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Tribal Relationships in the Gesneriaceae: Evidence from DNA Sequences of the Chloroplast Gene Ndhf

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Tribal Relationships in the Gesneriaceae: Evidence from DNA Sequences of the Chloroplast Gene Ndhf Tribal Relationships in the Gesneriaceae: Evidence from DNA Sequences of the Chloroplast Gene ndhF J. F. Smith; J. C. Wolfram; K. D. Brown; C. L. Carroll; D. S. Denton Annals of the Missouri Botanical Garden, Vol. 84, No. 1. (1997), pp. 50-66. Stable URL: http://links.jstor.org/sici?sici=0026-6493%281997%2984%3A1%3C50%3ATRITGE%3E2.0.CO%3B2-T Annals of the Missouri Botanical Garden is currently published by Missouri Botanical Garden Press. Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/about/terms.html. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/journals/mobot.html. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. The JSTOR Archive is a trusted digital repository providing for long-term preservation and access to leading academic journals and scholarly literature from around the world. The Archive is supported by libraries, scholarly societies, publishers, and foundations. It is an initiative of JSTOR, a not-for-profit organization with a mission to help the scholarly community take advantage of advances in technology. For more information regarding JSTOR, please contact [email protected]. http://www.jstor.org Tue Nov 13 08:53:31 2007 TRIBAL RELATIONSHIPS IN J. F. Smith2, J. C. Wolfram2," K. D. THE GESNERIACEAE: BrownZ, C. L. Carroll2, and D. S. DentonZ EVIDENCE FROM DNA SEQUENCES OF THE CHLOROPLAST GENE ndhF1 The tribal relationships of tlie Gesneriaceae are investigated using ndhF sequences. A full analysis of 70 taxa including 16 species froni the Scrophulariaceae, Bignoniaceae, and canthaceae as outgroups, resulted in two most- parsinionious trees of 5610 steps each. In all trees the Gesneriaceae were a nionophyletic group and Paulownia was tlie closest single-species outgroup for tlie analysis. Further analyses eliminated all but the members of the Gesneriaceae and Paulownia in order to better assess relationships within the family. The smaller analysis resulted in a single most- parsimonious tree of 4613 steps. The Klugieae are identified as tlie sister to tlie remainder of the family and could be separated as a distinct subfamily. The subfanlilies Cyrtandroideae (excluding Klugieae) and Gesnerioideae are monophyletic. The placement of Coronanthereae in Cyrtandroideae does not have suppol-t from this analysis, whereas its placement in Gesnerioideae is suppol-ted. Alternatively, Coronanthereae could be segregated as a separate subfamily 11ut in order to avoid a paraphyletic Gesnerioideae 15-auld either include the Sapeantheae and Beslerieae or elevate these two tribes to an additional subfamily. within Gesnerioideae the genus Sinningia is removed from tlie tribe Gloxinieae into tlie Sinningieae, which also contains the recently combined species Sinningia brasiliensis (Lietzia), as \+-ell as Paliavana and Vanhouttea. The Episcieae, Gesnerieae, Napeantheae, and Beslerieae are identified as mono- phyletic groups, as are the remainder of the Gloxinieae with Sinningia sensu lato removed. Within Cyrtandroideae, several well-supported, monophyletic lineages within the large, heterogeneous tribe Didyniocarpeae are identified, and with the current data tlie tribe Trichosporeae appears to be polyphyletic. The distribution of chromosome numbers. nodal anatomy, placental structure, and stem ~nodificationare examined based on these niolecular trees. Investigations" of higher- level cladistic relation- several -genera not having-. any strict affinity to the ships (generic, familial, and above) have recently Scrophulariaceae or other related families included drawn a great deal of attention (Annals of the Mis- in the analysis (Olmstead & Reeves, 1995). Like- souri Botanical Garden Vol. 80(3); Olmstead et al., wise, Olmstead and Reeves (1995) found that sev- 1992, 1993; Donoghue et al., 1992; Cantino, 1992; eral families traditionally segregated from the Judd et al., 1994). These analyses have provided Scrophulariaceae are best included as members of tremendous insights toward our classification sys- one of the two major lineages (e.g., Plantaginaceae). tem and process of classification, frequently draw- Although most members of the Lamiales s.1. are ing attention to families that have been separated temperate, there are some primarily tropical groups on the basis of primarily woody versus herbaceous (Gesneriaceae, Acanthaceae, Bignoniaceae). In or- taxa (Cantino, 1992; Judd et al., 1994) or tropical der to better assess whether the division between versus temperate (Judd et al., 1994). More recently these families represents another artificial segre-- an investigation of the Lamiales sensu lato has in- gation based on distribution (tropical vs. temperate) dicated that the largest family in this order, Scroph- or woody versus herbaceous (e.g., Bignoniaceae vs. ulariaceae, is unlikely to be a monophyletic group Gesneriaceae), a thorough investigation of the Ges- (Olmstead et al., 1992, 1993; Olmstead & Reeves, neriaceae was deemed necessary to complement 1995). A thorough investigation of the Scrophular- the investigations that have already demonstrated iaceae utilizing DNA sequences from both the rbcL monophyly of Acanthaceae (Scotland et al., 1995) and ndhF -genes has indicated that the family is and Bignoniaceae- (R. Olmstead, pers. comm.), but comprised of at least two monophyletic groups with have not sampled widely in the Gesneriaceae. 'Ki are indebted to the following for sharing plant material: L. E. Skog, '81. L. Wagner, J. K. Boggan, Strybing rboretum. M. Kallersjo, B. Sordenstam, R. Dunn. D. Turley, J. Katzenstein, B. Stewart. M. Evans, and tlie American Gesneriad and Gloxinia Society (AGGS) seed fund. we also thank Richard Ol~nsteadand Michael Kielin for helpful comments on the manuscript. Funding for this project. was provided by NSF grant DEB-9317775 and a grant from AGGS to JFS. ' Department of Biolog); Boise State University, 1910 University Drive, Boise, Idaho, 83725, US-I ? Current address: 1263 Londondery, Idaho Falls, Idaho, 83404, U.S.A. Volume 84, Number 1 Smith et al. 51 1997 Tribal Relationships in Gesneriaceae The Gesneriaceae are a mid-sized to large plant 1962). Another character that has been useful in family comprising approximately 2500-3500 spe- separating the subfamilies is the presence (Gesner- cies in 120-135 genera, distributed primarily in ioideae) or absence (Cyrtandroideae) of endosperm the tropics with a few temperate species in Europe, in the seed. In addition, the Gesnerioideae have a China, and Japan (Heywood, 1978; A. Weber, pers. neotropical distribution and most species have in- comm.). The majority of species in the Gesneri- ferior or semi-inferior ovaries, whereas the Cyrtan- aceae are herbaceous perennials, but can be an- droideae are primarily paleotropical with superior nuals, shrubs, lianas, and trees. Many species ovaries. However, the geographic distribution and (20%) are epiphytic, and the Gesneriaceae rank ovary position are not consistent within the subfam- among the top ten plant families in terms of abso- ilies. Therefore, although the Cyrtandroideae can lute numbers of epiphytic taxa (Madison, 1977; be defined by a synapomorphic character (uneven Kress, 1986). Given the diverse habits of the Ges- cotyledon development), the Gesnerioideae have neriaceae, it is not surprising that there is a wide been characterized by a symplesiomorphic char- array of morphological variation within the family. acter common to dicotyledons in general. Corolla tubes may be long and prominent as in Col- The two subfamilies have been divided further umnea L., or short as in Saintpaulia Wendl. Leaves into 9-17 tribes (Bentham, 1876; Burtt, 1962, are opposite in the majority of the family, but ani- 1977; Fritsch, 1893, 1894; Ivanina, 1965; Wiehler, sophylly, leading to an alternate arrangement with 1983; Burtt & Wiehler, 1995). The classification abscission of the smaller leaf, is common. Many of schemes differ due to the characters emphasized. these morphologically diverse features of the Ges- For example, Fritsch (1893, 1894) placed the Col- neriaceae are hypothesized as adaptations to the umneae in the Cyrtandroideae based on their su- epiphytic habit (Ackerman, 1986). perior ovary. Later, the Columneae were moved to The Gesneriaceae are a member of the Lamialee the Gesnerioideae due to the presence of uniform s.1. and are distinguished from other families in the cotyledons (Burtt, 1962, 1977) and combined into order by the combination of five-lobed corollas, pa- the Episcieae based on nodal anatomy (Wiehler, rietal placentation, and presence of endosperm in 1983). This paper presents a cladistic analysis of most taxa (Cronquist, 1981). However, because DNA sequences in order that phylogenetic relation- many of these characters vary within some mem- ships among taxa may be more clearly resolved, bers of the Gesneriaceae (including variation within and a more stable classification scheme proposed. individuals of some species), there has been con- The gene ndhF is a chloropfast gene that in to- siderable confusion regarding the placement of bacco encodes
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