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A Taxonomic Reassessment of the Tanacetum Aureum (Asteraceae, Anthemideae) Species Group: Insights from Morphological and Molecular Data

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A Taxonomic Reassessment of the Tanacetum Aureum (Asteraceae, Anthemideae) Species Group: Insights from Morphological and Molecular Data Turkish Journal of Botany Turk J Bot (2014) 38: 1259-1273 http://journals.tubitak.gov.tr/botany/ © TÜBİTAK Research Article doi:10.3906/bot-1404-78 A taxonomic reassessment of the Tanacetum aureum (Asteraceae, Anthemideae) species group: insights from morphological and molecular data 1 2, 1 1 Marzie KAZEMI , Ali SONBOLI *, Hassan ZARE MAIVAN , Shahrokh KAZEMPOUR OSALOO 1 Department of Plant Biology, Faculty of Biological Sciences, Tarbiat Modares University, Tehran, Iran 2 Department of Biology, Medicinal Plants and Drugs Research Institute, Shahid Beheshti University, Tehran, Iran Received: 15.04.2014 Accepted: 29.10.2014 Published Online: 17.11.2014 Printed: 28.11.2014 Abstract: A taxonomic reassessment of the Tanacetum aureum (Lam.) Greuter, M.V.Agab. & Wagenitz (Asteraceae, Anthemideae) species group was performed. This group of species is distinguished by herbaceous perennial habit, (1–)2–10(–80) disciform capitula, with compact to lax corymbs and strongly to moderately convex receptacle. A multivariate statistical analysis of 58 morphological characters using Ward cluster, principal component analyses, and molecular phylogenetic analyses of nrDNA ITS and cpDNA rpL32- trnL(UAG) + ndhF-rpL32 spacer sequence datasets were conducted. Both datasets corroborated that T. aureum, T. oligocephalum (DC.) Sch.Bip., and T. heimerlii (Nábělek) Parsa could be differentiated as separate species. An identification key of the studied taxa and a description and conspectus of the species are presented. Key words: Asteraceae, ITS, morphometry, phylogeny, plastid data, SW Asia, Tanacetum 1. Introduction 1986) treated Pyrethrum as a synonym of Tanacetum. It is Tanacetum L., with 160 species, ranks as the third well known that Tanacetum is taxonomically problematic largest genus of the tribe Anthemideae of Asteraceae, in its infrageneric classification, delimitation of species following Artemisia L. and Anthemis L. (Oberprieler complexes (e.g., Tanacetum aureum (Lam.) Greuter, et al., 2006). The genus includes perennial herbs and M.V.Agab., & Wagenitz. and T. polycephalum Sch.Bip.), subshrubs distributed in the circum-Mediterranean and infraspecific taxonomy of some controversial taxa. region; central, southwestern, and eastern Asia; and parts The inconsistent taxonomic treatment and unclear of North America (Oberprieler et al., 2009). Since the species delimitations of taxa in the Tanacetum aureum benchmark monograph of Bremer and Humphries (1993) (syn.: T. chiliophyllum (Fisch. & C.A.Mey. ex DC.) Sch. on Anthemideae, no comprehensive taxonomic revision Bip.) group in various taxonomic treatments of the or updated species conspectus of the genus Tanacetum genus caused confusion and, as such, the objective of has been made available. Several authors (e.g., Greuter this investigation is to reexamine their taxonomic status et al., 2005; Mozaffarian, 2005; Djavadi, 2008; Sonboli through morphometric and phylogenetic approaches. et al., 2010a, 2010b, 2011b) have proposed new species, Pyrethrum chiliophyllum Fisch. & C.A.Mey. ex DC. combinations, and records in Tanacetum s.l. occurring in and P. oligocephalum DC. were originally described from SW Asia, and some of them have also been excluded. Few type localities of Azerbaijan and Armenia, respectively, phylogenetically deviating species from Tanacetum have in by De Candolle (1838). While the first species was turn been excluded from it and transferred to other related described based on its canescent indumentum, erect genera (e.g., Sonboli et al., 2011a; Sonboli and Oberprieler, stems, pinnatisect leaves, corymbs of 6–7 capitula, and 2012). dark brownish margins of involucral bracts, the second Several species currently listed in Tanacetum species was defined based on its (1–)2–3 capitula, long were originally described as members of the genera peduncles, and involucre of 7–9 mm in diameter and 5–6 Pyrethrum Zinn and/or Chrysanthemum L. Although mm in length. Subsequently, Schultz (1844) transferred some taxonomic treatments (e.g., De Candolle, 1838; both species to Tanacetum as T. chiliophyllum (Fisch. & Tzvelev, 1995) considered Pyrethrum and Tanacetum as C.A.Mey. ex DC.) Sch.Bip. and T. oligocephalum (DC.) independent genera, others (e.g., Grierson, 1975; Podlech, Sch.Bip., respectively. Boissier (1875) in Flora Orientalis * Correspondence: [email protected] 1259 KAZEMI et al. / Turk J Bot considered Tanacetum to be a section of the genus population of T. chiliophyllum (without designation of Pyrethrum, relegated P. chiliophyllum to a synonym of P. infraspecific rank) was published by Khandjian (1975). millefoliatum (L.) Willd., and classified P. oligocephalum Later, Chehregani and Hajisadeghian (2009) reported as a variety within P. millefoliatum (P. millefoliatum var. diploid and triploid levels, 2n = 18 and 2n = 27, for an oligocephalum (DC.) Boiss.). Pyrethrum oligocephalum Iranian population of T. chiliophyllum. The taxonomic was later transferred to P. chiliophyllum, also as a variety confusion described earlier and cytotaxonomic variations (P. chiliophyllum var. oligocephalum (DC.) Sosn.). of different populations of Tanacetum chiliophyllum Nábělek (1925) described Chrysanthemum heimerlei necessitate closer molecular analysis of this species. Nábělek from its type locality in Van Province of Turkey Molecular phylogenetic studies, based on sequence based on its black-brown hairs, numerous leaf-bearing analysis of the internal transcribed spacer region of stems from the base up to the middle, naked at top, the nuclear ribosomal DNA repeat (nrDNA ITS), have dense corymbs, numerous capitula (up to 80), and yellow frequently been used to assess the generic delimitation involucre bracts with dense long and white hairs. This and infrageneric classifications in many groups of species was later transferred to the genus Tanacetum by Anthemideae (Oberprieler, 2001; Sonboli and Oberprieler, Parsa (1949) in his account for Flora of Iran as T. heimerlii 2010, 2012; Sonboli et al., 2010a, 2010b, 2011a, 2011b, (Nábělek) Parsa. Grierson (1975) relegated this species to a 2012). Recently, a molecular phylogenetic study of the varietal rank within T. chiliophyllum (T. chiliophyllum var. genus Tanacetum s.l. (Sonboli et al., 2012) based on nrDNA heimerlei (Nábělek) Grierson). ITS and cpDNA trnH-psbA sequence variation revealed Podlech (1986) in Flora Iranica distinguished 3 varieties within T. chiliophyllum (vars. chiliophyllum, 3 accessions of T. chiliophyllum (T. chiliophyllum vars. oligocephalum, and heimerlii). This treatment was followed chiliophyllum and oligocephalum from Iran, and another T. by Mozaffarian (2008) in his account for Flora of Iran (in chiliophyllum population without designation of its variety Persian). In the synopsis of the genus Tanacetum, Bremer from Armenia) nested in separate positions across the and Humphries (1993) recognized T. chiliophyllum as clade of Tanacetum sect. Xanthoglossa. To the best of our a distinct species and considered T. heimerlii and T. knowledge, no detailed investigation of the infraspecific oligocephalum as synonyms. Tzvelev (1995), in Flora of taxonomy and molecular phylogenetic relationships of the the USSR, classified 34 Tanacetum species into 4 sections. T. aureum species group has been conducted to date Tanacetum chiliophyllum and T. oligocephalum, along with Given the heterogeneous taxonomic status of the other closely related endemic species of Azerbaijan and studied species and the paucity of data pertaining to their Nakhichevan (e.g., T. longipedunculatum (Sosn.) Tzvelev), in-depth morphology and phylogeny, this investigation were recognized as independent species within series 9, aimed to establish whether, based on the combined Chiliophylla, which includes the third largest section of the analysis of phenetic and phylogenetic data, a more precise genus (Tanacetum sect. Xanthoglossa). classification could be obtained. Hence, this investigation Recently, Greuter et al. (2005) described T. aureum was undertaken to: 1) evaluate the morphology and (Lam.) Greuter, M.V.Agab. & Wagenitz based on a present infraspecific taxonomy of T. aureum; 2) assess the specimen (Tournefort (P-TRF 4740)) originally named phylogenetic position and affinity of different varieties as Achillea aurea Lam. Accordingly, P. chiliophyllum was of T. chiliophyllum with closely related species; 3) infer considered to be a synonym of T. aureum. Kandemir taxonomic status of different varieties within species, (2012) transferred the 2 varieties of T. chiliophyllum (vars. and 4) provide a robust identification key annexed with heimerlii and oligocephalum) to T. aureum and enlisted descriptions of the taxa and a conspectus of the species. them as T. aureum var. heimerlii (Nábělek) Kandemir and T. aureum var. oligocephalum (DC.) Kandemir, respectively. 2. Materials and methods Several cytogenetic and karyological investigations have been conducted on Tanacetum (Khandjian, 1975; 2.1. Phenetic studies Chehregani and Hajisadeghian, 2009; Olanj et al., 2013 2.1.1. Plant material and references cited therein). Most Tanacetum species Herbarium specimens of Tanacetum chiliophyllum and its are diploids, showing 2n = 2x = 18 chromosomes and close allies deposited at the Herbarium of Medicinal Plants a chromosome base number of x = 9 (Watanabe, 2013 and Drugs Research Institute (MPH) of Shahid Beheshti and references cited therein). However, other reported University of Tehran, Iran, were reviewed and identified chromosome numbers include 2n = 27, 36, 54, 72, and using relevant literature
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