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Floral Micromorphology and Its Systematic Implications in the Genus Sinosenecio (Senecioneae-Asteraceae)

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Floral Micromorphology and Its Systematic Implications in the Genus Sinosenecio (Senecioneae-Asteraceae) Plant Syst Evol (2011) 291:243–256 DOI 10.1007/s00606-010-0385-z ORIGINAL ARTICLE Floral micromorphology and its systematic implications in the genus Sinosenecio (Senecioneae-Asteraceae) Ying Liu • Qin-Er Yang Received: 24 January 2010 / Accepted: 31 October 2010 / Published online: 7 December 2010 Ó Springer-Verlag 2010 Abstract Several floral microcharacters (configuration of southern China is the diversity center of this genus, with stigmatic areas on the inner surface of the style branch, the majority of species occurring there (Chen 1999). Both shape of the anther apical appendage, anther size, and S. newcombei (Greene) J.P. Janovec & T.M. Barkley, a configuration of the endothecial thickenings and of the species endemic to the Queen Charlotte Islands, BC, filament collar) in 63 populations representing 35 species Canada (Janovec and Barkley 1996), and S. koreanus and one variety of Sinosenecio (Senecioneae-Asteraceae) (Kom.) B. Nord., a species occurring in Korea and north- were investigated. The floral micromorphological data eastern China, have been shown by ITS sequence data to be obtained are highly consistent with evidence from molec- elements of Tephroseris, whereas T. changii B. Nord., a ular systematics and cytology, strongly suggesting a poly- species endemic to Jinfoshan Mountain, Nanchuan County, phyletic nature of the currently circumscribed Sinosenecio Chongqing Municipality, China, may find a home in Sino- and the necessity of a taxonomic change at generic level. senecio (Pelser et al. 2007; Wang et al. 2009). In the tribe Senecioneae, Sinosenecio is generally considered to be Keywords Asteraceae Á Floral microcharacter Á most closely related to Nemosenecio and Tephroseris Senecioneae Á Sinosenecio Á Systematics Á Taxonomy (Nordenstam 1978, 2007; Jeffrey and Chen 1984; Chen 1999; Pelser et al. 2007; Wang et al. 2009; Nordenstam et al. 2009). In their taxonomic studies on the tribe Sene- Introduction cioneae of Eastern Asia, Jeffrey and Chen (1984) removed these three genera from Tussilagininae and placed them in a Sinosenecio B. Nord. (Senecioneae-Asteraceae), a genus new subtribe, Tephroseridinae, based on their aberrant segregated from the heterogeneous genus Senecio by chromosome number of n = 24 and some shared floral Nordenstam (1978), currently contains about 40 species. microcharacters, but other authors (e.g., Jeffrey 1992; All the species occur in China, only two of which extend Bremer 1994; Nordenstam 2007; Pelser et al. 2007; outside this country (to Myanmar and Indochina) (Jeffrey Nordenstam et al. 2009) did not recognize this subtribe and Chen 1984; Chen 1999; Wang et al. 2009). Central and because of its lack of diagnostic characters and its phylo- genetic position deeply nested in the subtribe Tussilagininae. Within the tribe Senecioneae, floral microcharacters Q.-E. Yang (&) have long been widely used for subtribal and generic Key Laboratory of Plant Resources Conservation and delimitations (e.g., Robinson and Brettell 1973; Norden- Sustainable Utilization, South China Botanical Garden, stam 1978; Vincent and Getliffe 1988, 1992; Vincent 1996; Chinese Academy of Sciences, Xingke Road, Tianhe District, Guangzhou 510650, People’s Republic of China Jeffrey 1992; Karis 1993; Bremer 1994; Liu 2001), e-mail: [email protected] although their taxonomic value was once questioned due to considerable intraspecific variability (Wetter 1983). When Y. Liu Nordenstam (1978) established Sinosenecio, he used two State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of sciences, floral microcharacters in its characterization, namely the Beijing 100093, People’s Republic of China configuration of the anther endothecial cell wall thickenings 123 244 Q.-E. Yang, Y. Liu and of the filament collar. Based upon his observation of B. Nord., S. homogyniphyllus and S. subcoriaceus C. Jef- S. homogyniphyllus (Cumm.) B. Nord., the type species of frey & Y.L. Chen, are clustered with the typical genera of the genus, he described the endothecial tissue in the genus subtribe Tussilagininae (‘‘tussilaginoid’’) sensu Jeffery and as ‘‘polarized’’ (with the ribs or thickenings confined to the Chen (1984), including Cremanthodium, Farfugium, Lig- horizontal walls of the endothecial cells) or ‘‘not polarized’’ ularia, Parasenecio, Petasites, Syneilesis and Tussilago. (with rather sporadic ribs or thickenings on both vertical Some other species, such as S. bodinieri (Vaniot) B. Nord., and horizontal walls). He observed the configuration of the S. euosmus (Hand.-Mazz.) B. Nord., S. fanjingshanicus filament collar in three Sinosenecio species and found that C. Jeffrey & Y.L. Chen, S. globigerus (Chang) B. Nord., the filament collar is of only one type in the genus, i.e., the S. guangxiensis C. Jeffrey & Y.L. Chen, S. oldhamianus ‘‘cylindrical’’ type. In these two characters, Nordenstam (Maxim.) B. Nord. and S. septilobus (Chang) B. Nord., are (1978) found that Sinosenecio is markedly different from nested with Tephroseris and Nemosenecio species in a the typical Senecio members that have ‘‘radial’’ endothecial well-supported clade (‘‘tephroseroid’’ clade = subtrib. thickenings and ‘‘balusterform’’ filament collars. In their Tephroseridinae Jeffrey & Chen). Yet another species, S. taxonomic studies on the tribe Senecioneae of Eastern Asia, hainanensis (Chang & Tseng) C. Jeffrey & Y.L. Chen, Jeffrey and Chen (1984) also attached great importance to somewhat surprisingly, is nested in the subtribe Seneci- the value of floral microcharacters for subtribal and generic oninae. Also, the results of our extensive cytological delimitations. Furthermore, they noticed the variation pat- studies do not indicate the monophyly of Sinosenecio (Liu tern of endothecial thickenings within Sinosenecio and and Yang 2010b). The results have provided us with reli- underscored the taxonomic value of this character for the able chromosome number data for 32 of the total ca. 40 genus. They used the character as one of the major distin- species currently recognized in the genus. The mitotic guishing features for its infrageneric classification. In their numbers (2n) counted include 26, 48 (seldom 72, 96), 58 studies Jeffrey and Chen (1984) distinguished three main and 60 (seldom 90), so we can postulate four basic chro- types of anther endothecial tissue within Sinosenecio based mosome numbers for the genus: x = 13, 24, 29 and 30 on the alignment of thickenings on the cells. They described (Table 1). Of these, the numbers of x = 30 and 24 are the the tissue as ‘‘strictly polarized’’ when the thickenings were commonest, occurring in 11 and 19 species respectively, restricted to the transverse or horizontal walls. When the while the numbers of x = 13 and x = 29 each occur only thickenings were distributed along all the inner walls, they in one species, S. oldhamianus and S. hainanensis, described the tissue, not consistently, as ‘‘radial’’ or ‘‘scat- respectively. Interestingly, all the species clustered toge- tered.’’ When the tissue contained both strictly polarized ther with the typical Tussilagininae genera have a basic and radial cells, they described it as ‘‘radial and polarized.’’ chromosome number of x = 30; those nested in the teph- They divided the genus into two sections (sect. Sinosenecio roseroid clade all have x = 24 except for S. oldhamianus and sect. Phyllocaulon C. Jeffrey & Y.L. Chen) as differ- with x = 13; and S. hainanensis, the only species nested entiated by the type of the endothecial tissue, claiming that within Senecioninae, has x = 29 (Table 1; Liu and Yang, the former is characterized primarily by strictly polarized in prep.). endothecial thickenings, whereas the latter is characterized Taking into account the evidence from both molecular by radial or radial and polarized thickenings. In their paper, systematics and cytology, therefore, the genus Sinosenecio they illustrated the endothecial thickenings of three spe- as currently circumscribed is not monophyletic, and cies, viz. S. subrosulatus (Hand.-Mazz.) B. Nord. (strictly includes at least three major and only distantly related polarized) within sect. Sinosenecio, and S. koreanus and assemblages of species that have different basic chromo- S. septilobus (Chang) B. Nord. (radial and polarized) within some numbers, one with x = 30, the other with x = 24 sect. Phyllocaulon, but did not state explicitly the actual (rarely 13), and yet another with x = 29. When viewing number of species that they had examined. Liu (2001) results of the molecular systematic and cytological studies reported the configuration of the endothecial thickenings in against the infrageneric classification of Sinosenecio pro- 12 Sinosenecio species belonging to the two sections and posed by Jeffrey and Chen (1984) and later totally adopted recognized two types of thickenings, ‘‘polarized’’ and by Chen (1999) (see Table 1 for the classification scheme), ‘‘scattered’’ [= ‘‘radial’’ in the sense of Nordenstam (1978)], we noticed that the three major assemblages of species but regrettably he largely ignored the value of his findings revealed by basic chromosome numbers and ITS phylo- for the systematic considerations of the genus. genetic analyses are largely incongruent with the distinc- Recent ITS phylogenetic analyses have seriously chal- tion between the two aforementioned sections (sect. lenged the monophyly of Sinosenecio (Pelser et al. 2007; Sinosenecio and sect. Phyllocaulon) that was ostensibly Wang et al. 2009; Liu and Yang, unpublished). In these made by Jeffrey and Chen (1984)
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