Collectanea Botanica 33: e001 enero-diciembre 2014 ISSN-L: 0010-0730 http://dx.doi.org/10.3989/collectbot.2013.v33.001

Taxonomical and nomenclatural notes on : A proposal of classification, a description of new sections and subsections, and a species list of the redefined sectionCentaurea

A. Hilpold1,2, N. Garcia-Jacas1, R. Vilatersana1 & A. Susanna1

1 Institut Botànic de Barcelona (IBB-CSIC-ICUB), pg. del Migdia, s/n, Parc de Montjuïc, ES-08038 Barcelona, 2 Museum of Nature South Tyrol, Bindergasse, 1, IT-39100 Bozen, Author for correspondence: A. Susanna ([email protected])

Editor: L. Sáez

Received 6 March 2014; accepted 11 April 2014

Abstract Ta x o n o m i c a l a n d n o m e n c l a t u r a l n o t e s o n Ce n t a u r e a : A p r o p o s a l o f c l a s s i f i c at i o n , a d e s c r i p t i o n o f n e w s e c t i o n s a n d s u b s e c t i o n s , a n d a s p e c i e s l i s t o f a r e d e f i n e d s e c t i o n Ce n t a u r e a .— In this paper, we summarize the results of our long-date research on the Centaurea. The first part of the paper deals with the overall classification of the genus, which we propose to divide into three subgenera: subgenus Centaurea, subgenus Cyanus and subgenus Lopholoma. The second part of this publication gives a recopilation of the species of the redefined section Centaurea, a group that includes former sections Acrolophus (sect. Centaurea s. str.), Phalolepis and Willkommia, together with taxonomical, geographical, ecological and karyological considerations. Finally, new descriptions or nomenclatural combinations are proposed to correlate nomenclature to the new classification: a new combination (sect. Acrocentron subsect. Chamae- cyanus) is proposed in subgenus Lopholoma; three new sections (sects. Akamantis, Cnicus, and Hyerapolitanae) are described in subgenus Centaurea; two subsections (subsects. Phalolepis and Willkommia) in sect. Centaurea; and three subsections (subsects. Exarata, Jacea, and Subtilis) in sect. Phrygia.

Key words: Acrocentron; Acrolophus; Akamantis; Chamaecyanus; Cyanus; Exarata; Hierapolitanae; Jacea; Lopholo- ma; Phalolepis; Phrygia; Subtilis; Willkommia.

Resumen No t a s t a x o n ó m i c a s y n o m e n c l a t u r a l e s e n Ce n t a u r e a : p r o p u e s t a d e clasificación , descripción d e s e c c i o n e s y subsecciones n u e v a s , y l i s t a d e e s p e c i e s d e u n a s e c c i ó n Ce n t a u r e a r e d e f i n i d a .— En este trabajo presentamos los resultados de nuestras investigaciones de larga fecha en el género Centaurea. La primera parte del trabajo trata de la clasificación del género, que proponemos dividir en tres subgéneros: subgénero Centaurea, subgénero Cyanus y subgénero Lopholoma. La segunda parte es una recopilación de las especies de la redefinida sección Centaurea, que incluye las antiguas secciones Acrolophus (sect. Centaurea s. str.), Phalolepis y Willkommia, junto con consideraciones geográficas, ecológicas y cariológicas. Por último, proponemos nuevas secciones, subsecciones y combinaciones para correlacionar nomenclatura y clasificación: proponemos una nueva (sect. Acrocentron subsect. Chamaecyanus) en el subgénero Lopholoma; se describen tres secciones nuevas (sects. Akamantis, Cnicus y Hyerapolitanae) en el subgénero Centaurea; dos subsecciones (subsects. Phalolepis and Willkommia) en la sección Centaurea; y tres subsecciones (subsects. Exarata, Jacea y Subtilis) en la sección Phrygia.

Palabras clave: Acrocentron; Acrolophus; Akamantis; Chamaecyanus; Cyanus; Exarata; Hierapolitanae; Jacea; Lo- pholoma; Phalolepis; Phrygia; Subtilis; Willkommia.

Cómo citar este artículo / Citation Hilpold, A., Garcia-Jacas, N., Vilatersana, R. & Susanna, A. 2014. Taxonomical and nomenclatural notes on Centaurea: A proposal of classification, a description of new sections and subsections, and a species list of the redefined section Centaurea. Collectanea Botanica 33: e001. doi: http://dx.doi.org/10.3989/collectbot.2013.v33.001

Copyright © 2014 CSIC. Este es un artículo de acceso abierto distribuido bajo los términos de la licencia Creative Commons Attribution-Non Commercial (by-nc) Spain 3.0. 2 A. HILPOLD, N. GARCIA-JACAS, R. VILATERSANA & A. SUSANNA

INTRODUCTION indicated that we classify Centaurea three subgen- era, namely Centaurea, Cyanus (Mill.) Cass. ex Systematics of Centaurea L. has changed dramati- Hayek, and Lopholoma (Cass.) Dobrocz. Thereaf- cally during the past two decades. Molecular meth- ter, our aims with this paper are: (1) to present an ods have resolved the old problem of the delimi- overall classification of the genus, detailing espe- tation of the genus (for details see Susanna et al., cially our proposal for subgenus Centaurea; (2) to 1995; Garcia-Jacas et al., 2000, 2001, 2006; Hilpold offer a list of species of the newly defined section et al., 2014). To put it shortly, Centaurea as defined Centaurea; and (3) to propose some new nomen- traditionally (e.g. Linnaeus, 1753; Bentham, 1873; clatural combinations within subgenera Centaurea Hoffmann, 1894; Dostál, 1976) was polyphyletic, and Lopholoma. and molecular studies showed that some clades, including the former section Centaurea (includ- ing the species, Centaurea centaurium L.) had MATERIAL AND METHODS to be excluded in order to make the genus mono- phyletic. To prevent the renaming of hundreds of For subgenera Cyanus and Lopholoma, our outline species, the type species was changed: the type is is based on the studies by Boršić et al. (2011) and now C. paniculata L. (Greuter et al., 2001). The old Font et al. (2002, 2009), respectively. Regarding section Centaurea acquired the rank of genus on its subg. Centaurea, our proposal follows the layout own, namely Vaill. (Greuter, 2003; suggested by Garcia-Jacas et al. (2006) and Hilpold Greuter et al., 2005). However, there are some open (2012), and is partly provisional for the Eastern questions on the classification of the genus that are Mediterranean (EMC) and Western Mediterranean discussed in the work here presented. (WMC) clades, which are in need of a more com- Centaurea constitutes an important member of plete molecular survey. tribe Cardueae Cass. (Compositae). It is formed As regards to the list of species of sect. Centau- by annual, biennial or perennial herbs, less often rea, we have followed current literature, basically shrubs, with usually unarmed leaves (Susanna & floras. follows Euro+Med Plantbase Garcia-Jacas, 2007). They are also characterized by (Euro+Med, 2006–2013) for the Eastern and Cen- a lateral hilum (Dittrich, 1968), and a specialized tral Mediterranean region, and López & Devesa floral morphology with showy sterile peripheral (2008a, b, c, d, 2010, 2011) and López et al., (2011) florets without staminodes (Wagenitz & Hellwig, for the . More details on the con- 1996a). The main morphological character used struction of the Table 2 are given below. for systematics within Centaurea is the form of the scarious appendages. Centaurea is insect pollinated, mainly by bees and bumblebees (Har- DISCUSSION AND CONCLUSIONS rod & Taylor, 1995; Bilisik et al., 2008; Albrecht et al., 2009; McIver et al., 2009). The genus occurs On the light of our previous results, we summa- mainly in the Mediterranean and Irano-Turanian rize our proposal of a classification for the genus regions, with some species (especially from sect. (new sections, subsections and nomenclatural Jacea) extending northwards to temperate . combinations are formally proposed at the end of Redefined Centaurea is the largest genus within the article): subtribe Centaureinae encompassing some 250 species (Susanna & Garcia-Jacas, 2007, 2009), but Centaurea L. these figures are conservative and somewhat dis- puted because of wide differences in taxonomical Subgenus Lopholoma (Cass.) Dobrocz. treatments. Besides, the numbers are steedly rising Sect. Acrocentron (Cass.) DC. with the description of new species in the Eastern Subsect. Acrocentron Mediterranean (Aksoy et al., 2008). Subsect. Chamaecyanus (Willk.) new sta- We published a general outline of a classifica- tus and combination tion of the genus in Susanna & Garcia-Jacas (2007, Sect. Stephanochilus (Coss. & Durieu ex 2009), but these were general works and we only Benth. & Hook. f.) O. Hoffm.

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 Taxonomical and nomenclatural notes on Centaurea 3

Subgenus Cyanus (Mill.) Cass. ex Hayek The genus Centaurea contains three well delim- Sect. Cyanus ited subgenera: Lopholoma (often named Acrocen- Subsect. Cyanus tron), Cyanus and Centaurea (the latter corresponds Subsect. Perennes Boiss. to the Jacea group sensu Garcia-Jacas et al., 2006). Cyanus has been also treated as separate genus Subgenus Centaurea (e.g. Greuter, 2003). We suggest treating them in Eastern Mediterranean Clade (EMC) the rank of subgenera. These three subgenera are Sect. Calcitrapa DC. [incl. sects. Seridi- well separated on the basis of molecular markers oides DC. and sect. Tetramorphaea (DC.) (Garcia-Jacas et al., 2001; Susanna et al., 2006; Boiss.] Barres et al., 2013) and correspond to the existing Sect. Chartolepis (Cass.) DC. pollen types. Sect. Cheirolepis (Boiss.) O. Hoffm. [incl. The pollen of the genus Centaurea is, like that sect. Plumosipappus (Czerep.) Wagenitz, of all Compositae (Blackmore et al., 2009), tri- sect. Pseudoseridia Wagenitz p. p. and sect. colporate and the ectexine is formed by two lay- Pteracantha Wagenitz] ers of columellae (Wagenitz, 1955). Within the Sect. Cynaroides Boiss. ex Walp. [incl. sect. genus, four clearly distinct pollen types can be Paraphysis (DC.) Wagenitz] found, which correspond to the three subgenera Sect. Grossheimia (Sosn. & Takht.) Dittrich (Wagenitz, 1955; Martín & Garcia-Jacas, 2000; Sect. Microlophus (Cass.) DC. Susanna & Garcia-Jacas, 2009). Centaureinae Sect. Phaeopappus (DC.) O. Hoffm. show a dysploid chromosome series, with basic Sect. Pseudophaeopappus Wagenitz chromosome numbers ranging between x = 16 Sect. Ptosimopappus (Boiss.) O. Hoffm. and x = 7 (Garcia-Jacas et al., 1996). Low num- Sect. Rhizocalathium Tzvelev (incl. sect. bers can be found in more evolved groups and Pseudoseridia Wagenitz p. p.) have been interpreted as an adaptation to dry con- ditions (Susanna & Garcia-Jacas, 2009). Chromo- Western Mediterranean Clade (WMC) some numbers in Centaurea range from x = 12 to Sect. Hymenocentron (Cass.) DC. x = 7 (Table 1). Whereas subgenera Cyanus and Sect. Melanoloma (Cass.) DC. (incl. sect. Centaurea have a broad range of chromosome Gymnocyanus Maire) numbers, subgenus Lopholoma is more conserva- Sect. Mesocentron (Cass.) DC. tive, showing only two chromosome numbers. Sect. Seridia (Juss.) DC. Chromosome numbers in the EMC and in the WMC follow largely the sectional division. The Circum-Mediterranean Clade (CMC) lowest chromosome number known to date in the Sect. Centaurea genus was found in the annual Centaurea patula Subsect. Centaurea DC. with x = 7. Subsect. Phalolepis (Cass.) new combina- Centaurea is rich in secondary metabolites, which tion may mostly be a protection against herbivores (Ol- Subsect. Willkommia (Blanca) new combi- son & Kelsey, 1997; Susanna & Garcia-Jacas, nation 2009). Some of them show antimicrobial activity Sect. Akamantis new section (Karioti et al., 2001; Ugur et al., 2009). Metabolites Sect. Ammocyanus Boiss. are predominantly sesquiterpene lactones (Tarasov Sect. Cnicus (L.) new status and combination et al., 1975; Koukoulitsa et al., 2005; Karamend- Sect. Hierapolitanae new section eres et al., 2007), but also flavonoids (Zapesoch- Sect. Phrygia Pers. naya et al., 1978; Nacer et al., 2006), essential oils Subsect. Phrygia (Altintas et al., 2004) and phenols (Bubenchikov et Subsect. Exarata, new subsection al., 1992). Many phytochemical studies have been Subsect. Jacea (L.) new status and combi- published over the last decade but only very few nation address explicitly systematic questions (but see Subsect. Subtilis, new subsection Yildirim et al., 2009).

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 4 A. HILPOLD, N. GARCIA-JACAS, R. VILATERSANA & A. SUSANNA

Table 1. Chromosome numbers in Centaurea. ? = unknown number.

Genus Centaurea Basic chromosome number x Subgenus Lopholoma Sect. Acrocentron 10, 11 Sect. Stephanochilus ? Subgenus Cyanus Sect. Cyanus 8, 9, 12 Sect. Perennes 10, 11 Subgenus Centaurea 7, 8, 9, 10, 11, 12 East Mediterranean Clade (EMC) 8, 9, 10 West Mediterranean Clade (WMC) 8, 10, 11, 12 Circum-Mediterranean Clade (CMC) Sect. Akamantis 9 Sect. Ammocyanus 7, 8 Sect. Centaurea 9 Sect. Cnicus 11 Sect. Hierapolitanae 8, 9 Sect. Phrygia 11

Subgenus Lopholoma Subgenus Cyanus

This subgenus comprises about 100 species (Font et The subgenus stands out by its predominantly bright al., 2002) and is distributed all over the Mediterrane- blue , as shown by its most prominent mem- an with one species, Centaurea scabiosa L., reach- ber, the cornflower (Centaurea cyanus L.; Fig. 1B). ing the high north of Europe. The heads of Further characters for this subgenus are two pol- Lopholoma are usually larger than in the other sub- len types (Cyanus and Montana types of Wagenitz, genera, and the bract appendages often ends in a long 1955) and the pectinate-ciliate, unarmed, decurrent spine (Fig. 1A). The group is also morphologically appendages of the phyllaries (Wagenitz & Hellwig, well differentiated by presenting an only and char- 1996a). The subgenus comprises about 40 species, acteristic pollen type, the Centaurea scabiosa type also concentrated in the Mediterranean, reaching (Wagenitz, 1955). This subgenus was studied apply- central Asia and the Caucasus. Molecular studies of ing molecular tools by Font et al. (2002, 2009). There the group were carried out by Boršić et al. (2011), are two groups within the subgenus that should be Olšavská et al. (2011) and Löser (2013). The study treated as two different sections: The former mono- of Boršić et al. (2011) confirmed the existence of typic genus Stephanochilus Coss. & Dur. ex Benth. two clearly distinct clades already marked by the is sister to all other members of the subgenus (Font pollen types and life cycles, corresponding to sub- et al., 2002) and constitutes the first section. The sec- sects. Cyanus (annuals) and Perennes (perennials). ond section, section Acrocentron, encompasses all other species of the subgenus. On molecular basis, Subgenus Centaurea Font et al. (2009) suggested that sect. Chamaecya- nus Willk. is not clearly distinguishable from sect. This subgenus corresponds to the Jacea group sensu Acrocentron despite clear morphological differences Garcia-Jacas et al. (2006) and it is the most species- (a very short and simplified pappus combined with rich. It is represented by several hundreds of species acaulescent habit). We propose to merge Chamae- mainly in the Mediterranean Region but also in West- cyanus as subsection in section Acrocentron. ern Asia and Central and Eastern Europe. The most

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 Taxonomical and nomenclatural notes on Centaurea 5

Figure 1. Photographs of the genus Centaurea, with special emphasis on subgenus Centaurea and sect. Centaurea. (A), Subgen. Lopholoma: C. ornata, Soria, Spain (Photograph: A. Susanna); (B), Subgen. Cyanus: Centaurea cyanus, Soria, Spain (Photograph: A. Susanna). (C) Subgen. Centaurea: East-Mediterranean Clade (EMC): (C), C. lycopifolia, Barcelona Botanical Garden (Photo- graph: A. Susanna). (D–N), Circum-Mediterranean Clade (CMC). (D), C. benedicta, Barcelona Botanical Garden (Photograph: A. Susanna); (E), C. akamantis, Avakas gorge, (Photograph: M. Galbany); (F), C. hierapolitana, Afon lake, (Photogra- ph: A. Susanna); (G), C. hyrcanica (Jacea-Phrygia group), (Photograph: A. Pirani); (H), C. exarata, Barcelona Botanical Gar- den (Photograph: A. Susanna); (I), C. patula, Barcelona Botanical Garden (Photograph: A. Susanna). (J–N), Sect. Centaurea. (J), C. tenorei, Minori, Italy (Photograph: A. Hilpold); (K), C. alba, Sierra de Aracena, Spain (Photograph: L. Barres); (L), C. pulvinata, Sierra de Abrucena, Spain (Photograph: G. Blanca); (M), C. horrida, (Photograph: S. Pisanu); (N), C. princeps, Barcelona Botanical Garden (Photograph: A. Susanna); (O), C. panormitana, Sferracavallo, (Photograph: A. Hilpold).

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 6 A. HILPOLD, N. GARCIA-JACAS, R. VILATERSANA & A. SUSANNA characteristic trait of the subgenus is the pollen Section Phrygia — Members of section Phrygia type, the so-called Jacea pollen type (Wagenitz, can be distinguished from those of section Centau- 1955; Garcia-Jacas et al., 2000). New systematic rea by (1) the basic chromosome numbers, x = 11 vs. insights of the subgenus on molecular grounds x = 9 respectively, (2) the leaf shape, since Phrygia were provided by Garcia-Jacas et al. (2006). One leaves are entire, while in section Centaurea at least of the main results of this study, based on ITS se- the basalmost leaves are deeply divided. Section quence analyses, was the separation of subgen. Phrygia also has a different ecology and distribution: Centaurea into three clades, which where named species of sect. Centaurea are much more dry adapt- after their main geographic distribution: the East- ed and bound to Mediterranean climate, whereas the ern Mediterranean Clade (EMC) comprises spe- vast majority of species of sect. Phrygia are typical cies often with spiny bract appendages from the elements of montane and subalpine meadows, influ- Eastern Mediterranean (Fig. 1C shows C. lycopi- enced by frequent mowing, grazing or avalanches. folia Boiss. & Kotschy ex Boiss.), including the They are dependent on a good and constant water widely distributed weed Centaurea calcitrapa L.; supply and are quite resistant to cold periods. These the Western Mediterranean Clade (WMC), the adaptations permitted the group also to disperse into members of which also show pronounced spiny the high north of Europe, with species like Centau- bract appendages, well visible in the common rea nigra L. or C. jacea L. Taxonomy within sect. weeds L. or C. sulphurea Phrygia is highly complex, and the latest attempts Willd.; and the Circum-Mediterranean Clade to elucidate it using molecular markers (Koutecký et (CMC), the members of which are predominantly al., 2011; López-Alvarado, 2012) are only partially not or only slightly spiny in their bract append- satisfying because of the very low levels of variation ages. Despite their wide distribution, their impor- found. The apparent lack of any intrinsic breeding tance as weeds, and the high species diversity, our barriers led to the description of many hybrids on one knowledge about systematics within the EMC and hand (Vanderhoeven et al., 2002; Koutecký, 2007; the WMC is poor and would need a more compre- Vonica & Cantor, 2011). On the other hand, a split hensive study with molecular methods. The CMC, of the breeding communities into diploid and tetra- however, was already the centre of attention of ploid lineages, connected by frequent polyploidi- profound molecular work (see below). zation events, was observed (Hardy et al., 2000; The most distinctive characters of the CMC Koutecký, 2007; Koutecký et al., 2011). Traditional are also the bract appendages (Fig. 1D–N). There systematics based on morphology within this group are three extreme forms: membranaceous, long uses merely the shape of the bract appendages for ciliate-fimbriate, and reduced or missing append- the subdivision into two groups: Phrygia (formerly ages. According to Garcia-Jacas et al. (2006) Lepteranthus) with long, fimbriate bract appendages most species within the CMC belong to two and Jacea with either short ciliate or membranaceous main clades: the Jacea-Phrygia group (hereafter bract appendages. Intermediate forms between these referred to as section Phrygia) and the Centau- extremes are commonly attributed to hybridization rea group (formerly Acrolophus subgroup; here- events (Vanderhoeven et al., 2002; Koutecký, 2007; after referred to as section Centaurea; Wagenitz Vonica & Cantor, 2011). We suggest keeping these & Hellwig, 1996a). Following Hilpold (2012), two groups the rank of subsections. besides these two groups, four more sections belong to the CMC: the new monotypic section Section Centaurea — The type section has its Akamantis consisting of Centaurea akamantis T. centre of distribution around the Mediterranean and Georgiadis & Hadjik. from Cyprus, section Am- the Black Sea (Fig. 2). Highest species numbers mocyanus composed of C. ammocyanus Boiss. can be found in the Balkan Peninsula, Italy, Turkey and C. patula, the new monotypic section Cnicus and the Iberian Peninsula (in order of abundance). consisting of Centaurea benedicta (L.) L. (for- Almost all African species are concentrated in the merly Cnicus benedictus L.) and the new sec- NW of the continent, in the Atlas mountain ranges. tion Hierapolitanae, including C. hierapolitana A few widespread species reach central Europe and Boiss. and C. tossiensis Freyn & Sint. ex Freyn the Baltic Sea (C. stoebe L.) and Middle Asia, Af- from Turkey. ghanistan and (C. virgata Lam.).

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 Taxonomical and nomenclatural notes on Centaurea 7

C. stoebe s. l.

C. stoebe ~30 C. virgata ~30 ~5 ~40

~30 ~10 ~50 ~30 ~5 ~40 ~5 ~10 ~5 ~5 ~10 C. cyprensis C. cyrenaica C. damascena

Figure 2. Distribution of sect. Centaurea (without introduced populations). Dark grey: areas with more than one species occu- rring. Light grey: only one species occurring, species name is given. Numbers show the approximate species number in the area. Note that the species numbers reflect, besides real differences in diversity, also the species concepts used in the different areas.

All members of sect. Centaurea grow in dry, removed from sect. Centaurea (Garcia-Jacas et al., open vegetation. Many species can be found in fre- 2006; Hilpold et al., 2009). Counts of chromosome quently pastured garrigues, on the coast or in the in- numbers differing from the basic number x = 9 are terior. Furthermore, open rocks are inhabited. Only extremely rare: 2n = 44 (one single count by Baden exceptionally, high mountain areas were colonized. (1983) in C. affinisFriv., usually with x = 9), 2n = 16 Some species are specialized in sandy beaches (e.g. (in C. deustiformis Adamović; Strid, 1983; only in C. spinosa L.). Both substrates, calcareous and sili- one out of many counts in C. diffusa Lam.; Banche- ceous soils are inhabited with predominance on the va & Greilhuber, 2006), and 2n = 32 (in C. arenaria first ones. Members of sect. Centaurea are rarely Willd.; Bancheva & Greilhuber, 2006). Figure 3 selfing or even obligate outcrossers (Harrod & shows the percentages of counted species and the Taylor, 1995; Hardy et al., 2004). Reproduction is distribution of the known chromosome numbers. usually sexual (Noyes, 2007), although facultative A few members of sect. Centaurea are used as apospory has been reported (Cela Renzoni & Viegi, ornamentals, like L. (“Velvet 1982). Centaurea”, cf. Ellis, 1999). Some species are used Species of sect. Centaurea have a basic chromo- as medicinals, especially in folk medicine, because some number of x = 9. Most counted populations of its secondary compounds (Nacer et al., 2006; are diploid (2n = 18; Fig. 3). Tetraploids (2n = 36) Akkol et al., 2009). The sesquiterpene lactone are frequent, and they have been subject of recent cnicin, found in Centaurea benedicta (the name of scientific work (Španiel et al., 2008; Mráz et al., the compound derives from its former name Cnicus 2012). Three species are hexaploid (2n = 6x = 54): benedictus L.) and other species of sect. Centaurea C. carystea Trigas & Constantin., C. cithaeronea (Olson et al., 1997; Erel et al., 2011) is sometimes Phitos & Constantin. and C. tuzgoluensis Aytaç & H. used for bitter tonics (Tešević et al., 2007). Duman; cf. Phitos & Constantinidis, 1993; Martın Most important, however, are not the benefits et al., 2009; Trigas et al., 2008, respectively. Two that members of sect. Centaurea provide, but their species with divergent chromosome numbers (x = negative impacts on agriculture and landscape. Cen- 11), Centaurea exarata Boiss. ex Coss. and C. subti- taurea stoebe (= C. maculosa Lam.) and to a lesser lis Bertol., traditionally assigned to sect. Centaurea, extent C. diffusa and C. virgata subsp. squarrosa were found to be closely related to the Centaurea (Boiss.) Gugler, are tremendous invasive weeds in jacea complex (sect. Phrygia) and thereafter were pastures of the US and Canada. Centaurea stoebe,

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 8 A. HILPOLD, N. GARCIA-JACAS, R. VILATERSANA & A. SUSANNA

2 sp. 3 sp. 0.78 % 1.17 % 98 sp. 12 sp. 38.3 % 4.7 % No information

2n = 18 32 sp. 12.5 % 2n = 36

2n = 18 and 36 107 sp. 41.8 % 2n = 54

Divergent chromosome number

Figure 3. Pie diagram indicating the percentages of known - unknown chromosome numbers a, ploidy levels, and discording results in Centaurea sect. Centaurea.

“one of ’s most devastating invasive few centimetres in annual and biennial species up to ” (Blair & Hufbauer, 2010), diminishes the 1.50 m in both biennial and perennial species, but fodder value of pastures infesting about 3 million ha mostly between 20 and 50 cm). A third section was (Di Tomaso, 2000) producing in this way economic upgraded to sectional rank 30 years ago: under the damage of hundreds of million dollars every year. name of sect. Willkommia Blanca (Blanca, 1981a; The estimated damage for the agriculture of Mon- Fig. 1K) a group of species was defined which shows tana amounts to 42 million dollars (Duncan et al., perennial life form (frequently dwarf shrubs) and 2001). The fight against these weeds is also the mo- usually black, fimbriate bract appendages ending in tor of most of the research conducted in Centaurea a small spine. This group, contrarily to the other two (e.g. Hufbauer & Sforza, 2008; Collins et al., 2011; sections, was defined also geographically as includ- Pollock et al., 2011; Reinhart & Rinella, 2011). ing only individuals from NW Africa and the Iberian Peninsula, but leaving behind morphologically simi- Traditional, morphology-based treatments and lar forms from the Eastern Mediterranean. The de- their limitations limitation between these three sections is more than problematic. The distinction between sections Pha- Section Centaurea is traditionally divided into two lolepis and Centaurea is based on one single morpho- main groups based on the morphology of their bract logical character and is thereafter highly questiona- appendages: section Phalolepis (Cass.) DC. (Fig. 1J) ble if this single character shows intermediate forms with lacerate membranaceous appendages, and or is not present at all (as in some species from the sect. Centaurea (formerly sect. Acrolophus (Cass.) central Mediterranean, where the bract appendages DC.; Fig. 1I) with ciliate to fimbriate ones. These are totally or almost totally reduced). The presence two groups are treated as subgenera in Dostál (1976) of intermediate forms was demonstrated by Wagen- in his treatment for and more often itz (1989) and appears in many species descriptions as sections. The appendage morphology is the only (e.g. Breitwieser & Podlech, 1986; López & Devesa, character used for their delimitation (Dostál, 1976), 2008c). Intermediate forms have been commonly at- but within these two groups, however, a broad range tributed to hybridization between the two sections, of morphological characters can be found to per- without doubting their monophyly (Garcia-Jacas et form further delimitations (Fig. 1L–N): leaf shape al., 2006; Suárez-Santiago et al., 2007b). (divided), indumentum (tomentose, glabrous), flow- Delimitation and monophyly of section Willkom- er colors (rarely yellow, mostly purple), size (a mia is not clear. In the Eastern Mediterranean, C.

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 Taxonomical and nomenclatural notes on Centaurea 9 attica Nyman from shows similar bract ap- Systematic treatments of sect. Centaurea in light pendages, and similar species are also found in NE of molecular approaches Libya (C. cyrenaica Bég. & A. Vacc.) and on the easternmost shore of the Mediterranean, for exam- From the late 1990s onwards, a series of molecu- ple C. damascena Boiss. and C. dumulosa. Boiss. lar investigations in sect. Centaurea has been con- All these species are currently assigned to sect. Cen- ducted, using mainly DNA sequence data (Ochs- taurea. Intermediates between sections Willkommia mann, 2000; Garcia-Jacas et al., 2006; Wagenitz et and Centaurea are frequent on the Iberian Peninsula, al., 2006; Suárez-Santiago et al., 2007b; Beltrame, where most members of section Centaurea show 2007; Mráz et al., 2012). Additionally, some work somewhat fimbriate and at least slightly spiny bract has been done, considering small species groups and appendages. Placement of species like C. corduben- using microsatellites techniques (Marrs et al., 2006; sis and C. monticola in any of these two sections is Suárez-Santiago et al., 2007a), isozymes (Bancheva unclear, which can also be seen in incongruencies et al., 2006, 2011), RAPD (Tornadore et al., 2000; between taxonomic treatments (Blanca, 1981a vs. Sozen & Ozaydin, 2010) and SDS-PAGE (Uysal López & Devesa, 2008c). Delimitation between et al., 2010). Interestingly, the traditional sectional Willkommia and Phalolepis following sole morpho- division based on morphological traits was not con- logical observations is not precise either. Partly or firmed in any of these molecular works. entirely membranaceous bract appendages within Willkommia are found in NW Africa (C. debdouen- Species delimitation within sect. Centaurea sis Breitw. & Podlech, C. pomeliana Batt.) and in the Iberian Peninsula (C. avilae Pau). Centaurea As taxonomic treatments within sect. Centaurea are tougourensis Boiss. & Reut., one of the only two highly incongruent, a really crucial question arises: Phalolepis species from NW Africa, has a peren- what has to be considered as a species and what not? nial, subshrubby life form and a small spine on their From the answer of this question depend many other otherwise membranaceous bract appendages—very investigations, like the diagnosis of the conservation similar to members of section Willkommia, of which status and conservation strategies (e.g. Townsend- it is surrounded by. Peterson & Navarro-Sigüenza, 1999). Modern species Besides the separation of these three subgenera, concepts agree in the point that species are separately Dostál (1975, 1976) distinguished two subgroups evolving metapopulation lineages (De Queiroz, 2007), within his subgenus Phalolepis and 11 within Acrol- but members of the genus Centaurea, like most other ophus, stressing for their delimitation upon morphol- plant species as well, were and are still described fol- ogy of leaves, indumentum and bract appendages. lowing a purely phenetic species concept (Michener, Further groupings within the three sections are used 1970), i.e. only by assessing morphological characters in local treatments, for example in Pignatti (1982). in the hope that all populations bearing these charac- ters would correspond to a single lineage. This ap- Hybridisation proach itself may be prone to produce inaccurate re- sults, but even more difficult is the question how much The morphologically described species of sect. morphological divergence is enough to separate two Centaurea are not separated through intrinsic morphologically distinct plant groups as separate spe- breeding barriers. Reports about hybrids are abun- cies. Additionally, the same or similar characters may dant (e.g. Halácsy, 1902; Georgiadis, 1981a; Blan- arise separately and may therefore not be synapomor- ca, 1984). These hybrids are frequently fertile and phies but rather homoplasic traits. Differences in spe- homoploid (Blanca, 1981a; Ochsmann, 1998; Pis- cies numbers produced by different modi operandi are anu et al., 2011), sometimes also polyploid (Blair huge. Examples can be easily seen if we compare dif- & Hufbauer, 2010; Mráz et al., 2012). Hybrids with ferent taxonomic treatments in Centaurea. not closely related species—for example from the WMC—have been reported (Pau, 1914; Prodan, Species List 1930; A. Susanna, pers. obs.), these however, pro- duce usually no fertile offspring and therefore they The following list includes all species of the newly do not lead to reticulation. defined section Centaurea (including former sections

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 10 A. HILPOLD, N. GARCIA-JACAS, R. VILATERSANA & A. SUSANNA

Acrolophus, Phalolepis and Willkommia). Only re- Italy, and central Europe — All Ital- cent literature is incorporated. Taxonomy follows ian Centaurea species are listed in Conti et al. Euro+Med Plantbase (Euro+Med, 2006–2013), which (2005) and in Network Nazionale della Biodiver- is the most complete listing of the genus Centaurea. sità (2012–) but are not treated in detail. Complete For areas not included in Euro+Med, local floras were treatments for Italy are given in Fiori (1927) and in used. Cases where our treatment diverges from that of Pignatti (1982). Treatments for single groups: Ar- Euro+Med are specifically mentioned. Synonyms are rigoni (2003; C. paniculata complex), Cela Ren- only given if the name in Euro+Med diverges from the zoni & Viegi (1982; C. cineraria complex); Gua- name used in this list. The second column of the Table rino & Rampone (2006; C. dissectae group). Many 2 shows the currently accepted assignation to any of new descriptions have been published over the last the three subsections Centaurea (C, formerly sect. Ac- 30 years. In the meantime, C. subtilis has been re- rolophus), Phalolepis (P) and Willkommia (W). The moved form the section (Hilpold et al., 2009). Here third column shows the approximate distribution of we follow the very tight treatment of Euro+Med. the . The fourth column gives information about Balkans (including , Aegean Sea and the chromosome number. We usually give two cita- ) — The most complete taxonomic treatments tions per chromosome number, because further infor- concentrating on the Balkan area are those of Boissier mation can be easily retrieved from the indices to plant (1875) and Hayek (1931). Newer treatments of the chromosome numbers (Goldblatt & Johnson, 1979–; entire area other than Flora Europaea (Dostál, 1976) Watanabe, 2002). The fifth column shows if there ex- do not exist. In the NW Balkans, besides some widely ists evidence from the ITS that the taxon belongs to distributed species, there are two main species groups: the section Centaurea. The information derives, if not the C. spinosociliata group and the C. cuspidata otherwise mentioned, from the studies of Ochsmann group. The latter one, though only distributed in a rela- (2000), Garcia-Jacas et al. (2006), Suárez-Santiago et tively small area, was split into several microspecies of al. (2007a, b) and Hilpold et al. (2014). Column six doubtful rank. These two species groups are treated in lists revisions of the species when available. The last Lovrić (1968). A treatment for Romania is given in Ci- column gives information about the existence of sub- ocârlan (2000), for Greece in Halácsy, (1902), for the species (only those accepted in Euro+Med). Greek Mountains in Gamal-Eldin & Wagenitz (1991), Iberian Peninsula —Systematic treatment follow for the Aegean Sea in Rechinger (1943), for N Greece the preparatory works for Flora iberica by López in Rechinger (1939). Listings of sect. Phalolepis from & Devesa (2008a, b, c, 2010, 2011) and López et Greece are given in Wagenitz (1971) and Georgiadis et al. (2011) for the C. paniculata and the C. alba al. (1996), of the Bulgarian members of genus Centau- groups, and Blanca & Suárez-Santiago (2011) for rea in Assyov et al. (2002) of the Croatian members in sect. Willkommia. Identification keys can be found in Flora Croatica Database (Nikolić, 2010). López & Devesa (2010; C. paniculata complex & C. Region N of Black Sea (Russia p. p., , diffusa), López & Devesa (2011; C. alba complex) Moldavia) — Most species in this area belong to and Blanca (1981a) for members of sect. Willkom- five species groups: the C. stoebe complex, the C. mia. Whether C. paniculata and C. leucophaea Jord. margaritacea aggr., the C. sterilis aggr., the C. ovi- from the Iberian Peninsula are really closely related na aggr. and the C. arenaria aggr. Monophyly of or even homonymous to populations from SE France these groups is not clear. The most complete treat- and NE Italy is an open question. The same is true ment for Russia, Ukraine and Moldavia is given in for Centaurea boissieri DC., C. monticola Boiss. ex Klokov et al. (1963). DC. and C. resupinata Coss., all listed for the Iberian Anatolia and Cyprus — Flora of Turkey (Wagen- Peninsula and NW Africa. itz, 1975) lists nine species for section Phalolepis North Africa — Most members of sect. Centaurea and 21 species for section Centaurea (= Acrolo- occur in the NW of the continent, namely in various phus). In recent years, several new species have ranges of the Atlas Mountains. Only C. cyrenaica been described. Two species of Phalolepis (C. Bég. & A. Vacc. occurs in the NE of the continent hierapolitana and C. tossiensis) do not belong to (Lybia). No recent complete treatments of the group sect. Centaurea. The same is true for C. akamantis are available, only the revision of section Willkommia from Cyprus—where only one species remains, C. from NW Africa by Breitwieser & Podlech (1986). cyprensis (Holub) T. Georgiadis (Meikle, 1985).

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 Taxonomical and nomenclatural notes on Centaurea 11

Caucasus Region — The Caucasus region, in- Centaurea sect. Cnicus (L.) Garcia-Jacas, Hilpold, cluding parts of Russia, , Armenia and Az- Susanna & Vilatersana, new status and combina- erbaijan, is relatively poor in endemic taxa of sect. tion ≡ Cnicus L., Sp. Pl. 2: 826. 1753 (nom. cons.) Centaurea. The most complete treatment for the [Basionym]. Type and only species: Centaurea area is given in Klokov et al. (1963). benedicta L., Sp. Pl. 2: 826. 1753. Fig. 1D. Near-East, Middle-East — No complete list- ings or treatments are available postdating Boissier Centaurea sect. Hierapolitanae Garcia-Jacas, (1875), since Euro+Med considers only the west- Hilpold, Susanna & Vilatersana, new section ern part of this area. Contributions for this area in Annual or perennial species with heterogamous the list derive mainly from regional floras; i.e. Post heads. Capitula minute, cylindrical, with only & Dinsmore (1932), Feinbrun-Dothan (1978) and 10-20 florets, the outer ones sterile and radiant. Wagenitz (1980). Involucral with orbicular, scariose, lacer- ate appendages, similar to those of sect. Pha- Nomenclatural proposals lolepis, but long decurrent. Type: Centaurea hierapolitana Boiss., Diagn. Pl. Orient. ser. 1, Centaurea subgenus Lopholoma (Cass.) Dobrocz. 4: 15. 1844. Other species: Centaurea tossien- Sect. Acrocentron (Cass.) DC. subsect. Chamae- sis Freyn & Sint., Oest. Bot. Zeitschr. 44: 258. cyanus (Willk.) Garcia-Jacas, Hilpold, Susanna & 1894. Fig. 1F. Vilatersana, new combination ≡ Centaurea sect. Chamaecyanus Willk. in Willkomm & Lange, Centaurea sect. Phrygia Prodr. Fl. Hisp. 2: 150. 1870 [Basionym]. Centaurea sect. Phrygia subsect. Exarata, new subsection Centaurea subgenus Centaurea Stands apart from subsect. Phrygia and subsect. Sect. Centaurea Jacea by the reduced appendages of the bracts, Centaurea sect. Centaurea subsect. Phalolepis which are very small, applicate, not plumose (Cass.) Garcia-Jacas, Hilpold, Susanna & Vilat- and recurved as in subsect. Phrygia or very ersana, new combination ≡ Phalolepis Cass., wide and covering partly or totally the bracts as Dict. Sci. Nat., ed. 2., 50: 248. 1827 [Basionym] in subsect. Jacea. Differs from subsect. Subtilis ≡ Centaurea sect. Phalolepis (Cass.) DC., Prodr. by having entire leaves, not pinnatisect as in C. 6: 568. 1838. ≡ Centaurea subgenus Phalolepis subtilis. Type and only species: Centaurea ex- (Cass.) Dobrocz., Bot. Zurn. 6: 63. 1949. Fig. 1K. arata Boiss. ex Coss., Notes pl. crit.: 116. 1851. Fig.1H. Centaurea sect. Centaurea subsect. Willkom- mia (Blanca) Garcia-Jacas, Hilpold, Susanna Centaurea sect. Phrygia subsect. Jacea (L.) & Vilatersana, new combination ≡ Centaurea Garcia-Jacas, Hilpold, Susanna & Vilatersana, sect. Willkommia Blanca, Lagascalia 10: 138. new combination ≡ Centaurea [infrageneric 1981 (publ. 1982) [Basionym]. Fig. 1L. unranked] jacea L., Sp. Pl. 2: 909. 1753 [Basio- nym] ≡ Centaurea sect. Jacea (L.) DC., Prodr. Centaurea sect. Akamantis Garcia-Jacas, Hilpold, 6: 570. 1837 [publ. 1838]. Susanna & Vilatersana, new section Perennial species with heterogamous heads. Cauline Centaurea sect. Phrygia subsect. Subtilis, new leaves linear-spatulathe, sparsely lanuginose. Capit- subsection ula very small, exactly ovate. Bracts lanuginose; ap- Can be easily set apart from subsect. Phrygia, pendages much reduced, triangular, scariose, termi- subsect. Exarata and subsect. Jacea by the pin- nated in a short recurved mucro. All the florets deep natisect basal and mid-cauline leaves, which purple, the central ones with darker anther-tube, the are entire in all the other subsections. Differs outer ones sterile, only slightly radiant. from sects. Jacea and Phrygia by the small and Type and only species: Centaurea akamantis T. triangular bract appendages not covering the Georgiadis & Hadjik., Willdenowia 23: 157. bracts. Type and only species: Centaurea subti- 1993. Fig. 1E. lis Bertol., Flora Italica 9: 451. 1854.

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 12 A. HILPOLD, N. GARCIA-JACAS, R. VILATERSANA & A. SUSANNA 6 2 2 3 3 10 Subsp. et et a I TS sequences , 2013 , 2011 , 2011 , 2001 , Willkommia all of them considered Anzalone, 1995; Hilpold al. , 2011 D uran & uman, 2002 Guarino & R ampone, 2006; Hilpold et al. Atasagun et al. unemark, 1967; Turland & Turland R unemark, 1967; C hilton, 2000 Arrigoni, 2003; Hilpold al. , 2011 Brullo et al. – – Phitos & D amboldt, 1971 – – + + + + + + López & D evesa, 2011 + + + + + + + + + + + + López & D evesa, 2008 + Armağan. & Ünal, 2009 + ITS Literature Other and sect. et al. , Phalolepis d , 2011) d ) et al. , 2007a), further taxonomic literature, and number of . Ten. var. var. Ten. , 2001 as C. dissecta et al. , 1998) a ) Acrolophus Acrolophus = Centaurea, sect. 18, 36 (Georgiadis, 1983) 18, 36 (Georgiadis, Chromosome number (2n) number Chromosome – 1982) Viegi, et al. , 1972; C ela R enzoni & 18 (Viegi & C ela R enzoni, 1976) 18 (Viegi Boiss. & Heldr. subsp. 36 (Matthäs, 1976 as C. subciliaris Boiss. & Heldr. acarnanica Matthäs) 36 ( U ysal et al. , 2009) – 18 (Köse, 2006) – – 18 ( U ysal et al. , 2009) 18 (Georgiadis, 1983; Montmollin, 1986) 18 (Georgiadis, – 2006; Kuzmanov 32, 36 (Bancheva & Greilhuber, 1979) enzoni, 1976; Arrigoni et al. , 1980) & C ela R enzoni, 1976; 18 (Viegi 18 (Georgiadis & Phitos, 1976; Georgiadis, 1983); 36 & Phitos, 1976; Georgiadis, 18 (Georgiadis & Phitos, 1976) (Georgiadis – 18 (Signorini et al. intermedia Fiori) – 18 (Blanca, 1980 18 (Garcia–Jacas . For each species we indicate sectional adscription, geographical distribution, chromosome number, Centaurea (including former sections Willkommia Anatolia, C aucasus 18 (Bakhshi Khaniki, 1996) Balkans Italy Italy I berian Peninsula 18 (López & D evesa, 2008 Distribution N Black Sea Greece Anatolia Anatolia Anatolia Italy Balkans Anatolia I berian Peninsula 36 (López & D evesa, 2008 Greece N Black Sea NE Balkans, N and E Black Sea Italy Greece, W Balkans W Greece, Anatolia Italy Italy I berian Peninsula Anatolia, C aucasus, Iran P P P P P P P P P C C C C C C C C C C C C C C W Subsect. ; W: W: Phalolepis ; ( C. , 2011; and Suárez-Santiago et al. , 2000, 2001, 2006; Hilpold 2011; C. ovina aggr.) Hub.-Mor. ; P: Centaurea ( Hub.-Mor. C : Klokov ( C. uman & A. H. D uman & Halácsy Moretti List of species of the redefined sect. (Sommier) Bég. Lojac. Pau Nyman subsections). yes/no (from Garcia-Jacas Table 2. Table Species C. aggregata D C. C. aggregata C. affinis Friv. C. aeolica C. aetaliae C. alba L. C. aemulans Klokov ( diffusa s. l. ) C. acarnanica (Matthäs) Greuter C. albanica C. aplolepa C. aphrodisea Boiss . C. aphrodisea C. anthemifolia C. antalyensis D uran C. ambigua Guss. C. amaena Boiss. & Balansa Hoffmanns. & Link C. aristata Hoffmanns. L. C. argentea C. appendicata aggr.) margaritacea Willd. C. arenaria C. avilae C. attica C. austroanatolica C. aspromontana Brullo et al. C. aspromontana deusta s. l. ) C. arrigonii Greuter C. aziziana R ech. f.

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 Taxonomical and nomenclatural notes on Centaurea 13 5 2 3 2 2 7 Ib, 1 NA Subsp. c a , 2011 , 2011 Köse & Alan, 2013 Köse & D avis et al. , 1988 Hilpold et al. Phitos & Constantinidis, 1993 Blanca & Suárez-Santiago, 2011 Hilpold et al. C aruso et al. , 2013 Brullo, 1988 – – – – – – – – – – + 1972 Wagenitz, + 1972 Wagenitz, + + + López & D evesa, 2008 + + + López & D evesa, 2008 + + + + + + + + ITS Literature Other et al. , 2009) d ) d ) a ) b ) a , 1981 b ) , 2009); 36 (Georgiadis & , 2009); 36 (Georgiadis Chromosome number (2n) number Chromosome 18 (Martın et al. Christodoulakis, 1984; Martın – 2006, as C . ovina Pall. ex 18 (Bancheva & Greilhuber, subsp. besserana ( D C.) ostál Willd. – et al. , 2008) 54 (Trigas – – 36 (Lovri ć , 1982 a ) 18 (Blanca, 1980 18, 36 (Cela-Renzoni & Viegi, 1982; D amboldt & Viegi, 18, 36 (Cela-Renzoni & Matthäs, 1975) 54 (Phitos & Constantinidis, 1993) 18 ( C iocârlan, 2000, secondary citation!) 18 ( R omaschenko et al. , 2004) 1983) 1974; Georgiadis, 18 ( D amboldt & Melzheimer, 18 (Geogiadis & Phitos, 1977) – – 18 ( U ysal et al. , 2009) – enzoni & Viegi, 1982) Viegi, et al. , 1972; C ela- R enzoni & 18 (Viegi – – Anatolia I berian Peninsula 18+ 0–1B (Blanca, 1980 Distribution Anatolia NE Balkans, N Black Sea I berian Peninsula 18 (López & D evesa, 2008 N Black Sea Greece Anatolia C aucasus I berian Peninsula 18 (López & D evesa, 2008 W Balkans W I berian Peninsula 18 + 2B (Blanca, 1981 I berian Peninsula, NW- Africa Greece Italy NE Balkans Anatolia Greece Greece N Balkans, Black Sea Greece, NE Balkans 1983) 18 (Georgiadis, Anatolia Italy Italy Italy N Black Sea P P P P P P P BalkansW Greece, & Phitos, 1977) 18 (Georgiadis P P C C C C C C C C C C C C C C C W W W Subsect. C. ovina aggr.) ( Talavera C. margaritacea ( C. margaritacea Hayek Iljin Prodan Boiss. Table 2. ( C ont.) Table Species C. cariensis Boiss. C. carratracensis Lange Köse & Alan C. baseri Köse & C. besseriana D C . ( ovina aggr.) C. bethurica E. López & D evesa C. caprina Steven & C. carystea Trigas C onstantin. C. cariensiformis Hub.-Mor. C. caspia Grossh. C. castellanoides Teyber ( C. cuspidata C. biokovensis Teyber aggr.) C. bombycina D C. boissieri D Phitos & Phitos & C. cithaeronea C onstantin. C. cineraria L. C. caliacrae C. calolepis C. chalcidicaea C. chrysocephala Phitos & T. Georgiadis C. arenaria C. borysthenica Gruner ( arenaria aggr.) C. bovina Velen. C. breviceps C. cadmea Boiss . C. calabra G. C aruso, S.A.Giardina, R aimondo & Spadaro C. brunnea (Halácsy) Halácsy Guss. C. busambarensis C. brulla Greuter ( deusta s. l. ) aggr.)

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 14 A. HILPOLD, N. GARCIA-JACAS, R. VILATERSANA & A. SUSANNA 2 Subsp. c , 2011 , 2011 , 2011 Hilpold et al. Hilpold et al. Guarino & R ampone, 2006 Alavi, 1983 Valsecchi & Filigheddu, 1991 Valsecchi Meikle, 1985 as C. veneris (Sommier) Bég. – – – – – – – – + + + López & D evesa, 2008 + López & D evesa, 2010 + + Phitos & D amboldt, 1971 + + + + + + + + + + López & D evesa, 2011 + ITS Literature Other nigra D C. subsp. et al. , 1991) Vis. subsp. C. incompta Vis. d ) d ) a ) , 1979), 36 (Georgiadis & Phitos, , 1979), 36 (Georgiadis , in press) Chromosome number (2n) number Chromosome – – 36 (Hilpold et al. 18 (Georgiadis & Phitos, 1976; Taylor & Taylor, 1977); Taylor, & Taylor & Phitos, 1976; 18 (Georgiadis 2006) 16 (Bancheva & Greilhuber, – 16 (Strid, 1983) 18 (Baltisberger, 1990 as C. ambigua Guss. subsp. 18 (Baltisberger, – 18 (Strid & Franzen, 1981 as & Panči ć ) D ostál) derventana (Vis. 18 + 0–1B (Matthäs, 1976; Brullo (Fiori) Pignatti & Lausi – – 18 (Brullo & Pavone, 1977) – – 18 (Kuzmanov et al. 1983) 1976; Georgiadis, 18 (Lovri ć , 1982 a ) 1983 as C. biebersteinii 36 (Georgiadis, (Bornm.) D ostál) cylindrocephala – – 1980) 36 (Siljak–Yakovlev, chsmann, 1999; A. Hilpold unpubl. 36 (Lausi, 1966; O chsmann, 1999; res.) Distribution I berian Peninsula 36 + 1B (Blanca, 1980 I berian Peninsula 36 (López & D evesa, 2008 Anatolia NE Balkans Italy NE Mediterranean and NE Mediterranean and Black Sea, introduced Europe, W to C and America, N and S Anatolia Italy Balkans W Italy N Black Sea Near-East Libya I berian Peninsula 18 (López & D evesa, 2008 France W Balkans W Balkans W Greece, C yprus Balkans, Anatolia Balkans, NE Balkans Balkans W taly, W Balkans W I taly, P P Balkans W Greece, P P P C C C C C C C C C C C C C C C C C W Subsect. C. C. spinosociliata C. alba p. p .) Vis. Breitw. & PodlechBreitw. WAfrica NW Sm. Font Quer C . Guarino & R ampone Table 2. ( C ont.) Table Species C. citricolor C. cordubensis Font Quer C. cordubensis C. consanguinea D C. codruensis Prodan ( kanitziana s. l. ) Vals. & Filigh. Vals. C. corensis C. diffusa Lam. Boiss. & Heldr. C. dichroa C. deustiformis Adamovi ć cuspidata aggr.) C. delucae & Panči ć ( C. derventana Vis. Ten. C. deusta C. demetrii D umbadze ( ovina aggr.) C. damascena Boiss. C. delicatula Bég. & A. Vacc. A. Bég. & C. cyrenaica & PodlechC. debdouensis Breitw. WAfrica NW Willk. ( Willk. C. costae C. corymbosa Pourr. Vis. ( C. cuspidata C. cuspidata Vis. aggr.) Bornm. ( C. C. cylindrocephala stoebe s. l. ) Georgiadis T. (Holub) C. cyprensis C. cuneifolia Nyár. ( C. stoebe s. l. ) C. coziensis Nyár. C. crithmifolia C. cristata Bartl. ( aggr.)

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 Taxonomical and nomenclatural notes on Centaurea 15 2 4 2 2 Subsp. C. Batt. , 2011 , 2011 mauritanica et al. , 2011 L. var. var. L. Gamal-Eldin & Wagenitz, Wagenitz, Gamal-Eldin & 1983 Micevski, 1985 Bancheva Arrigoni, 1972 Hilpold et al. Quézel & Santa, 1963 as alba Hilpold et al. – – – – – – – – – 1980 Wagenitz, – 1980 Wagenitz, – – – – – – + + + + + + + + + ITS Literature Other

, 2012) et al. , 2005) C. sagredoi Blanca subsp. tenuiloba b as C. sagredoi 36 (Siljak-Yakovlev, 1980; Siljak-Yakovlev et al. , 1980; Siljak-Yakovlev 36 (Siljak-Yakovlev, 2005) 18 (Papes & Radi ć , 1982; Siljak-Yakovlev – – – 1983) 36 (Strid & Franzen, 1981; Georgiadis, – 18 ( R aimondo & Spadaro, 2006) (Boiss.) Blanca – Arrigoni & Mori, 1971) 18 (both subsp.; – – 1980) 36 (Siljak-Yakovlev, – 18 (Blanca, 1981 18 ( R aimondo & Bancheva, 2004) 36 (Papes & Radi ć , 1982) – 18 ( U ysal, 2008) – – 36 ( U ysal & Köse, 2009) 36, 18 ( R omaschenko et al. , 2004; Mráz Chromosome number (2n) number Chromosome – ’Amato & Pavesi, 1990; A. Hilpold unpubl. res.)18 ( D ’Amato & Pavesi, 1990; + W Balkans W Balkans W NE Balkans Greece Anatolia, C aucasus 1980) 18 (Tonian, N Black Sea Balkans W Greece, Italy W Balkans W NE Balkans Italy I raq Balkans W I raq I berian Peninsula Italy W Balkans W N Balkans Anatolia N Black Sea E Mediterranean (Middle East) Anatolia N Black Sea Distribution Italy P P P PP Balkans W P P P P PAfrica NW P C C C C C C C C C C C C C C W Subsect. C. cuspidata D umbadze ( C. (Nyman) Heldr. Velen. Blakelock Micevski Vis. Table 2. ( C ont.) Table C. grisebachii C. glaberrima Tausch C. gloriosa Radi ć ( aggr.) C. gracilenta E. Gamal-Eldin & C. greuteri Wagenitz C. gulissashvilii ovina aggr.) C. margaritacea C. gerberi Steven ( margaritacea aggr.) R aimondo & Spadaro C. giardinae C. galicicae C. euxina Velen. Viv. C. filiformis C. formanekii Halácsy C. foveolata C. friderici C. fusiformis Blakelock Blanca C. gadorensis C. erycina R aimondo & Bancheva C. epapposa Velen. C. ertugruliana U ysal aggr.) aggr.) C. dumulosa Boiss. C. dursunbeyensis U ysal & Köse C. edith-mariae Radi ć ( cuspidata C. donetzica Klokov ( aggr.) margaritacea C. margaritacea C. dubjanskyi Iljin ( margaritacea Species C. djebel-amouri Greuter C. diomedea Gasp.

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 16 A. HILPOLD, N. GARCIA-JACAS, R. VILATERSANA & A. SUSANNA 2 6 Subsp. et ; C atoni , 2011 , 2001 enzoni & Viegi, 1982; Viegi, C ela- R enzoni & Hilpold et al. & Gratani, 2013 Brullo et al. Kalpoutzakis & C onstantinidis, 2004 Francini, 1951; Tornadore et Tornadore Francini, 1951; al. , 2000 2003 Tan, Strid & Lovri ć , 1971; Marcucci al. , 1999 Aidin et al. , 2013 – – 1980 Wagenitz, – – – – – – – – – – – + + + + + + + + López & D evesa, 2010 + + + ITS Literature Other et al. , d ) , 2000) a ) , 2010) Chromosome number (2n) number Chromosome 18 (López & D evesa, 2008 18 (Viegi et al. , 1972) 18 (Viegi 36 (Lovri ć , 1982 b ) 18 (Bakhshi Khaniki, 1996) – 18 (Strid & Franzen, 1981) – – 18 (Blanca, 1980 amboldt & Melzheimer, 1974; Matthäs, 1976)18, 36 ( D amboldt & Melzheimer, & C ela R enzoni, 1976) 18 (Viegi – – 18 ( D esole, 1954) 18 (Phitos & D amboldt, 1971) – 18 (Tornadore et al. 18 (Tornadore – – 18 (Georgiadis & Phitos, 1976) 18 (Georgiadis – 1982) 18 (Lausi, 1966; Siljak-Yakovlev, – – 36 + 1–5B (Papes & Radi ć , 1982; Siljak-Yakovlev 2005) Distribution NW-Mediterranean NW-Mediterranean (Spain, France) Italy W Balkans W I ran, raq Anatolia, NE Balkans Italy I berian Peninsula Greece Italy Greece Italy C aucasus Greece N Balkans Italy Greece NE Balkans Greece NE Balkans Balkan W I taly, Anatolia, NE Balkans 36 (Meriç et al. N Black Sea N Black Sea Balkans W P Balkans W P P P P P C C C C C C C C C C C C C C C C C C Subsect. C. cuspidata Steven ( C. ovina Boiss. Halácsy Halácsy D . Brândză ) Boiss. (Lacaita) Brullo s. l. ) J. Wagner J. (Sommier) Arrigoni ( C. (Sommier) Boiss. Table 2. ( C ont.) Table C. stoebe Species C. hanrii Jord . C. gymnocarpa Moris & D e Not. C. ipecensis R ech. f. Vis. ( C. cuspidata C. incompta Vis. aggr.) C. intricata C. inermis Velen. C. ionica Brullo ( deusta s. l. ) C. jaennensis D egen & ebeaux W C. incompleta C. ilvensis aetaliae s. l. C. huljakii C. horrida Badarò C. hohenackeri aggr.) C. heldreichii C. heldreichii C. arenaria C. jankana Simonk. ( arenaria aggr.) C. japygica Strid & Kit Tan C. johnseniana Strid & Kit C. jurineifolia C. kalambakensis Freyn & Sint. C. kanitziana C. kartschiana Scop. C. kilaea C. margaritacea C. konkae Klokov ( margaritacea aggr.) C. kubanica Klokov ( C. kusanii Radi ć ( aggr.)

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 Taxonomical and nomenclatural notes on Centaurea 17 6 6 2 Subsp. et al. , b et al. , 2011 , 1996 Arrigoni & C amarda, 2003 Georgiadis et al. Georgiadis U ysal et al. , 2010 Arrigoni, 2003; López & D evesa, 2010; Hilpold 2011 Francini, 1951; Tornadore et Tornadore Francini, 1951; al. , 2000; Hilpold – – – – – – – – – – + + + + + + + + + + + & Phitos, 1978 Georgiadis + + López & D evesa, 2008 + + ITS Literature Other d ) et al. , 2005) et al. , 2011) a , 1983) d ; López d ) , 1996) , 2000) – – 36 (Papes & Radi ć , 1982; Siljak-Yakovlev et al. 18 (Georgiadis – – – 18 ( U ysal et al. , 2009) Chromosome number (2n) number Chromosome 36 (Georgiadis, 1983) 36 (Georgiadis, – 18 (Tornadore et al. 18 (Tornadore – 18 (Georgiadis & Phitos, 1976) 18 (Georgiadis 1981 a ) & Phitos, 1976; Georgiadis, 36 (Georgiadis – 18 (Kalpoutzakis & C onstantinidis, 2004) 18 ( O chsmann, 1999; López & D evesa, 2008 18, 18+1B, 19, 20 (Blanca, 1980 18 ( U ysal et al. , 2009; Martın 2009) 18 ( U ysal et al. , 2009) 16 (Strid & Franzen, 1981) – Teyber subsp . 36 (Lovri ć , 1982 a as C. biokovensis Teyber Hayek) mucuriensis (Teyber) W Balkans W Greece N Black Sea N Black Sea N Black Sea Italy Anatolia Distribution Greece W Balkans W Italy Greece I berian PeninsulaGreece N Black Sea 18 (López & D evesa, 2008 Greece Mediterranean NW (Spain, France, Italy) I berian Peninsula 18 (López & D evesa, 2008 berian Peninsula, NW I berian Peninsula, NW Africa Anatolia Anatolia Greece Italy W Balkans W P Balkans W P P P P P Balkans W P P P C C C C C C C C C C C C C C W Subsect. C. Hayek

Jord. Bornm. Halácsy D C. Boiss. & Heldr. Halácsy ) Lacaita Kalpoutz. & C onstantin. (Fiori) Arrigoni ( C. (Fiori) Boiss. Table 2. ( C ont.) Table C. cuspidata aggr.) Species C. marmorea Bornm. & Soška C. marmorea C. mayeri Radi ć ( Georgiadis C. messenicolasiana T. et al. C. margaritalba Klokov ( C. C. margaritalba aggr.) margaritacea . ( C. Ten C. margaritacea aggr.) margaritacea D umbadze ( C . majorovii aggr.) arenaria C. magistrorum Arrigoni & C. magistrorum C amarda C. lycia C. lacerata (Hausskn.) Halácsy C. lactiflora C. monticola C. leucomalla C. leucomelaena C. leucadea C. langei Nyman C. laureotica Klokov ( C. ovina C. lavrenkoana aggr.) C. leonidia C. leucophaea & Link C. limbata Hoffmanns. C. lycaonica C. luschaniana Heimerl ex Stapf T. Georgiadis & Phitos Georgiadis T. C. litochorea P C. litigiosa aetaliae s. l. Teyber ( C. cuspidata Teyber C. mucurensis aggr.)

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 18 A. HILPOLD, N. GARCIA-JACAS, R. VILATERSANA & A. SUSANNA 5 Subsp. et al. , , 2000 , 2011 , 2011 , 2001 , 2011; et al. , 2011; Kalpoutzakis & C onstantinidis, 2004 onstantinidis & Vassiliades, Vassiliades, C onstantinidis & 1996 Georgiadis, 1979 Georgiadis, Guarino & R ampone, 2006; Bancheva Hilpold et al. Quézel & Santa, 1963; 1981 as C. Pottier-Alapetite, cineraria L.; Hilpold 2011 D avis et al. , 1988 Arrigoni, 2003; López & D evesa, 2010 Hilpold et al. Tornadore et al. Tornadore Talavera & Muñoz, 1984 Talavera D amboldt & Matthäs, 1979; Phitos & D amboldt, 1976 Brullo et al. – – – – – – – Geogiadis & Phitos, 1977 + 1972 Wagenitz, + + + + + + + + + + + + + ITS Literature Other O chsmann, , 2000) d ) a ) , 1991; Tornadore et al. Tornadore , 1991; 18 (Georgiadis & Phitos, 1976) 18 (Georgiadis 18 (López & D evesa, 2008 – – – 1983) 18 (Georgiadis, 1980) Tonian, 18 (Bakhshi Khaniki, 1996; – 18 (Brullo et al. 18 (Georgiadis & Phitos, 1976) 18 (Georgiadis Chromosome number (2n) number Chromosome 1996) Vassiliades, 18 ( C onstantinidis & 18 (Georgiadis, 1979) 18 (Georgiadis, 18 (Blanca, 1980 olombo & Trapani, 1990) Trapani, 18 ( C olombo & – – 1982) Viegi, et al. , 1972; C ela R enzoni & 18 (Viegi 18 (Geogiadis & Phitos, 1977) amboldt & Matthäs, 1975; Matthäs, & ( D amboldt +1B 36 1B, + 18 2000) 1983) & Phitos, 1976; Georgiadis, 18 (Georgiadis – + Greece Mediterranean NW (Spain, France, Italy) N Black Sea Anatolia Greece C aucasus N Black Sea Italy Greece Distribution Greece Greece I berian Peninsula Italy NW Africa NW Anatolia Italy I berian Peninsula – Greece Greece Italy P P P P P C C C C C C C C C C C C C WAfrica NW W W Subsect. ( C. Brullo et al. Phitos & D amboldt P Balkans W Greece, . ( C. ovina aggr.) ) L. Boiss. & O rph. s. Pomel ( C. resupinata Table 2. ( C ont.) Table Species C. orphanidea Boiss. C. paniculata C. arenaria C . odessana Prodan ( C. arenaria aggr.) C. olivieri l. ) C. olympica ( D C.) K. Koch cuneifolia s. l. C. ossaea Halácsy C. ovina Willd C. paczoskii Klokov ( aggr.) margaritacea C. nobilis (H. Groves) Brullo C. niederi Heldr. C. musarum T. Georgiadis C. musakii T. C. paui Willk. C. parlatoris Heldr. C. papposa ( C oss.) Greuter C. nydeggeri Hub.-Mor. C. panormitana Lojac. Talavera & J. M. Talavera C. pauneroi Muñoz C. pawlowskii C. paxorum Phitos & Georgiadis T. deusta s. l. ) C. pelia D C. pentadactyli

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 Taxonomical and nomenclatural notes on Centaurea 19 2 2 Subsp. , et b a ; Figuerola echinger, 1974; Tan et al. Tan 1974; R echinger, 2007 Blanca, 1980 U ysal et al. , 2010 Greuter, 1967 Greuter, U ysal et al. , 2010 Blanca & Suárez-Santiago, 2011 Blanca, 1981 al. , 1991 – Phitos & D amboldt, 1976 – – – – – – 1981 Wagenitz, – + + + 1971 Wagenitz, + + + + + + + + + + + Puntillo, 1996 + ITS Literature Other

et al. , D C.) – a as C. brachtii , 1996) et al. , 1997; Martın 2009) a ) a ) b as C. dufourii ( D ostál) Blanca + b ) 18 (Probatova et al. – – – 18 ( R omaschenko et al. , 2004) – Chromosome number (2n) number Chromosome 18, 36 ( O chsmann, 1999; Lovri ć , 1982 18 (Blanca, 1980 18 ( R omaschenko et al. , 2004) 18 (Georgiadis & Phitos, 1977) 18 (Georgiadis – – 18 (Georgiadis & Phitos, 1976) 18 (Georgiadis R chb. f.) 18 (Blanca, 1980 outsi & Georgiadis, 1988) 18 (Montmollin, 1986; R outsi & Georgiadis, – – 2005 as C. elegantissima Bornm.) , 1982; Siljak-Yakovlev 27+1B, 36 (Papes & Radi ć , 1982; Siljak-Yakovlev – 18 (Jasievicz & Mizienty, 1975, as C. calvescens 18 (Jasievicz & Mizienty, 18 (Blanca, 1981 18 (Garcia-Jacas – 18 (Blanca, 1981 – N Black Sea Greece Greece N Black Sea NE Balkans N Black Sea Distribution taly, W Balkans W I taly, I berian Peninsula N Black Sea Greece Greece Greece I berian Peninsula Greece N Black Sea Anatolia W Balkans W N Balkans I berian Peninsula, NW- Africa Aegean Sea Italy I berian Peninsula W Balkans W P P P P P P P P P C C C C C C C C C C C W WAfrica NW W W W Subsect.

C. cuspidata Kleopow ( C. Boiss. & O rph. C. margaritacea ( C. margaritacea D C. ) Iljin D C (Blanca) Blanca Puntillo C es. Plazibat ( Table 2. ( C ont.) Table C. pseudomaculosa D obrocz. ( stoebe s. l. ) Species C. pulvinata C. ptarmicoides Halácsy C. pseudoleucolepis margaritacea aggr.) margaritacea C. pseudocadmea Wagenitz C. pseudobovina Hayek ( jurineifolia s. l. C. protomargaritacea Klokov ( C. C. protomargaritacea aggr.) margaritacea C. pestalotii C. protogerberi Klokov ( C. C. protogerberi aggr.) margaritacea C. princeps Boiss. & Heldr. C. pomeliana Batt. R ech. f. C. prespana C. peucedanifolia C. pineticola C. pinnata Vicioso C. radichii C. poeltiana C. poculatoris Greuter aggr.) C. pinetorum Hub.-Mor. aggr.) Wagenitz C. reducta C. reichenbachii C. reichenbachii C oss. C. resupinata C. polyclada ( C. deusta s. l. ) C oincy C. rouyi C. rufidula Bornm.

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 20 A. HILPOLD, N. GARCIA-JACAS, R. VILATERSANA & A. SUSANNA 2 Subsp. et c , 2011 b , 2004 , 2001; Hilpold et al. , 2011 Manifold, partly published as C. maculosa R unemark, 1967 Brullo et al. Hilpold et al. D oğan & uran, 2009 R aimondo & Spadaro, 2008; Bancheva Bancheva & R aimondo, 2013 R aimondo et al. , 2004 Blanca, 1980 Brullo et al. al. , 2011 1 – – – – – – – – – – + + + + + + 1974 Wagenitz, + + + + + + + López & D evesa, 2008 + ITS Literature Other et al. , d ) et al. , 1997) , 1997) a ) et al. , 2006, secondary citation!) , 1999) 18 ( C onstantinidis 18+ 2B, 36 + 4B (Phitos & D amboldt, 1971; Matthäs, 1976) 2008) – Bor.; Španiel Bor.; 18, 36 (Lovri ć , 1982 b as C. rhenana 18 (Bal et al. – 18 ( R omaschenko et al. , 2004) – – & Phitos, 1976)36 (Runemark, 1967; Georgiadis 36 (Lovri ć , 1982 a ) + – – 18 ( R aimondo & Spadaro, 2008) – 18 (Anzalone, 1961) 18 (Blanca, 1980 36 ( R omaschenko et al. , 2004) 36 ( R omaschenko et al. , 2004) – 18 (Bancheva – Chromosome number (2n) number Chromosome amboldt & Melzheimer, 1974; Georgiadis, 1974; Georgiadis, 18+0–1B ( D amboldt & Melzheimer, 1983; Kuzmanov et al. , 1983) 18 (Boscaiu et al. Greece Greece, NE Balkans N Black Sea Europe, except I b. Pen., introduced to N and S America Anatolia Anatolia N Black Sea W Balkans W Aegean Sea N Black Sea Balkans W N Black Sea Anatolia Italy Italy Italy I berian Peninsula I berian Peninsula 36 (López & D evesa, 2008 N Black Sea N Black Sea E Balkans Italy Italy Distribution S and NE Balkans I berian Peninsula P P P P P P C C C C C C C C C C C C C C C C W W C/P Subsect. ( C. et al. Seenus A. D uran & B. Klokov ( C. ovina Sm. Brullo ( C. deusta s. l. ) Table 2. ( C ont.) Table C. subsericans Halácsy C. subciliaris Boiss. & Heldr. C. steveniana aggr.) C. stoebe L. Wagenitz C. sivasica Wagenitz Wagenitz C. sipylea Wagenitz C. sterilis Steven ( C. sterilis aggr.) C. soskae Košanin C. spinosa L. C. arenaria C. sophiae Klokov ( arenaria aggr.) C. spinosociliata C. savranica Klokov ( stoebe s. l. ) C. schousboei Lange C. sarfattiana Brullo et al. C. serpentinica D oğan C. sicana R aimondo & Spadaro deusta s. l. ) C. scannensis Anzal. et al. C. sagredoi Blanca C. sagredoi Species C. semijusta Juz. ( sterilis aggr.) N. B. Illar. ( C. C. sarandinakiae N. B. Illar. sterilis aggr.) Bancheva & C. sakarensis R aimondo C. saccensis R aimondo C. scillae C. rutifolia C. segariensis Figuerola et al.

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 Taxonomical and nomenclatural notes on Centaurea 21 2 2 3 2 Subsp. , 2006 et al. , 2006 et al. , 2006 C ela- R enzoni, 1970; Hilpold et al. , 2011 O chsmann, 2000 Wagenitz Wagenitz Guarino & R ampone, 2006 Micevski, 1985 Vural et al. Vural 3 . – – – – – – – – – – + 1972 Wagenitz, + + + + + Wagenitz + + + + + + + + + ITS Literature Other et al. , 2009) , 2012) et al. , 1999) , 2009) , 1991) et al. , 2010) – – & Chariat–Panahi, 1985; Martın 36 (Ghaffari 18, 36 + 6B (Siljak-Yakovlev et al. , 2005; Papes & Radi ć 18, 36 + 6B (Siljak-Yakovlev 1982) – 18 ( U ysal et al. , 2009) – Centaurea × psammogena 18 ( O chsmann, 1999 as Centaurea Gáyer) – 36, 18 ( R omaschenko et al. , 2004; Mráz – 1982) Viegi, et al. , 1972; C ela R enzoni & 18 (Viegi 18 ( O chsmann, 1999) – Chromosome number (2n) number Chromosome M. Bieb. ex Willd. Willd. M. Bieb. ex 18 [Sz.–Borsos, 1971 as C. arenaria (Gugler) Soó] subsp. pseudorhenana 18 (Brullo et al. 18 (Georgiadis, 1981 b as C. ipsaria Stoj. & Kitan.) 18 (Georgiadis, – – 54 (Martın et al. 18 (Benamara 18, 36 (Peruzzi & Cesca, 2002) 1983) & Phitos, 1976; Georgiadis, 36 (Georgiadis – – – 18 (Georgiadis, 1983) 18 (Georgiadis, Greece Asia, W E Europe and WC Europe and alien in America N W Balkans W Anatolia Anatolia Greece N Balkans, introduced elsewhere N Black Sea NE Balkans Italy NE Balkans , Italy Anatolia Distribution N Balkans Italy Greece Balkans W C aucasus Anatolia Greece Balkans NW I taly, 36 (Lovri ć , 1982 ­­­­­­­­ ; Marcucci Balkans Italy Italy Balkans 2 P P P P PP Africa NW P C C C C C C C C C C C C C C C C C C WAfrica NW C ? Subsect. et al. ( C. stoebe et al. Boiss. ( C. aggr.) Hub.-Mor. D egen & örfl. Heuff. ( D C.) Jord. Klokov ( C. sterilis Radi ć ( C. cuspidata D egen et al. Table 2. ( C ont.) Table C. vermia R ech. f. C. vesceritensis Lam. C. virgata s. l. ) resupinata C. visianii aggr.) Wagenitz C. werneri Wagenitz C. wagenitzii C. wettsteinii C. varnensis Velen. C. vankovii aggr.) C. vatevii C. veneris (Sommier) Bég. s. l. ) C. vandasii Velen. C. valesiaca Species C. arenaria A. Kern. ( C. arenaria C. tauscheri aggr.) C. tenacissima (H. Groves) Brullo C. thasia Hayek Micevski C. tomorosii Boiss. & R eut. C. tougourensis C. transcaucasica Grossh . C. tuzgoluensis Aytaç & H. D uman C. thessala Hausskn. C. tommasinii A. Kern. ( spinosociliata C. triniifolia Lacaita C. tenorei C. tenoreana Willk. C. tenoreana C. tymphaea Hausskn. C. ulrichiorum Wagenitz

Collectanea Botanica vol. 33 (enero-diciembre 2014), e001, ISSN-L: 0010-0730, http://dx.doi.org/10.3989/collectbot.2013.v33.001 22 A. HILPOLD, N. GARCIA-JACAS, R. VILATERSANA & A. SUSANNA Subsp. b ; Wagenitz & Hellwig, 1996 Wagenitz U ysal et al. , 2010 – + 1974 Wagenitz, + Sozen & O zaydin, 2010 + ITS Literature Other – Chromosome number (2n) number Chromosome – 18 (Özaydyn, 2007) – Greece, W BalkansW Greece, 1983) & Phitos, 1976; Georgiadis, 18 (Georgiadis + Anatolia Distribution N Black Sea Anatolia Anatolia C C C C C Subsect. et al. , 2006) . aggr.) oincy var. suffrutescens Blanca. C oincy var. et al. (2007 a ) as C. rouyi C. arenaria arenaria ( C. D C. Wagenitz Wagenitz Table 2. ( C ont.) Table C. zuccariniana D Species C. zeybekii C. wolgensis C. . A. C. wiedemanniana Fisch. & C . Mey. C. yozgatensis Wagenitz In Suárez-Santiago Unassigned to any section (Wagenitz et al. (2006). Published in Wagenitz 1 2 3

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