Centaurea onopordifolia Boissier (1849: 114)
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publication ID |
https://doi.org/ 10.11646/phytotaxa.363.1.1 |
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persistent identifier |
https://treatment.plazi.org/id/03E887F4-DF66-FFB2-19D4-3922F2DFFCE6 |
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treatment provided by |
Felipe |
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scientific name |
Centaurea onopordifolia Boissier (1849: 114) |
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1. Centaurea onopordifolia Boissier (1849: 114) View in CoL . Type :— SYRIA. Vallees d’Antiliban, July 1846, Boissier s.n. (holotype G!: six sheets comprising one specimen, isotypes P!: two sheets). ( Fig. 79 View FIGURE 79 )
Biennial plants, with thick taproot, whole plant thistle-like, grayish-green, 70–120 cm; collar of fibrous petiolar remains present at stem base. Stems spiny-winged, with minor and undulate triangular spiny-margined lobes, several, 5 to 15, branched at base, densely leafy throughout, 3–8 mm in diam. at base, densely covered with adpressed white arachnoid-tomentose hairs. Leaves rigid, spiny-margined, densely covered with adpressed white arachnoid-tomentose hairs, sometimes bicolor, above grayish, below whitish. Basal and lower cauline leaves with a short petiole, pinnatisect or pinnatifid, or pinnatilobed, (10–) 20–30 cm long, sometimes withered at anthesis, oblong or lanceolate. Median and upper cauline leaves sessile, pinnatisect or pinnatifid, oblong or triangular, uppermost increasingly smaller. Capitula numerous, arranged in a racemose dichasium, loose panicle or corymb, rarely raceme, on subsessile and arachnoid-tomentose branches, usually wingless above, sometimes with peduncles up to 6 cm long, sometimes with bracts similar to phyllaries with small stramineous or chaffy appendages. Involucres ovoid-oblong, convex at base, 28–35 × (12–) 14–16 mm. Phyllaries multiseriate, imbricate, coriaceous-scarious, green in vivo, brownish-green in sicco, densely lanate-tomentose. Appendages rigid, large, concealing a major part of phyllaries, triangular to elongate triangular, yellow or pale brown, 1.5–2.5 mm wide at base (excluding cilia), not decurrent, spinulate, innermost pectinate or rounded; spinules few, 3–4 on each side, (2–) 4–10 mm long; spine narrowly triangular, robust, 5–25 mm long, longer than adjacent spinules. Outer phyllaries ovate-oblong, 1.5–3.5 × 6–8 mm; median phyllaries oblong or lanceolate, 6–8 × 4–5 mm; inner phyllaries narrowly lanceolate to linear, 14–17 × 2–4 mm long. Flowers yellow; central florets hermaphroditic, 23–25 mm long, corolla 11–12 mm long, 5-lobed, lobes 4–5 mm long, with brownish nerves, style longer than corolla, stigma exserted from corolla, up to 3 mm; peripheral florets sterile, shorter than central ones, finely dissected, not radiant, 5-lobed, limb lobe filiform. Achenes lanceolate or oblong, 5–5.5 mm long, 2.2–2.5 mm wide, compressed, dark brown to blackish, smooth, rounded at apex, glabrescent; insertion areole lateral, 0.8–1 mm long. Pappus persistent, multiseriate, scabrous, brown, twice as long as achenes, ca. 10–11.5 mm long, bristles of inner rows much shorter than others, 2.5–3 mm long.
Nomenclatural notes: —At G there are six original sheets of Centaurea onopordifolia . Wagenitz (1960: 491) designated one of them as a lectotype and the other as an isolectotype. Our study showed that these six sheets belong to one specimen, so that there is only one specimen at G and it must be automatically considered as a holotype. Consequently, the lectotype formerly designated is unnecessary according to Art. 9.1 ( McNeill et al. 2012).
Taxonomic and distribution remarks: — Centaurea onopordifolia occurs in Lebanon, Palestine and Syria ( Fig. 80 View FIGURE 80 ). It is an Irano-Turanian element, and grows on dry steppe, rocky brown soil and stony slopes, scattered between Amygdalus shrubs, and mid-montane zone, at elevations of 1200–1800 m. Centaurea onopordifolia is the only species of the genus Centaurea with that thistle-like habit, stem spiny-winged, with minor and undulate triangular spiny-margined lobes, and also leaves spiny-margined. By these morphological characters, C. onopordifolia is similar to members of the genera Onopordum Linnaeus (1753: 827) , Carduus Linnaeus (1753: 820) and Cirsium Miller (1754: 334) , but differs from them by many characters such as: presence of appendages and peripheral florets, and also persistent pappus. For this reason we transferred this species to a new section, i.e. Centaurea sect. Cardueae here.
Chromosome numbers: —Unknown.
Selected specimens examined:— SYRIA. Antilibanon, inter Damaskus et Zale , 22 August 1937, Rechinger 2158 ( K!, W!) ; Antilibani, in arvis et siccis ad Zebedani, 1400 m (in saxosis supra Zebedani ), 23 June 1897, Bornmüller 924 ( B!, G!, P!, W!) ; Libanon oder angrenzendes Syrien: Coelesyrie , au pied oriental du Liban, pres de Deir-el-Ahmar, 19 July 1861, Blanche 3426 ( G!) ; Coelesyria: Zair , 6 July 1890, Peyron 1747 ( G!, P!) ; Zahli, Louis s.n. ( P!) ; Leyhaya , 26 June 1939, Louis s.n. ( P!) ; Jebel ed Druz, zwischen Suwaydah und Sale , Gebüsche und Felder , 1450 m, 5 August 1983, Frey & Kürschner 83-524 ( E!) ; Djebel-Druze , 10 July 1929, Gombault s.n. ( P!) ; 6 July 1930, Gombault s.n. ( P!) . LEBANON. Aitanit et Jesir-el-Kuwah ( Jisr-ul-Kuwwah ), Post 68 ( G!) . PALESTINE. Hermon, Busheri memorial, 1800 m, 16 July 1997, Gertman & Marta 122232 ( HUJ!) ; Hermon, between Shiba and Yina , dried steppe, 1200–1500 m, 21 July 1924, Eig 5329 ( HUJ!) ; Hermon, Wadi Yina , 21 July 1924, Smoli 5228 ( HUJ!) ; Hermon , 1600 m, 18 August 1969, Margalit s.n. ( HUJ!) ; S of Hermon, above Majdal Shams , rocky brown soil, 1300–1800 m, 23 July 1968, Danin s.n. ( HUJ!) ; Hermon, above Majdal Shams , S. exposure, stony slopes, 1350 m, 31 May 1989, Cohen s.n. ( HUJ!: two sheets) .
Biogeographical distribution and ecology of Centaurea sect. Cynaroides
Primarily focusing on the western parts of Iran and eastern parts of Turkey, the distributional range of C. sect. Cynaroides was profoundly revised ( Fig. 81 View FIGURE 81 ). Mostly, species were found to be restricted to very small areas which can be taken as evidence for a rapid speciation in this section ( Negaresh & Rahiminejad 2016). Except for C. charrelii which as a Mediterranean element grows exclusively in Greece, rest of the species are Irano-Turanian elements occurring in Iran, Turkey, Iraq and Syria ( Negaresh & Rahiminejad 2014, 2016) ( Fig. 81 View FIGURE 81 ).
In this section, species display a wide range of ecological needs. Their substrates vary from stony calcareous, rocky slopes, dry slopes, dry or green hills, dry grassy open hillsides, dry banks, clay slopes, grassland, stony plains, stony plains, limestone cliffs, serpentine soil, gravel slopes to basalt soil ( Figs. 3 View FIGURE 3 , 7D View FIGURE 7 , 41 View FIGURE 41 , 48C View FIGURE 48 , 62C View FIGURE 62 , 65C View FIGURE 65 , 66B, 66C View FIGURE 66 , 70B View FIGURE 70 , 71C View FIGURE 71 ). They are accompanied with shrubberies, steppes, limestone grasslands, or scattered in the woodlands especially of pines, oaks or cypresses ( Figs. 3 View FIGURE 3 , 41 View FIGURE 41 , 82 View FIGURE 82 , 83 View FIGURE 83 ). Field observations revealed that this section prefers foothills, meadow-steppe slopes and low to high elevations of mountainous zones. Even some taxa may grow in disturbed habitats such as: riversides, roadsides, champ side, dry waste at foothills, in vineries and pastures ( Figs. 3 View FIGURE 3 , 11C View FIGURE 11 , 12C View FIGURE 12 , 17B View FIGURE 17 , 41 View FIGURE 41 , 44C View FIGURE 44 , 48 View FIGURE 48 , 50C, 50D View FIGURE 50 , 56C View FIGURE 56 , 65 View FIGURE 65 , 66 View FIGURE 66 ).
The species of C. sect. Cynaroides are sporadically or densely grouped as patches of 2–5 or 15–30(–40) individuals ( Figs. 3B, 3D, 3F View FIGURE 3 , 41 View FIGURE 41 , 50 View FIGURE 50 , 82 View FIGURE 82 , 83 View FIGURE 83 ). Centaurea spicata , C. haradjianii , C. charrelii , C. zaferii and C. cataonica usually occur at the elevations less than 600 m (sometimes ca. 50 m), while C. geluensis occurs higher than 2400 m; however, the favorite altitude of C. sect. Cynaroides is between 600–2200 m.
Iran harbours 19 accepted species, 3 subspecies and 4 varieties generally occurring along Zagros Mountains , from southwest to northwest and central parts ( Fig. 81 View FIGURE 81 ). This country can be considered as one of the centers (likely the most important) of origin and diversity of a part of the section. A considerable number of taxa as bellow: Centaurea ravansarensis , C. shahuensis , C. alfonsoi , C. karamianiae , C. mozaffarianii , C. geluensis , C. iranshahrii , C. phlomoides , C. kabirkuhensis , C. kamalnejadii , C. rahiminejadii , C. solitaria , C. shehbazii , C. bavegehensis , C. kamyaranensis , C. amadanensis subsp. gymnoclada , C. amadanensis subsp. ranjbarii , C. regia var. longispinosa , and C. regia var. purpurea are exclusively endemic to Iran. Unlike the endemics that are known merely from their type locality ( C. geluensis , C. shahuensis , C. karamianiae , C. mozaffarianii , C. rahiminejadii , C. kamyaranensis , and C. amadanensis subsp. ranjbarii ) some are widely distributed like C. imperialis (especially in W and NW) and C. amadanensis (especially in C and W). The hotspot of diversity of the section seems to be located in the western parts of the country. With 16 endemic species, Kermanshah Province and Shahu Mountains in this area with 10 species, 1 subspecies and 2 verities are to be regarded as the beating heart of the section ( Fig. 84 View FIGURE 84 ).
Turkey with 18 recognized species, 3 subspecies and 2 varieties may be accounted for the other centre of diversity of the section. The following are strict endemics of this country: Centaurea sclerolepis , C. fenzlii , C. haradjianii , C. aladaghensis , C. zaferii , C. tomentella , C. amanicola , C. zarrei , C. sennikoviana , C. haussknechtii , C. kurdica subsp. kurdica , C. kurdica subsp. ciliata , and C. cataonica subsp. cataonica . Similar to Iran, some Turkish species like C. fenzlii (especially in SE), C. kurdica (especially in S), and C. sclerolepis (especially in S) are of widespread distribution, while some are mainly growing in the southern part of Turkey. Five species grow in Amanus Mountains on the border line between Turkey and Syria and are endemic to this region. The hotspot of diversity of Turkey is Amanus Mountains. Taking into account that the gene pool of the genus Centaurea is mainly Mediterranean and Amanus Mountains is located in the southern part of the country, it can be assumed that C. sect. Cynaroides had originally originated in this area and subsequently migrated eastwards to the Irano-Turanian region ( Fig. 84 View FIGURE 84 ).
Cytotaxonomy, homoploidy and evolution of Centaurea sect. Cynaroides
Cytotaxonomic studies can be a powerful tool for investigating different evolutionary trends such as breeding system or polyploidy and hybridization ( Uysal et al. 2009). A close correlation between karyology and systematics in the subtribe Centaureinae has been demonstrated repeatedly. In the genus Centaurea s. str., basic chromosome numbers are a valuable character for sectional classification ( Guinochet & Foissac 1962, Tonjan 1980, Garcia-Jacas & Susanna 1992, Garcia-Jacas et al. 1996, Wagenitz & Hellwig 1996a, Garcia-Jacas et al. 1997, Romashchenko et al. 2004, Uysal et al. 2009, Ranjbar & Negaresh 2013). Descending dysploidy, polyploidy and large differences in the size of chromosomes are three important karyological trends in Centaurea ( Uysal et al. 2009) . One of the better-documented karyological characters in Centaurea is descending dysploidy, ranging from x = 12 to x = 7 ( Garcia-Jacas et al. 1996, Uysal et al. 2009). The general trend towards descending dysploidy in Centaurea could be correlated with adaptation of mesophyllous taxa to the xeric conditions of the Mediterranean ( Uysal et al. 2009). Other examples among subtribe Carduinae support this view ( Garnatje et al. 2004).
The chromosome number 2 n = 18 had been determined for 34 populations (including 17 species, 2 subspecies and 2 varieties) of C. sect. Cynaroides (96%); the number of species for which ploidy level was scored was not statistically significant among taxa (see Tables 1 and 5). Our morphological studies and field observations indicated the presence of some hybrid characters between species. Moreover, in spite of more diversity in morphological characters and pollinators, there is no difference in chromosome numbers (see Fig. 9 View FIGURE 9 ). Basic chromosome number within the section may suggest the occurrence of an extensive evolution by homoploidy. This evidence demonstrates that homoploidy may be common in C. sect. Cynaroides and along with pollinators play an important role in the speciation. So, we found homoploidy in the center diversity of W Iran, whereas the only tetraploid occurs in very remote regions ( Greece). Finally, as it has been repeatedly suggested ( Fernández Casas & Susanna 1986, Georgiadis 1995, Garcia-Jacas et al. 1998 a, 1998 b, Constantinidis et al. 2002, Romashchenko et al. 2004, Uysal et al. 2009), homoploidy in C. sect. Cynaroides agrees with the evolution suggested by morphology and nuclear-DNA sequences.
Pollination
The genus Centaurea is discussed frequenly in the literature, but its pollination biology has received little attention. Watson & Renney (1974) noted that the members of Centaurea are entomophilous and self-compatible, but they did neither go into further details about specific floral visitors nor did they provide supporting data. Proctor & Yeo (1979) referred to visitation of several insects to Centaurea species but provided no quantitative data. Competition for pollinators, plant-insect interactions, connection between pollinators and evolution in flower structure and reproduction and pollination biology was reported ( Lack 1976, 1982 a, 1982b, 1982c, Harrod & Taylor 1995, Ehlers 1999, Agrawal et al. 2000, Hirsch et al. 2003) for some Centaurea species.
The most commonly observed insect visitors of yellow starthistle capitula were Hymenoptera spp., especially the European honeybee. Principal visitors for Centaurea diffusa , C. maculosa and C. solstitialis are Bombus spp. , Anthophora sp. , Apis mellifera and Megachile sp. ( Harrod & Taylor 1995). According to Harrod & Taylor (1995), Apis mellifera may increase the reproductive capacities of Centaurea . Recently, Aslan et al. (2011) defined Larinus latus as a pollinator for Centaurea regia subsp. regia . Our studies showed that different pollinators play role in pollination in C. sect. Cynaroides . Some pollinators are common between different species, whereas the others are entirely specific ( Figs. 85 View FIGURE 85 and 86 View FIGURE 86 ). Most species belonging to C. sect. Cynaroides are often rare with a narrow distribution. In addition, differences in color and size of flowers may be for adaptation to various pollinators. Some pollinators for this section are family Curculionidae : Larinus latus and Larinus sp. ( Figs. 85B, 85C View FIGURE 85 , and 86G–86I View FIGURE 86 ); family Scarabaeidae : Oxythyrea cinctella , Cetonia spp. and Potocia spp. ( Figs. 85H View FIGURE 85 , 86B, 86F and 86K View FIGURE 86 ); Ant: family Formicidae ( Figs. 85B, 85E and 85F View FIGURE 85 ); Bees: family Megachilidae : Megachile spp. ( Fig. 86C View FIGURE 86 ), family Apidae : Apis spp. ( Figs. 85J and 85K View FIGURE 85 ); family Syrphidae ( Figs. 85D and 85G View FIGURE 85 ); and Butterflies ( Figs. 85A View FIGURE 85 and 86D View FIGURE 86 ).
| K |
Royal Botanic Gardens |
| W |
Naturhistorisches Museum Wien |
| B |
Botanischer Garten und Botanisches Museum Berlin-Dahlem, Zentraleinrichtung der Freien Universitaet |
| G |
Conservatoire et Jardin botaniques de la Ville de Genève |
| P |
Museum National d' Histoire Naturelle, Paris (MNHN) - Vascular Plants |
| E |
Royal Botanic Garden Edinburgh |
| HUJ |
Hebrew University |
| S |
Department of Botany, Swedish Museum of Natural History |
| C |
University of Copenhagen |
| A |
Harvard University - Arnold Arboretum |
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