See discussions, stats, and author profiles for this publication at: http://www.researchgate.net/publication/248021556 Molecular phylogeny of Chrysanthemum , Ajania and its allies (Anthemideae, Asteraceae) as inferred from nuclear ribosomal ITS and chloroplast trn LF IGS sequences ARTICLE in PLANT SYSTEMATICS AND EVOLUTION · FEBRUARY 2010 Impact Factor: 1.42 · DOI: 10.1007/s00606-009-0242-0 CITATIONS READS 25 117 5 AUTHORS, INCLUDING: Hongbo Zhao Sumei Chen Zhejiang A&F University Nanjing Agricultural University 15 PUBLICATIONS 56 CITATIONS 97 PUBLICATIONS 829 CITATIONS SEE PROFILE SEE PROFILE All in-text references underlined in blue are linked to publications on ResearchGate, Available from: Hongbo Zhao letting you access and read them immediately. Retrieved on: 02 December 2015 Plant Syst Evol (2010) 284:153–169 DOI 10.1007/s00606-009-0242-0 ORIGINAL ARTICLE Molecular phylogeny of Chrysanthemum, Ajania and its allies (Anthemideae, Asteraceae) as inferred from nuclear ribosomal ITS and chloroplast trnL-F IGS sequences Hong-Bo Zhao • Fa-Di Chen • Su-Mei Chen • Guo-Sheng Wu • Wei-Ming Guo Received: 14 April 2009 / Accepted: 25 October 2009 / Published online: 4 December 2009 Ó Springer-Verlag 2009 Abstract To better understand the evolutionary history, positions of some ambiguous taxa were renewedly con- intergeneric relationships and circumscription of Chry- sidered. Subtribe Artemisiinae was chiefly divided into two santhemum and Ajania and the taxonomic position of groups, (1) one corresponding to Chrysanthemum, Arc- some small Asian genera (Anthemideae, Asteraceae), the tanthemum, Ajania, Opisthopappus and Elachanthemum sequences of the nuclear ribosomal internal transcribed (the Chrysanthemum group), (2) another to Artemisia, spacer (nrDNA ITS) and the chloroplast trnL-F intergenic Crossostephium, Neopallasia and Sphaeromeria (the spacer (cpDNA IGS) were newly obtained for 48 taxa and Artemisia group). Within the Chrysanthemum group, combined with those already deposited in GenBank. Phy- Chrysanthemum Arctanthemum and Ajania were closely logenies with an emphasis on Chrysanthemum, Ajania and related to each other, and the generic circumscription was its allies, by both maximum parsimony and Bayesian ambiguous; Phaeostigma was excluded from this group analysis, were constructed using either the ITS sequence that was also confirmed by the 6-bp insertion in the IGS alone, the IGS sequence alone or combined sequences. The sequence; radiate or rare discoid Brachanthemum was IGS sequence was low phylogenetically informative, but excluded, and discoid Elachanthemum without ray florets some deletions and insertions were informative for inter- was added to this group; at the same time, Opisthopappus specific and intergeneric delimitations. The ITS and the in subtribe Tanacetinae should be transferred to subtribe ITS/IGS phylogenies both suggested the presence of two Artemisiinae and became one of the components of the major clades. The monophyly of subtribe Artemisiinae Chrysanthemum group. Based on the molecular phyloge- (clade A) could be retrieved when the phylogenetic netic framework, the evolution of pollen and capitulum characters was inferred. Keywords Ajania Á Artemisiinae Á Chrysanthemum Á H.-B. Zhao Á F.-D. Chen (&) Á S.-M. Chen Á G.-S. Wu Á Floral characters Á Molecular phylogeny Á Radiate genera Á W.-M. Guo ITS Á trnL-F College of Horticulture, Nanjing Agricultural University, 210095 Nanjing, Jiangsu, China e-mail: [email protected]; [email protected] S.-M. Chen Introduction e-mail: [email protected] G.-S. Wu There are two main groups (the Artemisia group and the e-mail: [email protected] Chrysanthemum group) in Artemisiinae (Anthemideae). W.-M. Guo According to Bremer and Humphries’s (1993) cladogram e-mail: [email protected] based on morphological characters, the Artemisia group includes Artemisia, along with eight minor Asian genera H.-B. Zhao [Ajaniopsis Shih, Crossostephium Less., Filifolium Kitam., School of Agriculture and Food Science, Zhejiang Forestry College, 311300 Lin’an, Zhejiang, China Kaschgaria Poljakov, Mausolea Bunge, Neopallasia Pol- e-mail: [email protected] jakov, Stilpnolepis H. Kraschen. (including Elachanthemum 123 154 H.-B. Zhao et al. Ling et Y.R. Ling) and Turaniphytum Poljakov] and two branches and erect corolla lobes, with both of these North American genera (Picrothamnus Nutt., Sphaerome- characters, as well as the suffruticose habit and smooth ria Nutt.) characterized by their disciform or discoid, (non-spiny) pollen, being very similar to those of Artemisia commonly paniculate capitula, microechinate and thin- (Bremer and Humphries 1993). Pollen morphology sug- walled pollen, and ribless cypsela lacking a pappus. The gests that Phaeostigma is more related to Ajania than to Chrysanthemum group is comprised of genus Chrysanthe- Artemisia (Muldashev 1981, 1983), so Shih and Fu (1983) mum, some other radiate genera with solitary or laxly cor- reduced Phaeostigma to Ajania. ymbose capitula such as Arctanthemum (Tzvelev) Tzvelev, The sister group of Artemisiinae is to be found within Tridactylina (DC.) Schultz-Bip. and Brachanthemum DC., Tanacetum L., where there are some taxa such as T. par- and the disciform Ajania Poljakov and Phaeostigma Muld. thenium (L.) Schultz-Bip. and T. tatsienense (Bureau et with dense corymbose capitula, which were once included Franchet) Bremer et Humphries, which are very similar to within a broad concept of Chrysanthemum sharing the representatives of Chrysanthemum and Brachanthemum echinate exine pollen ornamentation. However, Oberprieler (Bremer and Humphries 1993). In addition, some genera et al. (2006) placed Ajaniopsis and Elachanthemum into the such as Leucanthemella Tzvelev, Nipponanthemum Kitam Chrysanthemum group (i.e., Ajania group), and separated (subtribe Leucantheminae), Hippolytia Poljakov and the new genus Hulteniella Tzvelev from Arctanthemum.At Opisthopappus Shih (subtribe Tanacetinae) are closely the same time, they excluded Ajaniopsis, Kaschgaria, related to the Chrysanthemum group (Kondo and Abd Stilpnolepis and Turaniphytum from the Artemisia group El-Twab 2002; Zhao et al. unpublished). Both Leucan- that also was confirmed by the results of Watson et al. themella and Nipponanthemum are radiate, supporting a (2002) and Valle`s et al. (2003). radiate ancestry for the Artemisiinae (Watson et al. 2002). Chrysanthemum (formerly called Dendranthema) Oberprieler et al. (2007) transferred Leucanthemella, includes 41 species according to Oberprieler et al. (2006) Nipponanthemum and Hippolytia to subtribe Artemisiinae and is mainly distributed across eastern Asia (China, based on ITS phylogeny and morphological evidence. Korea, Japan and Siberia) (Shih and Fu 1983; Bremer and Previous molecular phylogenetic research on Artemis- Humphries 1993; Iwatsuki et al. 1997; Oberprieler et al. iinae was mainly focused on the phylogeny of Artemisia 2006). Based on capitulum characters, Chrysanthemum is and its allies (Kornkven et al. 1998, 1999; Torrell et al. divided into two sections (Sect. Chrysanthemum and Sect. 1999; Watson et al. 2002; D’Andrea et al. 2003; Valle`s Chlorochlamys) (Shih and Fu 1983). Arctanthemum (four et al. 2003; Sanz et al. 2008; Tkach et al. 2008a, b). The species) and Brachanthemum (ten species) were formerly results show the generic delimitation in this subtribe sensu classified within Chrysanthemum, but more recently they and the generic independence of several small genera such were segregated and reassembled into two new genera as Crossostephium, Filifolium, Neopallasia and Sphaer- (Shih and Fu 1983; Bremer and Humphries 1993; Iwatsuki omeria are questionable (Oberprieler et al. 2006). At the et al. 1997; Oberprieler et al. 2006). Tzvelev (in Bremer same time, probably due to fewer taxa, the phylogeny of and Humphries 1993) noted that Brachanthemum is prob- the Chrysanthemum group and intergeneric relationships of ably closely related to Chrysanthemum by its thin-walled, Chrysanthemum and allies remain poorly resolved. myxogenic fruits lacking a pappus. Arctanthemum com- We present here a reconstruction of the phylogeny of prises the rosulate herbs distributed across the Arctic, subtribe Artemisiinae and some allied or segregate taxa whereas most Chrysanthemum species are leafy herbs or from other subtribes endemic to eastern Asia, with an herbaceous perennials from China and Japan (Bremer and emphasis on Chrysanthemum, Ajania, some radiate genera Humphries 1993). and their allies based on sequence variation in both the Disciform Ajania including about 39 species (Oberpri- nuclear ribosomal DNA (ITS) and chloroplast DNA (the eler et al. 2006) was once treated as a new section of trnL-F IGS). The ITS and IGS have been successfully Chrysanthemum by Kitamura (1978) and Ohashi and used in phylogenetic studies of the Asteraceae and for Yonekura (2004). The relationship between Chrysanthe- Anthemideae specifically (Oberprieler and Vogt 2000; mum and Ajania is difficult to define. Some Chrysanthe- Oberprieler 2002; Watson et al. 2002; Valle`s et al. 2003). mum species may be more closely related to Ajania than to The main goals are: (1) to investigate the circumscription other Chrysanthemum species, and thus Chrysanthemum is of the Chrysanthemum group and intergeneric relation- even more plesiomorphic, and possibly paraphyletic with ships within this group and between the Chrysanthemum Ajania and its relatives (Bremer and Humphries 1993). group and its allies from other subtribes; (2) to reevaluate Ajania quercifolia (W. Smith)