johannes Lundberg, Robert J. McKenzie, Mesjin Tadesse, Mark E. Mort, Bertil Nordenstam, Christoph Oberprieler, Santiago Ortiz, Pieter B. Pelser, Christopher P. Randle, Harold Robinson, Nddia Roque, Gisela Sancho, John C. Semple, Miguel Serrano, Tod F. Stuessy, Alfonso Susanna, Matthew Unwin, Lowell Urbatsch, Estrella Urtubey, Joan Valles, Robert Vogt, Steve Wagstaff, Josephine Ward and Linda E. Watson INTRODUCTION volumes listed the tribes mostly in the order of Bentham 1873a rather than beginning with Heliantheae, which Constructing a large combined tree of Compositae, a Bentham thought was most primitive (Bentham 1873b). 'metatree' (also called 'meta-supertree' by Funk and The papers in the 1977 volumes did accept some changes Specht 2007 and 'megatree' by R. Ree, pers. comm.) such as the recognition of Liabeae and the conclusion allows one to examine the overall phylogenetic and bio- that Helenieae were not a 'good' group, both more or geographic patterns of the family. The first modern at- less accepted by Cronquist in 1977. However, most pro- tempts to understand the family were by the authors in posed changes such as the new tribe Coreopsideae, etc. Heywood et al. (1977) plus the paper by Cronquist (1977), were not accepted by the synantherological community. which was initially intended to be in the Heywood Cronquist (1977) believed that the primitive characters publication. Literature prior to 1977 has been discussed of the family were as follows (slightly modified): shrubby; in detail in other chapters (for the early literature, see leaves opposite; inflorescence cymose; heads few, each Chapter 1). In Cronquist's 1977 paper he reaffirmed his with many florets; involucre leafy, several-seriate; recep- agreement with Bentham's 13-tribe classification of the tacle chaffy; ray florets present and fertile; disk florets family and the concept that Heliantheae s.l. were the perfect and fertile; lobes of the disk corollas with well primitive members (Cronquist 1955; Bentham 1973a, b). developed mid-vein; pappus chaffy, of five members; and Cronquist (1977) pointed out that the Heywood et al. anthers connate, not tailed. Cronquist stated that the 748 Funk et al. presence of ray florets may have predated the origin of has been some discussion on the pros and cons of the Compositae, so that even discoid tribes might have had 'supertree' and 'supermatrix' methods (Steel et al. 2000; a radiate ancestry. Gatesy et al. 2002; Bininda-Emonds et al. 2003), and The acceptance of the modified Bentham system was both methods are compared with the metatree approach not universal. There were at least two papers in the by Funk and Specht (2007). The metatree for this analysis Heywood et al. volumes (Jeffrey 1977; Skvarla 1977) was constructed in the following manner: and two individuals who published elsewhere (Carlquist 1. A 'base tree' was formed from the phylogeny of 1966, 1976; Robinson 1981) who had reservations about Panero and Funk (2008) with a few alterations. The the concept of "13 tribes rooted in the Heliantheae". All most important change was the addition of taxa of these dissenting authors observed that the data they from the Heliantheae Alliance. The Heliantheae were generating did not support all of the above-listed Alliance section of the Panero and Funk tree characteristics as primitive in the family. However, for (which had only a few taxa) was replaced with the the most part, the synantherological community contin- branching pattern of the Heliantheae Alliance from ued to use the Bentham classification. Baldwin (Baldwin et al. 2002; Chapter 41). Also, Not too long after 1977, opinions began to change with some refinements were made using the work of the advent of cladistic methodology and molecular data. Ortiz (Chapters 18 and 19) and Ortiz et al. (Chapter Jansen and Bremer and their collaborators (Bremer 1987, 17) for Carduoideae, and Funk and Chan (Chapter 1992, 1994; Jansen and Palmer 1987, 1988; Hansen 1991a, 23) for Cichorioideae. The base tree was reduced b; Jansen et al. 1991a, b; Bremer and Jansen 1992; Jansen to a matrix using Brooks Parsimony Analysis and Kim 1996; Bremer and Gustafsson 1997) reordered (BPA; Brooks 1982; Brooks and McLennan 2002), Compositae by placing Barnadesiinae as the sister group of wherein any branching diagram can be reduced to the family and placing Heliantheae (including Eupatorieae) a series of zeros and ones in a data matrix. We used highly nested in the phylogeny of the family. MacClade to generate the data matrix (Maddison Bremer's cladistic analysis (1994) was the first revi- and Maddison 2001). The data matrix was run in sion of the whole family based on morphology since a tree program (PAUP 4.0M0; Swofford 2002) to Bentham, and he recognized many of the problem areas check for errors. All trees have been "ladderized to in the cladograms of the family and tribes, but the mor- the right" for consistency, although anyone familiar phology did not generate enough data to resolve many of with cladistics will understand that the tree can be the issues. Over ten years later Kadereit and Jeffrey (2007) "rotated" at any node. This feature is amply dem- reordered the genera, tribes, and subfamilies within the onstrated by comparing the rooted tree (Fig. 44.1) family based on morphology and molecular results, and and the unrooted tree (Fig. 44.2). this work is now the standard reference for descriptions 2. The most recent (and available) tree for each clade of the tribes and genera of the family. (see below) was reduced to a matrix (as above) and This chapter seeks to link the most recent molecular these matrices were added to the original matrix. trees together in a metatree framework (Funk and Specht Each time a new clade tree was added, the overall 2007) and to use that tree to provide a basis for under- analysis was re-run to insure an accurate replication standing the systematics, evolution, and biogeography of of the newly added tree, as well as to confirm that the family. the addition did not result in topological changes elsewhere in the metatree. It should be noted that when a phylogeny for a tribe contained many taxa MATERIALS AND METHODS from the same area in a monophyletic group or a grade, these were often pruned to decrease the size Construction of the metatree of the tree without subtracting any biogeographi- The metatree for Compositae was developed using a com- cal information. For instance, the phylogeny of pilation of trees. The name metatree was adopted for this Gnaphalieae contained a clade of 58 terminal taxa type of tree because it is a "tree of trees", one that is based all endemic to Australia; this clade was reduced to on a fixed 'base tree' topology (Funk and Specht 2007). 25 taxa. This type of tree has also been called a meta-supertree or 3. A summary tree (Fig. 44.1) was produced in which megatree (R. Ree, pers. comm.), and some authors refer each major clade was reduced to a single branch. to it as a supertree. It is, however, neither a tree produced This tree also shows the phylogenetic position of by a combined analysis of coded cladograms obtained critically placed taxa and is displayed as an unrooted from individual datasets (classic 'supertrees') nor is it the tree in Fig. 44.2. result of analyzing a dataset in which data from multiple See the section on optimization for an explanation of datasets have been combined ('supermatrix' trees). There the biogeographic areas and how they were assigned. Chapter 44: Compositae metatrees: the next generation 749 Sources of the trees extending eastwards through Paraguay to Uruguay and General references for this study were Bremer (1994), southern Brazil and down through Argentina to southern Heywood (1993), Heywood et al., (1977), Hind (1996), Patagonia (Heywood 1993). and Kadereit and Jeffrey (2007). Below, the origin of each Cassini, in his famous 1816 diagram (Chapter 41: Fig. phylogeny on the metatree is discussed. 41.1), showed Calyceraceae and Campanulaceae to be closely related to Compositae. Even though he did not Outgroups have it in the diagram, he also thought Goodeniaceae Lundberg (Chapter 10) examined the relationships among were close (see Chapter 1). the families now contained in Asterales, including Com- positae. His work indicated that Calyceraceae were the Compositae sister group of Compositae (1st outgroup) and that Good- The base tree. — The basic structure of the tree was eniaceae (2 outgroup) were the sister group of the taken from Panero and Funk (2002, 2008) and Baldwin Calyceraceae + Compositae clade. The next most closely (Baldwin et al. 2002; Chapter 41); see above for details. related family is Menyanthaceae, and it is followed by a The trees in Panero and Funk (2008) contained exten- clade containing Stylidiaceae, Alseuosmiaceae, Phellinac- sive sampling from the base of the tree, Mutisieae (sensu eae, and Argophyllaceae. The distribution ofthese eight fam- Cabrera), three to ten genera representing all other tribes ilies (Fig. 44.1) shows that the Compositae + Calyceraceae (including the Heliantheae Alliance), and many taxa that clade is nested in a grade of Australasian taxa (Australia, had been "hard to place" in previous studies (includ- New Guinea, New Caledonia, and New Zealand). Each ing Hecastodeis, Gymnarrhena, and Corymbium).