Abstract
Phylogenetic analyses of chloroplast gene (rbcL, matK), intron (rpl16, rps16, rpoC1) and nuclear ribosomal DNA internal transcribed spacer (ITS) sequences and chloroplast DNA restriction sites, with supplementary data from variation in size of the chloroplast genome inverted repeat, have been used to elucidate major clades within Apiaceae (Umbelliferae) subfamily Apioideae Drude. This paper summarizes the results of previously published molecular cladistic analyses and presents a provisional classification of the subfamily based on taxonomic congruence among the data sets. Ten tribes (Aciphylleae M. F. Watson & S. R. Downie, Bupleureae Spreng., Careae Baill., Echinophoreae Benth., Heteromorpheae M. F. Watson & S. R. Downie, Oenantheae Dumort., Pleurospermeae M. F. Watson & S. R. Downie, Pyramidoptereae Boiss., Scandiceae Spreng. and Smyrnieae Spreng.) are erected or confirmed as monophyletic, with Scandiceae comprising subtribes Daucinae Dumort., Scandicinae Tausch and Torilidinae Dumort. Seven additional clades are also recognized but have yet to be treated formally, and at least 23 genera examined to date are of dubious tribal or clade placement. The utility of these different molecular markers for phylogenetic inference in Apioideae is compared based on maximum parsimony analyses of subsets of previously published molecular data sets. Of the six loci sequenced, the ITS region is seen to be evolving most rapidly and rbcL is the most conservative. Intermediate in rate of evolution are matK and the three chloroplast introns; with rpl16 and rps16 evolving slightly faster than matK or rpoC1. The analysis of restriction sites, however, provided 2–4 times more parsimony informative characters than any single DNA locus sequenced, with estimates of divergence just slightly lower than that of the ITS region. The trees obtained from separate analyses of these reduced data sets are consistent with regard to the major clades inferred and the relationships among them. Similar phylogenies are obtained by combining data or combining trees, representing the supermatrix and supertree approaches to phylogenetic analysis, respectively. The inferred relationship among the tribes and informally recognized major clades within Apioideae is presented.
Highlights
‘ Umbelliferae were the first family of flowering plants to achieve general recognition, after nearly three and a quarter centuries of successive and multinational effort, considerable disagreement still exists as to the proper delimitation of the family and even more uncertainty prevails as to its natural subdivisions and the criteria on which they should be erected
Comparison of the accounts of, for example, Koch (1824), de Candolle (1830), Bentham (1867), Boissier (1872), Drude (1898), Calestani (1905) and KosoPoljansky (1916), which were erected largely on the basis of fruit morphology and anatomy, shows widely diverging opinions on the definition and composition of its tribes and subtribes. While such focus on fruit structure has been rejected by many (Heywood, 1971b, 1978a; Theobald, 1971; Davis, 1972; Cronquist, 1982; Hedge et al, 1987; Shneyer et al, 1992, 1995), the highly criticized century-old system of Drude (1898), or some modification thereof (Pimenov & Leonov, 1993), remains the most commonly used treatment
The apioid umbellifers display a remarkable array of morphological and anatomical modifications of their fruits, some of which are likely adaptations for various modes of seed dispersal (Jury, 1986). These characters are prone to convergence and their almost exclusive use to delimit suprageneric taxa has confounded attempts to identify monophyletic groups
Summary
This is necessary in order to provide the framework for lower-level revisions, as well as to interpret patterns in the evolution of the many phytochemical, anatomical, cytological, morphological and palynological characters available for the group (such as those summarized in Heywood, 1971a, and Cauwet-Marc & Carbonnier, 1982). Major clades within the subfamily have been elucidated on the basis of phylogenetic analyses of chloroplast gene (rbcL, matK ), intron (rpl, rps, rpoC1) and nuclear ribosomal DNA ITS sequences and chloroplast DNA (cpDNA) restriction sites, with supplementary data obtained from variation in size and position of the chloroplast genome inverted repeat.
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