On the identity of Centaurea paradoxa (Asteraceae, Cardueae, Centaureinae) from Iran Authors: Ranjbar, Massoud, and Negaresh, Kazem Source: Willdenowia, 44(1) : 13-20 Published By: Botanic Garden and Botanical Museum Berlin (BGBM) URL: https://doi.org/10.3372/wi.44.44102 BioOne Complete (complete.BioOne.org) is a full-text database of 200 subscribed and open-access titles in the biological, ecological, and environmental sciences published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Complete website, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/terms-of-use. Usage of BioOne Complete content is strictly limited to personal, educational, and non - commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Downloaded From: https://bioone.org/journals/Willdenowia on 12 Dec 2021 Terms of Use: https://bioone.org/terms-of-use Willdenowia 44 – 2014 13 MASSOUD RANJBAR1* & KAZEM NEGARESH1 On the identity of Centaurea paradoxa (Asteraceae, Cardueae, Centaureinae) from Iran Abstract Ranjbar M. & Negaresh K.: On the identity of Centaurea paradoxa (Asteraceae, Cardueae, Centaureinae) from Iran. – Willdenowia 44: 13 – 20. 6 March 2014. – Version of record published online ahead of inclusion in April 2014 issue; ISSN 1868-6397; © 2014 BGBM Berlin-Dahlem. DOI: http://dx.doi.org/10.3372/wi.44.44102 New findings and a revised description are presented for Klasea sect. Leuzeopsis (Boriss.) L. Martins (Asteraceae, Cardueae, Centaureinae). This previously monotypic section now contains one more species: K. paradoxa (Mozaff.) Ranjbar & Negaresh, which is proposed here as a new combination. Lectotypes are designated for Leuzea serratuloides Fisch. & C. A. Mey. ex DC. (K. serratuloides (Fisch. & C. A. Mey. ex DC.) Greuter & Wagenitz) and two of its synonyms. Descriptions, distributions, ecology and illustrations for both species are given, as is the conservation status for K. paradoxa. The differences between the two species are tabulated and an identification key is provided. The geographical distributions of the two species are mapped. Additional key words: Klasea, new combination, lectotypification, Compositae Introduction Klasea Cass. is the second largest genus of Asteraceae subtribe Centaureinae in Iran. According to modern taxonomy, the genus includes eight sections in Iran: K. sect. Coriaceae L. Martins, K. sect. Demetria (Boriss.) L. Martins, K. sect. Grandifoliae L. Martins, K. sect. Klasea, K. sect. Melanolepis L. Martins, K. sect. Nikitinia (Iljin) L. Martins, K. sect. Quinquefoliae L. Martins and K. sect. Schumeria (Iljin) L. Martins (Martins 2006; Ranjbar & al. 2011). It has been reported that Klasea comprises 17 species in Iran (Dittrich & al. 1980; Mozaffarian 1992; Martins 2006; Ranjbar & al. 2011, 2012). The present study confirms that K. sect. Leuzeopsis (Boriss.) L. Martins is also present in Iran, increasing the number of Klasea sections in that country to nine and the number of species to 21. During recent floristic studies in W Iran, a plant similar to Centaurea L. was collected that initially could not be identified to an existing species. After thorough consultation of the relevant floristic literature (Dittrich & al. 1980; Mozaffarian 1991, 1992; Martins 2006; Ranjbar & al. 2011, 2012), and comparison with specimens in the herbaria B, BASU, E, G, LD, TARI, W and WU, it was concluded that (1) this material as well as two specimens at B collected in the same area in W Iran in 1904 by Th. Strauss match what has been named C. paradoxa Mozaff. (Mozaffarian 1991) and (2) C. paradoxa in fact belongs to Klasea, in particular to K. sect. Leuzeopsis, and is most similar to K. serratuloides (Fisch. & C. A. Mey. ex DC.) Greuter & Wagenitz. It was therefore decided to transfer C. paradoxa to Klasea, as K. paradoxa. The main differences between the latter species and K. serratuloides are presented in Table 1. 1 Department of Biology, Herbarium Division, Bu-Ali-Sina University, PO Box 65175/4161, Hamedan, Iran; *e-mail: ranjbar@ basu.ac.ir (author for correspondence). Downloaded From: https://bioone.org/journals/Willdenowia on 12 Dec 2021 Terms of Use: https://bioone.org/terms-of-use 14 Ranjbar & Negaresh: On the identity of Centaurea paradoxa from Iran Table 1. Morphological comparison of Klasea paradoxa and K. serratuloides. Klasea paradoxa Klasea serratuloides 35 – 70 always divided, usually pinnatilobed, sometimes pinnatisect or pinnatifid oblong, elliptic or ovate 18 – 40 4-6 pairs of segments dentate or lobed, rarely entire 5 – 35 undivided or lyrate-pinnatisect 15 – 40 sparsely arachnoid to subglabrous 5 – 10 35 – 47 20 – 27 yellowish cream lanceolate or lanceolate-arcuate 2–7 glabrous 5–6 30 – 33 up to 16 brown ovate or oblong-ovate 10 – 14 × 4 – 4.5 20 – 24 6 – 7( – 7.5) × 2.5 – 3( – 3.5) 11 – 15 Character Plant height [cm] Leaf dissection Leaf shape [in outline] Basal and lower cauline [B+LC] leaf length [cm] B+LC leaf segments or lobes number B+LC leaf segments or lobes margin Peduncle length [cm] Phyllary indumentum Phyllary spine length [mm] Floret length [mm] Corolla tube length [mm] Achene colour Achene shape Achene size [mm] Pappus length [mm] oblong or obovate 10 – 17( – 20) 2 or 3 pairs of lobes irregularly crenate-toothed or sinuate-lobulate Material and methods Key to species of Klasea sect. Leuzeopsis The present study is mainly based on herbarium material. Several sheets have been examined for each species, either received on loan from the herbaria BASU, G, LD, W and WU or obtained as digitized type material from the herbaria B, BR, E, JE, LE and P. Moreover, during several excursions from 2008 to 2013 in W Iran, the authors have studied living plants in the field. 1. Leaves always divided, usually pinnatilobed, sometimes pinnatisect or pinnatifid; florets 35 – 47 mm long; achenes yellowish cream, 10 – 14 mm long; pappus 20 – 24 mm long . . . . . . . . . . . . 1. K. paradoxa – Leaves undivided or lyrate-pinnatisect; flowers 30 – 33 mm long; achenes brown, 6 – 7( – 7.5) mm long; pappus 11 – 15 mm long . . . . . . 2. K. serratuloides Results and Discussion 1. Klasea paradoxa (Mozaff.) Ranjbar & Negaresh, comb. nov. ≡ Centaurea paradoxa Mozaff. in Iran. J. Bot. 5: 30. 1991. – Holotype: Iran, Lorestan, Malavi to Eslamabad road, c. 5 km from Buluran to Eslamabad, 1330 m, 9 May 1988, V. Mozaffarian 64381 (TARI!) – Fig. 1, 2. Klasea sect. Leuzeopsis (Boriss.) L. Martins in Bot. J. Linn. Soc. 152: 459. 2006 ≡ Serratula sect. Leuzeopsis Boriss., Fl. URSS 28: 607. 1963. – Type: Serratula serratuloides (Fisch. & C. A. Mey. ex DC.) Takht. Description — Herbs perennial, monocephalous, 5 – 70 cm tall. Leaves undivided to pinnatilobed with 2 – 6 pairs of lobes or segments, oblong, elliptic, ovate or obovate in outline, 10 – 40 cm, coriaceous, with prominent veins mostly in proximal part, margin (crenate-)dentate or lobed, rarely entire, apex obtuse or acute. Capitulum large; peduncle short to long; involucre subglobose, 3 – 4 × 3 – 5 cm; phyllaries coriaceous, sparsely arachnoid to glabrous, with an often deflected spine; spine 5 – 10 mm long. Florets pink or mauve, 30 – 50 mm long; corolla tube to 27 mm long; apical anther appendages subacute to slightly emarginate. Achenes yellowish cream or brown, ovate to lanceolate, 6 – 14 × 2.5 – 4.5 mm, glabrous, abruptly narrowed toward base; pappus 11 – 24 mm long, c. 2 × as long as achene. Distribution — Turkey, Armenia, Azerbaijan and Iran. Downloaded From: https://bioone.org/journals/Willdenowia on 12 Dec 2021 Terms of Use: https://bioone.org/terms-of-use Description — Herbs perennial, single-stemmed, usually green throughout, 35 – 70 cm tall. Stem erect, always simple, monocephalous, stout, 4 – 5 mm in diam. at base, sparsely leafy but leafless on peduncle, sulcate, loosely to densely covered with tomentose hairs. Leaves always divided, large, coriaceous, with prominent veins, ± loosely covered with tomentose hairs. Basal and lower cauline leaves long petiolate (petiole to 10 cm long), divided, usually pinnatilobed, sometimes pinnatisect or pinnatifid, oblong or ovate in outline, 18 – 40 × 6 – 14 cm; segments in 4 – 6 pairs, oblong, triangular-lanceolate or triangular, 2 – 6 × 0.8 – 3.5 cm, margin dentate or lobed, rarely entire, apex acute or shortly acuminate. Median cauline leaves petiolate, pinnatilobed, oblong or elliptic in outline, 7.5 – 12 × 3 – 7 cm; segments in 2 – 5 pairs, triangular, 0.5 – 2 × 0.2 – 1 cm, margin entire, rarely dentate, apex shortly acuminate. Capitulum solitary; peduncle leafless, 15 – 40 cm Willdenowia 44 – 2014 15 Fig. 1. Klasea paradoxa – A, C: habit; B: capitula with peduncles; C, D: habitat. – A – D: Iran, Lorestan, Malavi to Eslamabad, 25 Apr 2013, photographs by M. Ranjbar. – Scale bars: A = 5 cm; B = 4 cm; C = 6 cm. Downloaded From: https://bioone.org/journals/Willdenowia on 12 Dec 2021 Terms of Use: https://bioone.org/terms-of-use 16 Ranjbar & Negaresh: On the identity of Centaurea paradoxa from Iran Fig. 2. Klasea paradoxa – A: leaves showing venation and indumentum; B: capitulum; C: florets with pollinator; D, E: achenes with pappus. – A – E: Iran, Lorestan, Malavi to Eslamabad, 25 Apr 2013, photographs by M. Ranjbar. – Scale bars: A = 10 cm; B = 2 cm; C = 10 mm; D, E = 5 mm. long. Involucre subglobose, 30 – 40 × 30 – 50 mm. Phyllaries 8 – 10-seriate, imbricate, green, purplish distally, with longitudinal dark green veins, coriaceous, sparsely arachnoid to subglabrous, apex almost constricted, with a spine; spine often deflected, 5 – 10 mm long, scabrous; outer phyllaries lanceolate, 7 – 10 × 3.5 – 5 mm; median phyllaries oblong or oblong-ovate, 12 – 17 × 7 – 9 mm; inner phyllaries oblong or oblong-linear, 20 – 25 × 5 – 7 mm. Receptacle setose, with long smooth bristles. Florets pink or mauve when living, purplish yellow when dried, 35 – 47 mm long; corolla tube narrow, 20 – 27 mm long; limb 15 – 20 mm long; lobes 6 – 9 mm long; anther tube shorter than corolla; filaments glabrous; apical anther appendages slightly emarginate or subacute; style and stigma partly exserted from corolla, pubescent. Achenes yellowish cream, lanceolate or lanceolate-arcuate, 10 – 14 × 4 – 4.5 mm, glabrous, smooth and shiny; pappus deciduous, multiseriate, plumose, white, 20 – 24 mm long, bristles unequal. Downloaded From: https://bioone.org/journals/Willdenowia on 12 Dec 2021 Terms of Use: https://bioone.org/terms-of-use Phenology — Flowering from April to May; fruit ripening from May to June. Distribution and ecology — Klasea paradoxa is a Zagrosian element endemic to Lorestan province, W Iran (Fig. 3). It grows on dry rubbly and rocky slopes and talus in the mid-montane zone at altitudes of 1300 – 1400 m (Fig. 1C, D). Other plants observed to occur in the area include: Ajuga chamaecistus Ging. ex Benth., Alyssum constellatum Boiss., Amygdalus scoparia Spach, Astragalus multijugus DC., Centaurea koieana Bornm., Euphorbia macroclada Boiss., Klasea cerinthifolia (Sm.) Greuter & Wagenitz, Matthiola longipetala (Vent.) DC., Onosma macrophylla Bornm., Quercus brantii Lindl., Salvia bracteata Banks & Sol., Silene chlorifolia Sm., Scrophularia sp. and Trigonella disperma Bornm. Conservation status — Klasea paradoxa is a narrow endemic with an estimated area of occupancy is less than Willdenowia 44 – 2014 17 2. Klasea serratuloides (Fisch. & C. A. Mey. ex DC.) Greuter & Wagenitz in Willdenowia 33: 59. 2003 ≡ Leuzea serratuloides Fisch. & C. A. Mey. ex DC., Prodr. 6: 666. 1838 ≡ Serratula serratuloides (Fisch. & C. A. Mey. ex DC.) Takht. in Trudy Armyansk. Fil. Akad. Nauk S.S.S.R., Ser. Biol. 2: 90. 1937. – Lectotype (designated here): Azerbaijan, Nakitschiwan [Nakhichevan, on some labels Karabagh], in declivitatibus montium vallis Koschadara, Szowits (GDC!; isolectotypes: K!, LE!, P!) – Fig. 4. = Leuzea sclerophylla Boiss. & Heldr., Diagn. Pl. Orient., ser. 1, 10: 97. 1849 ≡ L. serratuloides var. subindivisa Boiss., Fl. Orient. 3: 607. 1875. – Lectotype (desigFig. 3. Distribution of Klasea paradoxa (square) and K. serratuloides (circles). nated here): Turkey, in siccis vallis superioris Ermenek Ciliciae Tracheae prope 10 km2. The population is under continuous threat of ex- pagum Betlam, 3 Jul 1845, Heldreich (G!; isolectotype: cessive grazing. Therefore, the species should be catego- G!). rized as Critically Endangered (CR) according to IUCN = Phaeopappus freynii Sint. ex Freyn in Oesterr. Bot. Z. criteria B2ab(v) (IUCN 2012). 42: 235. 1892. – Lectotype (designated here): Turkey, Armenia turcica, Kharput, in declivibus mont. Karatasch, Remarks — Centaurea paradoxa is transferred here 12 Jun 1889, Sintenis 761 (LD!; isolectotypes: B!, BR!, from Centaurea to Klasea because of its bisexual ho- E!, JE!, K [two sheets]!, W!). mogamous capitula with tubular florets and phyllaries with parallel veins. Klasea paradoxa belongs to K. sect. Description — Herbs perennial, 5 – 35 cm tall; root strong, Leuzeopsis, which includes taxa that often have simple woody. Stem simple, angled or ± straight, strong, usually stems, large coriaceous leaves, large capitula (3–4 cm short, less often somewhat surpassing lower leaves, leafin diam.), deflected spines and apical anther appendag- less in upper part (2 – 7 cm), sulcate, sparsely or rarely es usually slightly emarginate. According to Borissova densely pubescent. Leaves petiolate (petiole 3 – 6( – 12) cm (1963) and Martins (2006), K. sect. Leuzeopsis con- long), undivided or lyrate-pinnatisect, oblong or obovate tained a single species, namely K. (Serratula) serratu- in outline, 10 – 17( – 20) × 6 – 10( – 20) cm, thick, corialoides. Klasea paradoxa resembles K. serratuloides in ceous, with prominent veins, thinly hairy on lower surlife form, involucre size and floret colour (cf. Fig. 4), face mostly along veins, subglabrous on upper surface; but differs from it in certain characters (see Table 1), lateral lobes in 2 or 3 pairs, oblong, apex obtuse; terminotable among which are: leaf dissection always divid- nal lobe ovate or ovate-oblong, larger than lateral lobes, ed, usually pinnatilobed, sometimes pinnatisect or pin- margin irregularly crenate-toothed or sinuate-lobulate. natifid (vs undivided or lyrate-pinnatisect in K. serratu- Capitula solitary; peduncle leafless, to 7 cm long. Involuloides), corolla tube 20 – 27 mm long (vs up to 16 mm cre subglobose, 3 – 4 × 4 – 5 cm. Phyllaries c. 8-seriate, long), phyllaries sparsely arachnoid to subglabrous moderately imbricate, green, yellow-brownish when dry, (vs glabrous), pappus 20 – 24 mm long (vs 11 – 15 mm with longitudinal dark green veins, coriaceous, glabrous, long) and achenes 10 – 14 × 4 – 4.5 mm (vs 6 – 7( – 7.5) margin narrowly membranous, apex with a spine; spine × 2.5 – 3( – 3.5) mm). Because of these differences, we often deflected, 5 – 6 mm long; outer phyllaries ovate; recognize K. paradoxa and K. serratuloides as separate median phyllaries oblong; inner phyllaries lanceolatespecies. Moreover, because the original description of linear, apex acuminate. Receptacle setose. Florets light C. paradoxa (Mozaffarian 1991) is rather short, we give magenta or pale rose, 30 – 33 mm long; corolla tube nara more complete description here. row, to 16 mm long; limb 15 – 17 mm long; lobes c. 7 mm long; anther tube slightly shorter than or as long as corolAdditional specimens seen — Iran: Lorestan prov- la; filaments glabrous; apical anther appendages rounded ince: road from Eslamabad to Poldokhtar, 40 km to or slightly emarginate; style and stigma exserted from Kuhdasht, Buluran, 1350 m, 25 Apr 2013, Ranjbar & corolla, usually shortly 2-lobed. Achenes brown, ovate Negaresh 33056, 33057 & 33058 (BASU!); near Bulu- or oblong-ovate, strongly compressed, distally narrowed, ran, 65 km to Baba Zeyd, 1370 – 1400 m, 2 May 2013, 6 – 7( – 7.5) × 2.5 – 3( – 3.5) mm, weakly sulcate or smooth, Ranjbar & Negaresh 33059 & 33060 (BASU!); Kuh glabrous, base subacute; pappus multiseriate, plumose, Tarikha s.ö. v. [SE of] Kermanschah, 11 May 1904, Th. white, 11 – 15 mm long, bristles connate at base into a Strauss (B [2 sheets]!). ring. Downloaded From: https://bioone.org/journals/Willdenowia on 12 Dec 2021 Terms of Use: https://bioone.org/terms-of-use 18 Ranjbar & Negaresh: On the identity of Centaurea paradoxa from Iran Fig. 4. Isolectotype of Leuzea serratuloides (≡ Klasea serratuloides) at P. – Image provided by Muséum National d’Histoire Naturelle, Paris. Downloaded From: https://bioone.org/journals/Willdenowia on 12 Dec 2021 Terms of Use: https://bioone.org/terms-of-use Willdenowia 44 – 2014 Phenology — Flowering from May to July; fruit ripening from July to August. Distribution and ecology — Klasea serratuloides occurs in Turkey, Armenia and Azerbaijan (Fig. 3). It is an Irano-Turanian element known only from the midmontane zone, occurring on dry rubbly and rocky slopes, talus, eroded shaly hills and banks in open forest of Pinus brutia Ten. at altitudes of 600 – 2000 m (Borissova 1963; Davis & Kupicha 1975). Remarks — Klasea serratuloides has been subjected to controversial classification in either subtribe Carduinae (e.g. Dittrich 1977) or Centaureinae (Adylov & Zuckerwanik 1993). The affinity of Serratula serratuloides to Klasea has been shown by Susanna & al. (2002), Martins & Hellwig (2005) and Martins (2006). Concerning reports of Klasea serratuloides in Iran, Borissova (1963) suspected that Serratula serratuloides occurred in Iran, and Davis & Kupicha (1975) stated that it occurred in Iran, but this taxon was not mentioned in Flora iranica (Dittrich & al. 1980). Martins (2006: Fig. 8) provided a distribution map recording K. serratuloides in W Iran. The basis for that record was presumably two specimens from the same area as the relevant dot on the distribution map: Kuh Tarikha s.ö. v. [SE of] Kermanschah, 11 May 1904, Th. Strauss (B), both determined by J. Bornmüller as Leuzea serratuloides, revised by G. Wagenitz in 1960 as S. serratuloides “var. ?” and revised by L. Martins in 2005 as K. serratuloides. We have examined digital photographs of these specimens and conclude that they in fact represent K. paradoxa. Additional specimens seen — Armenia: montes “Gegamski khrebet”, distr. Abovyan, inter monasterium Ge[g]hard et pagum Gokht, 1700 – 1800 m, 20 Jul 1975, V. Vašák (B!); Abovjan distr., near Geghard, 1650 m, steppe. 21 Jun 1996, E. Gabrieljan & al. (B!); Mikoian district, surroundings of Kavushug village [Hermon], left valley slope of Dzhaly-chai river, 1250 m, 17 Jun 1957, Avetisian & al. (W!); distr. Ararat, montes “Gegamski khrebet”, loco Aruni Dzor dicto, 1500 – 1900 m, 11 Jul 1975, V. Vašák (B!); Azizbekov distr., 2 km to Khndzorut from road Martiros-Khndzorut, right side of road with tragacanth shrub vegetation, 23 Jun 1987, Oganezova (B!); Azizbekov, surroundings of Bartsruni village [Bardzruni] on lime and cement slopes, 20 Jun 1957, Aslanian & al. (W!). Azerbaijan: Nakhichevan: pr. p. [prope pagum] Bitseh’enach, 18 Jun 1939, L. Prilipko (B!). Turkey: B9 Van: 8 km from Van to Gürpinar (Havasor), Davis 44592 (E!); Satak road, 2 km N of Micingersuyu river, 2000 m, 25 Jul 1954, Davis 2389 (E!). C4 Konya: 1300 m, 23 Jul 1995, Ayta 7227 (E!); 1000 m, 15 Aug 1949, Davis 16208 (E!); Ermenek-Fariske, Brachfeld, linkes Ufer des Göksu, 16 km N Ermenek, 710 m, 12 Jun Downloaded From: https://bioone.org/journals/Willdenowia on 12 Dec 2021 Terms of Use: https://bioone.org/terms-of-use 19 1950, Huber-Morath 10020 (E!). C5 Konya: Ereğli, Aydos Daği, between Berendi and Kiraman village, 1400 m, 28 Jun 1976, Erik 1756 (E!). C6 Maraş: Maraş to Göksun nr Yemiş Da., 600 m, 3 May 1957, Davis & Hedge 27513 (E!). Acknowledgements We are indebted to the curators of the following herbaria: B, BASU, BR, E, G, JE, LD, LE, P, W and WU, for their contribution to the revision of materials by making specimens available, either on loan or as digital images. The field work in Iran was supported by grants from Bu-Ali Sina University. We are also most grateful to Nicholas Turland and two anonymous reviewers, who improved the overall quality of the manuscript. References Adylov T. A. & Zuckerwanik T. I. 1993: Opredelitel’ rastenii srednei Azii (Conspectus florae Asiae Mediae) 10. – Tashkent: Academy of Sciences of the Republic of Uzbekistan. Borissova A. G. 1963: Serratula L. – Pp. 259 – 301 in: Bobrov E. G. & Czerepanov S. K. (ed.), Flora URSS 28. – Moskva & Leningrad: Izdatel’stvo Akademii Nauk SSSR. Davis P. H. & Kupicha F. K. 1975: Serratula. – Pp. 452 – 460 in: Davis P. H. 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