Plants inhabited by ants (myrmecophytes) have evolved in a diversity of tropical plant lineages. Macaranga includes approximately 300 paleotropical tree species; in western Malesia there are 26 myrmecophytic species that vary in morphological specializations for ant association. The origin and diversification of myrmecophytism in Macaranga was investigated using phylogenetic analyses of morphological and nuclear ITS DNA characters and studies of character evolution. Despite low ITS variation, the combined analysis resulted in a well-supported hypothesis of relationships. Mapping myrmecophytism on all most parsimonious trees resulting from the combined analysis indicated that the trait evolved independently between two and four times and was lost between one and three times (five changes). This hypothesis was robust when tested against trees constrained to have three or fewer evolutionary transformations, although increased taxon sampling for the ITS analysis is required to confirm this. Mapping morphological traits on the phylogeny indicated that myrmecophytism was not homologous among lineages; each independent origin involved a suite of different specializations for ant-plant association. There was no evidence that myrmecophytic traits underwent sequential change through evolution; self-hollowing domatia evolved independently from ant-excavated domatia, and different food-body production types evolved in different lineages. The multiple origins of myrmecophytism in Macaranga were restricted to one small, exclusively western Malesian lineage of an otherwise large and nonmyrmecophytic genus. Although the evolution of aggregated food-body production and the formation of domatia coincided with the evolution of myrmecophytism in all cases, several morphological, ecological, and biogeographic factors appear to have facilitated and constrained this radiation of ant-plants.
Compositae (Asteraceae) are the largest flowering plant family (23,000 to 30,000 species) and its members are found throughout the world in both temperate and tropical habitats. The subfamilies and tribes of Compositae remained relatively constant for many years; recent molecular studies, however, have identified new subfamilial groups and identified previously unknown relationships. Currently there are 35 tribes and 10 subfamilies (Baldwin & al., 2002; Panero & Funk, 2002). Some of the tribes and subfamilies have not been tested for monophyly and without a clear understanding of the major genera that form each tribe and subfamily, an accurate phylogeny for the family cannot be reconstructed. The tribe Arctoteae (African daisies) is a diverse and interesting group with a primarily southern African distribution (ca. 17 genera, 220 species). They are especially important in that most of the species are found in the Cape Floral Kingdom, the smallest floral kingdom and the subject of intense conservation interest. Arctoteae are part of the monophyletic subfamily Cichorioideae s.s. Other tribes in the subfamily include Eremothamneae, Gundelieae, Lactuceae, Liabeae, Moquineae, and Vernonieae, and these were all evaluated as potential outgroups. Ultimately 29 ingroup taxa and 16 outgroup taxa with a total of 130 sequences (125 newly reported), from three genetic regions, two from chloroplast DNA (trnLF and ndhF) and one from the nuclear genome (ITS), were used to evaluate the tribe and its proposed outgroups. Each molecular region is examined separately, the chloroplast markers are examined together, and the data are combined. The data were analyzed with and without outgroups and problem taxa using parsimony and maximum likelihood methods. The analyses showed robust support for two outgroup clades, LiabeaeVernonieae and GundelieaeLactuceae and two main subtribes within Arctoteae: Arctotineae and Gorteriinae. Support for monophyly of Arctoteae is weak. Within Arctoteae, some taxa of interest are easily placed: Didelta, Cuspidia and Heterorhachis are consistently part of subtribe Gorteriinae, Cymbonotus, the Australian genus, is nested within subtribe Arctotineae, and Haplocarpha is at the base of Arctotineae. Berkheya, Haplocarpha, and Hirpicium are probably paraphyletic. Furthermore, Platycarpha most likely does not belong in Arctoteae, and Heterolepis and the tribe Eremothamneae are within Arctoteae but not within either of the two main subtribes. After some rearrangements, the two main subtribes, Arctotineae and Gorteriinae, are monophyletic and the latter has three clades. The study shows that the unusual taxa are of critical importance, and they should be included in any molecular analysis. Adequate representation of the ingroup is also important as all previous studies of Arctoteae had involved only a few taxa from the core subtribes, and so did not reveal the problems. Multiple outgroups evaluated in an iterative manner had pronounced effects on the relationships within the ingroup, not only on the pos...
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.. The University of Chicago Press is collaborating with JSTOR to digitize, preserve and extend access to International Journal of Plant Sciences. DNA sequence data from the internal and external transcribed spacers of 18S-26S nuclear ribosomal DNA and the 3 trnK intron of chloroplast DNA provide evidence for patterns of evolutionary diversification and relationships in Lasthenia. Maximum parsimony analysis shows strong support for monophyly of Lasthenia and monophyly of each of seven sections (as treated here) in Lasthenia, with minor revision of Ornduff's (1966) sectional circumscriptions. Lasthenia sect. Amphiachaenia (correct name for L. sect. Baeria sensu Ornduff 1966 and redelimited to include L. leptalea) was resolved as a basally divergent section, i.e., as the sister group of a clade comprising all other Lasthenia taxa. Placement of other sections within Lasthenia is only weakly resolved, except for L. sect. Burrielia (redefined to exclude L. leptalea) and L. sect. Hologymne, which are robust sister clades. The clade comprising L. sect. Burrielia and L. sect. Hologymne and clades corresponding to each of the other sections except L. sect. Amphiachaenia constitute a polytomy, with most clades characterized by a long basal branch and relatively short terminal branches. We suggest that overall patterns of divergence in the molecular trees for Lasthenia conform to expectations of saltational diversification, with an initial rapid radiation followed by long periods of minimal diversification in each group preceding relatively recent episodes of speciation. Results also show that several important taxonomic characters in Lasthenia are homoplastic and that the base chromosome number for Lasthenia is xp8.
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