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Within the Senecioneae (Asteraceae) Based on Chloroplastdna EV

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Within the Senecioneae (Asteraceae) Based on Chloroplastdna EV American Journal of Botany 82(12): 1567-1573. 1995. THE ORIGIN OF DENDROSENECIO WITHIN THE SENECIONEAE (ASTERACEAE) BASED ON CHLOROPLAST DNA EVIDENCE! ERIC B. KNOX2,3 AND JEFFREY D. PALMER Herbarium and Department of Biology, University of Michigan, Ann Arbor, Michigan 48109-1048; and Department of Biology, Indiana University, Bloomington, Indiana 47405 Chloroplast DNA restriction-site variation was surveyed using 15 enzymes for 37 accessions from the tribe Senecioneae (Asteraceae), plus two outgroup species, in order to determine the placement within the tribe of the giant senecios (Den­ drosenecio). The survey revealed 176 phylogenetically informative mutations and 121 autapomorphic mutations. Dendro­ senecio is diagnosed by a minimum of 15 mutations, which suggests that the giant senecios evolved from a relatively isolated lineage within the Senecioneae, and this conclusion is supported by earlier evidence from chromosome counts and phytochemistry. Among the taxa sampled, the closest relatives of Dendrosenecio are Cineraria deltoidea and two species of Euryops. Support was not found for suggestions in the literature that the closest relatives of Dendrosenecio are species in Solanecio or Senecio subgen. 'Crociseris.' The position of the Dendrosenecio/Cineraria/Euryops clade is weakly sup­ ported as basal to the majority of other senecionoid genera. The tussilaginoid genera sampled (Ligularia, Petasites, Roldana, and Tussilago) are monophyletic in our analysis, with the surprising inclusion of Pericallis hybrida as the sister-taxon to Roldana suffulta. The sister-group to the Dendrosenecio/Cineraria/Euryops clade includes all species of Delairea, Gynura, Kleinia, Packera, Senecio, and Solanecio sampled. Within Senecio, subgenera Senecio and 'Crociseris' form a monophyletic core, with subgenus 'Kleinioidei' being broadly paraphyletic or possibly polyphyletic. Since the discovery of Dendrosenecio johnstonii est relatives of the giant senecios. Several species in Se­ (Oliv.) B. Nord. (Asteraceae) on Mt. Kilimanjaro, Tan­ necio subgen. "Crociseris' (see Table I for authors) and zania, by H. H. Johnston (1886), the giant senecios have the segregate genus Solanecio have been suggested. attracted considerable interest and discussion by taxono­ Within Senecio, these include S. subsessilis (Engler, mists and evolutionary biologists. These plants typically 1892) and other coarse herbs from upland eastern Africa have large leaf-rosettes borne on thick, woody stems that in sect. Plantaginei (S. ochrocarpus and S. karaguensis branch after flowering to produce candalabra-like growth O. Hoffm.; Mabberley, 1974; Nordenstam, 1978), the forms the size of telephone poles. They have diversified coarse European herbs S. umbrosus and S. doronicum in on ten mountains in eastern Africa (Knox, 1993a; Knox sect. Crociseris (Mabberley, 1974), and the South Amer­ and Palmer, 1995) to form 17 taxa (1 1 species and six ican S. hualtata DC. in sect. Hualtatini (DC.) Bak. (Hau­ nonautonymic infraspecific taxa; Knox, 1993b), and the man, 1935). The two Solanecio species are robust shrubs adaptive radiation of this group involved much conver­ from tropical Africa (S. mannii; Carlquist, 1974, p.554) gent or parallel evolution, resulting in a "mosaic of vari­ and Ethiopia, (S. gigas; Nordenstam, 1978). With the ex­ ation" in morphological features (Mabberley, 1973) that ception of Senecio karaguensis and S. hualtata, the re­ confounded earlier attempts to reconstruct their evolu­ maining species were included in a preliminary survey tionary history. The giant senecios were initially treated (Knox, 1993c) of restriction-site variation, which includ­ as Senecio (Oliver, 1887), a placement explicitly sup­ ed the North American herb Packera aurea and used the ported by several workers (Cotton, 1932, 1944; Hedberg, cultivar Pericallis hybrida (formerly treated as Cineraria; 1957; Mabberley, 1973, 1974) until Nordenstam (1978) Jansen et al., 1990; Jansen, Michaels, and Palmer, 1991) recognized Dendrosenecio as a genus distinct from a as the outgroup species. This survey found that these pu­ more narrowly delimited Senecio (Knox, 1993b). tative close relatives to Dendrosenecio shared a more re­ A recurrent question in the literature concerns the clos- cent common ancestry with each other, and that Perical­ lis, Dendrosenecio, and the clade comprising all other I Manuscript received 16 December 1994; revision accepted 23 May 1995. taxa sampled were separatedby long basal branches. In The authors thank the Tanzanian Commission for Science and Tech­ the present study, this preliminary survey of the tribe nology, the Institute Zairois pour la Conservation de la Nature, the Senecioneae was therefore expanded using readily avail­ Office Rwandais du Tourisme et des Pares Nationaux, and Addis Abeba able material (Table 1) in order to determine the place­ University and the Ethiopian Ministry for Foreign Affairs for permis­ ment of Dendrosenecio within the tribe and to identify sion to conduct research in their respective countries; R. K. Jansen for DNA and helpful comments on the manuscript; and M. C. Burd, R. R. potential close relatives. Kowal, the Missouri Botanical Garden, and the Royal Botanic Gardens, In the most recent treatment of the Senecioneae, Jef­ Kew for seed or leaf tissue. This work was supported in part by grants frey (1992) accepted the two subtribes (Blennospermati­ from the National Science Foundation (BSR-8800487, BSR-8901123, nae and Senecioninae) delimited by Nordenstam (1977) BSR-8996262) and awards from the Rackham School of Graduate Stud­ and proposed many provisional infrasubtribal groupings ies, The University of Michigan. that were not formally described. His stated purpose was 2 Present address: Department of Biological Sciences, Rutgers Uni­ versity, Newark, NJ 07102. to provide a list of putative monophyletic genera (and 3 Author for correspondence (Fax: 201-648-1007). subordinate groups within the still heterogeneous genus 1567 Ul 0\ 00 TABLE 1. Accessions examined for cpDNA restriction-site variation (classification of tribe Senecioneae according to Jeffrey, 1986; 1992. Classification Species and author Accession" Geographic origin Outgroup species (Asteraceae) Bamadesia caryophylla (Veil.) S. F. Blake UMMBG (Knox 2439) Peru Eupatorium atrorubescens (Lem.) Nicholson UMMBG (Knox 2438) Mexico Tribe Senecioneae Subtribe Blennospermatinae Rydb. Blennosperma nana (Hook.) S. F. Blake Ornduff 83739 (UC) California Subtribe Senecioninae Dum. Tussilaginoid genera Ligularia stenocephala (Maxim.) Matsum & G. Koidz Kowal 3092 Asia Petasites japonicus (Siebold & Zucc.) Maxim. Kowal 3093 Sachalin Islands Roldana suffulta (Greenm.) H. Rob. & Brettell Kowal et al. 3007 Mexico Tussilago farfara L. Kowal s.n. 3094 Europe and Asia Senecionoid genera Neotropical group Dendrosenecio kilimanjari (Mildbr.) E. B. Knox Knox 50 E. Africa ssp. cottonii (Hutch. & G. Taylor) E. B. Knox Synotoid group Delairea odorata Lem. UMMBG (Knox 2422) S. Africa Packeroid group Packera aurea (L.) (A. Love & D. Love) Knox 880 N. America Senecionoid group Cineraria deltoidea Sond. Knox 2037 S. and E. Africa ;l> Euryops dacrydioides Olivo Burd s.n. (Knox 2436) E. Africa ~ E. pectinatus Casso MBG 893711 S. Africa t'!'I Gynura aurantiaca (Blume) DC. UMMBG (Knox 2424) Indonesia n;.:l Othonna capensis L. H. Bailey UMMBG (Knox 2423) S. Africa > Pericallis hybrida B. Nord. Knox 884 Canary Islands Z Solanecio gigas (Vatke) C. Jeffrey Knox 633 Ethiopia ...... 0 S. mannii (Hook. f.) C. Jeffrey Knox 555 E. Africa c Kleinia neriifolia Haw. UMMBG (Knox 2425) Canary Islands ;.:l Kleinia Mill. subgen. Kleinia Z Subgen. Notonia (DC.) O. Hoffm. K. fulgens Hook. f. UMMBG (Knox 2426) S. Africa > K. pendula (Forssk.) DC. UMMBG (Knox 2430) Arabia and NE. Africa r- K. stapeliiformis (Phillips) Stapf UMMBG (Knox 2429) 0 S. Africa "!1 Senecio L. subgen. 'Crociseris' OJ Sect. Crociseris (Reichenb.) Hall. & Wohlf. Senecio coriaceous Kowal 2926 Asia 0 S. doronicum L. RBG 007.83.60046 Europe .., > S. umbrosus Waldst. & Kit. Hubl s.n. (Knox 2398) Europe Z Sect. Plantaginei Harv. S. inomatus DC. Knox 870 S. and E. Africa -< S. ochrocarpus Olivo & Hiern Knox 637 Ethiopia S. subsessilis Olivo & Hiern Knox 454, 795 E. Africa Subgen. 'Jacobaea' sect. Jacobaea (Mill.) Dumort. ser. Erucifolii (Rouy) Schischk. S. erucifolius L. RBG 0064875 Europe S. jacobaea L. RBG 0066949 Europe and N. Asia Subgen. 'Kleinioidei' sect. Kleinioidei (DC.) Harv. S. scaposus DC. UMMBG (Knox 2431) S. Africa Sect. Rowleyani C. Jeffrey S. citriformis Rowley UMMBG (Knox 2428) sw. Africa S. mandraliscae (Tineo) Jacobsen UMMBG (Knox 2433) S. Africa S. rowleyanus M. Jacobs UMMBG (Knox 2427) W. Africa Subgen. Senecio sect. Suffruticosi Greenm. S. stoechadiformis DC. Kowal 3051 Mexico Sect. Senecio S. madagascariensis Poir. RBG 0020518 Madagascar S. sylvaticus L. RBG 0066994 Europe and N. Asia S. vulgaris L. Knox 2411 Europe (Cosmop.) 'Burd = M. C. Burd; Hubl = E. Htibl; Kowal = R. R. Kowal; MBG = Missouri Botanical Garden; Ornduff = R. Ornduff; RBG = Royal Botanic Gardens, Kew; UMMBG = University of Michigan Matthaei Botanical Garden. '< 2. 00 N December 1995] KNOX AND PALMER-THE ORIGIN OF DENDROSENEClO 1569 Senecio) for the selection of representative species in fu­ shared mutations found a single island of eight shortest ture systematic studies. We have used Jeffrey's informal trees of 349 steps, with a consistency index (CI; Kluge classification for the species included in our survey (Ta­ and Farris, 1969) of 0.50. The eight shortest trees
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