Abstract
Cousinia sect. Stenocephalae is a mainly Iranian section of the genus and the second largest section of Cousinia after C. sect. Cynaroidae. Its members are mainly distributed in Iran. Using nrDNA ITS sequences, together with morphological evidence where possible, we tested the monophyly of C. sect. Stenocephalae as well as the related C. sect. Albidae and C. sect. Cousinia. We reconstructed phylogenetic relationships within these sections by performing a Bayesian analysis on a sample of 153 species of Cousinia, of which 49 species belong to the study sections. The analysis revealed that C. sect. Stenocephalae, Albidae and Cousinia are not monophyletic, which lays stress upon the need of a deep reexamination of the sectional classification in Cousinia using a different approach.
References
<p>Ahmad-Khanbeygi, Z., Sheidai, M. & Attar, F. (2011) Morphometry and palynological study of the genus <em>Cousinia</em> sect. <em>Cousinia </em>(Asteraceae) in Iran. <em>Iranian Journal of Botany</em> 17 (2): 158–166.</p>
<p>Attar, F. (2011a) Taxonomic revision of <em>Cousinia</em> sect. <em>Albidae</em> Bunge (Asteraceae) in the flora of Iran. <em>Iranian Journal of Botany</em> 17 (2): 150–157.</p>
<p>Attar, F. (2011b) New taxa of <em>Cousinia </em>Cass. sect. <em>Stenocephalae </em>Bunge (Asteraceae) from Iran. <em>Taxonomy and Biosystematics</em> 6: 59–68.</p>
<p>Attar, F. & Djavadi, S.B. (2010) A taxonomic revision of<em> Cousinia</em> sect. <em>Cynaroides</em> (Asteraceae, Cardueae) in the flora of Iran. <em>Iranian Journal of Botany</em> 16 (1): 130–184.</p>
<p>Attar, F., Ghahreman, A., Saber, A. & Zarre, Sh. (2005) A new species of the genus <em>Cousinia </em>(Compositae, sect. <em>Stenocephalae</em>) from SW Iran. <em>Iranian Journal of Botany</em> 11 (1): 65– 69.</p>
<p>Attar, F. & Maroofi, H. (2010) <em>Cousinia</em> (sect. <em>Albidae</em>) <em>Iranshahriana</em> (Asteraceae, Cardueae), a new endemic species from Kurdistan province, Iran. <em>Iranian Journal of Botany</em> 16 (1): 197–199.</p>
<p>Bahraminejad, S., Amiri, R., Ghasemi, S. & Fathi, N. (2013) Inhibitory effect of some Iranian plant species against three plant pathogenic fungi. <em>International Journal of Agriculture and Crop Sciences</em> 5: 1002–1008.</p>
<p>Boissier, E. (1845) <em>Diagnosis plantarum orientalium novarum, ser. 1</em>, vol. 6. B. Hermann, Lipsiae [Leipzig].</p>
<p>Boissier, E. & Buhse, F.A. (1860) Aufzählung der auf einer Reise durch Transkaukasien und Persien gesammelten Pflanzen. <em>Nouveaux</em> <em>Meìmoires de la Socieìteì Impeìriale des Naturalistes de Moscou</em> 12: 1–246.</p>
<p>Boissier, E. (1875) <em>Flora Orientalis</em>. H. Georg, Geneva & Basilea [Basel].</p>
<p>Bordbar, F., Baqeri, M. & Mirtadzadini, M. (2017) New Chromosome Counts in <em>Cousinia</em> (Asteraceae)from Flora of Iran. <em>Cytologia</em>. 82 (5): 485–488.</p>
<p>Bornmüller, J.F.N. (1907) Beiträge zur Flora der Elbursgebirge Nord-Persiens. <em>Bulletin de l’Herbier Boissier</em>, ser. 2, 7: 209–224.</p>
<p>Bornmüller, J.F.N (1910) Bearbeitung der von J.A. Knapp im nordwestlichen Persien gesammelten Pflanzen. <em>Verhandlungen der Zoologisch-Botanischen Gesellschaft</em> <em>in Wien</em> 60: 1–512.</p>
<p>Bornmüller, J. (1911) Collectiones Straussianae novae. Weitere Beiträge zur Kenntnis der Flora West Persiens. <em>Beihefte zum Botanischen Centralblatt, Original Arbeiten</em> 28 (2): 225–267.</p>
<p>Bornmüller, J.F.N. (1912) Ein Beitrag zur Kenntnis der Gattung <em>Cousinia. Oesterreichische Botanische Zeitschrift</em> 62: 1–523. https://doi.org/10.1007/BF01632329</p>
<p>Bornmüller, J.F.N. (1916) Ein Beitrag zur Kenntnis der Gattung <em>Cousinia</em>. <em>Beihefte zum Botanischen Centralblatt</em>,<em> Zweite Abt.</em> 34: 131–203.</p>
<p>Bornmüller, J.F.N. (1934) Ein Beitrag zur Kenntnis der Gattung <em>Cousinia</em>. XII. Neue Arten und bemerkenswerte Formen aus Persien und Turkestan.<em> Repertorium Specierum Novarum Regni Vegetabilis</em> 36: 323–338.</p>
<p>Bornmüller, J.F.N. (1941) Neue Arten der Gattung <em>Cousinia</em> der Flora Persiens. <em>Mittheilungen des Thüringischen Botanischen Vereins</em> 47: 115–128.</p>
<p>Bunge, A. von (1841). Correspondenz.<em> Flora oder Botanische Zeitung </em>24 (1): 153–159.</p>
<p>Bunge, A. von (1865) Übersichtliche Zusammenstellung der Arten der Gattung <em>Cousinia</em> Cass. <em>Mémoires de l’Académie Impériale de Sciences de Saint Pétersbourg</em>, Ser. 7, 9: 1–56.</p>
<p>Cassini, A.H.G. (1827) <em>Cousinia</em>. <em>In</em>: Cuvier, G. (Ed.) <em>Dictionnaire des sciences naturelles</em>, vol. 47. Levrault, Strasbourg, pp. 503–505.</p>
<p>Candolle, A.P. de (1838) <em>Prodromus systematis naturalis regni vegetabilis,</em> vol. 6. Treuttel & Würtz, Paris.</p>
<p>Djamali, M., Baumel, A., Brewer, S., Jackson, S.T, Kadereit, J.W., López-Vinyallonga, S., Mehregan, I., Shabanian, E. & Simakova, A. (2012) Ecological implications of <em>Cousinia </em>Cass. (Asteraceae) persistence through the last two glacial–interglacial cycles in the continental Middle East for the Irano-Turanian flora. <em>Review of Palaeobotany and Palynology</em> 172: 10–20. https://doi.org/10.1016/j.revpalbo.2012.01.005</p>
<p>Djavadi, S.B. (2005) New or rare chromosome count in ten species of <em>Cousinia</em> from Iran. <em>Rostaniha </em>8 (1): 19–27.</p>
<p>Djavadi, S.B. (2012) New chromosome counts in <em>Cousinia</em> (Asteraceae, Cardueae) section <em>Cynaroideae </em>and <em>Stenocephalae</em> from Iran. <em>Iranian Journal of Botany</em> 18 (1): 86–93.</p>
<p>Djavadi, S.B. & Attar, F. (2006) Two new species of <em>Cousinia</em> Cass. of sect. <em>Stenocephalae</em> Bunge from Iran. <em>Feddes Repertorium</em> 117: 453–458. https://doi.org/10.1002/fedr.200611113</p>
<p>Djavadi, S.B. & Attar, F. (2010) New chromosome counts in the genus <em>Cousinia</em> (Asteraceae, Cardueae) from Iran. <em>Willdenowia</em> 40 (2): 351–357. https://doi.org/10.3372/wi.40.40214</p>
<p>Djavadi, S.B., Attar, F. & Najafi, Z. (2007) A new species of <em>Cousinia</em> sect. <em>Cousinia</em> from NW Iran. <em>Iranian Journal of Botany</em> 13 (1): 43–46.</p>
<p>Doyle, J.J. & Doyle, J.L. (1987) A rapid DNA isolation procedure for small quantities of fresh leaf tissue. <em>Phytochemical Bulletin</em> 19: 11–15.</p>
<p>Duistermaat, H. (1996) <em>Arctium</em> getting entangled to <em>Cou</em>s<em>inia</em> (Asteraceae: Cardueae). <em>Bocconea</em> 5: 685–689.</p>
<p>Edgar, R.C. (2004) Muscle: multiple sequence alignment with high accuracy and high throughput. <em>Nucleic Acids Research </em>32: 1792–1797. https://doi.org/10.1093/nar/gkh340</p>
<p>Ghaffari, S.M. & Chariat-Panahi, S. (1985) Chromosome count of some angiosperms from Iran. <em>Iranian Journal of Botany</em> 3 (1): 67–73.</p>
<p>Ghaffari, S.M. & Djavadi, S.B. (1998) Chromosome studies and distribution of nine species of <em>Cousinia</em> section <em>Stenocephalae</em> (Asteraceae) in Iran.<em> Bulletin de la Société Neuchâteloise des Sciences Naturelles</em> 121: 61–68.</p>
<p>Ghaffari, S.M., Susanna, A. & Garcia-Jacas, N. (2006) New chromosome counts in the genus <em>Cousinia </em>(Asteraceae) from Iran. <em>Botanical Journal of the Linnean Society </em>151: 411–419. https://doi.org/10.1111/j.1095-8339.2006.00506.x</p>
<p>Herder, F. von (1869) Plantae raddeanae monopetalae. <em>Bulletin de la Société Impériale des Naturalistes de Moscou</em> 41 (3): 1–53.</p>
<p>Herrando-Moraira, S., The Cardueae Radiations Group (Calleja, J.A., Galbany-Casals, M., Garcia-Jacas, N., Liu, J.-Q., López-Alvarado, J., López-Pujol, J., Mandel, J.R., Massó, S., Montes-Moreno, N., Roquet, C., Sáez, L., Sennikov, A., Susanna, A. & Vilatersana, R.) (2019) Nuclear and plastid DNA phylogeny of tribe Cardueae (Compositae) with Hyb-Seq data: A new subtribal classification and a temporal diversification framework. <em>Molecular Phylogenetics and Evolution</em> 137: 313–332. https://doi.org/10.1016/j.ympev.2019.05.001</p>
<p>Hiemeral, A. (1885) <em>Cousinia</em> Cass. <em>In: </em>Stapf, O., Die botanischen Ergebnisse der Polak’schen Expedition nach Persien im Jahre 1882. I. Theil. (Plantae collectae a Dr. J.E. Polak et. Th. Pichler). <em>Denkschriften der Kaiserlichen Akademie der Wissenschaften Mathematisch-Naturwissenschaftliche Classe</em> 50 (2): 1–355.</p>
<p>Huber-Morath, A. (1975) <em>Cousinia</em> Cass. <em>In</em>: Davis, P.H. (Ed.) <em>Flora of Turkey and the East Aegean Islands</em>, vol. 5. Edinburgh University Press, Edinburgh, pp. 329–353.</p>
<p>Jaubert, H.F. & Spach, E. (1850) <em>Illustrationes Plantarum Orientalium</em>, vol. 3. Roret, Paris. https://doi.org/10.1080/00378941.1856.10828752</p>
<p>Joudi, L., Habibi Bibalani, G.H. & Shadkami, H. (2011) Introduction to the medicinal species of Asteraceae family in Ilkhji and Sharafaldin regions of East Azarbaijan in Iran. <em>Journal of American Science</em> 7 (5): 455–458.</p>
<p>Kalouti, S., Kazempour Osaloo, S.H., Attar, F., Susanna, A. & Garcia-Jacas, N. (2021) Molecular phylogeny and taxonomy of species of <em>Cousinia </em>section <em>Stenocephalae</em> (Asteraceae) in NE of Iran. <em>Nova Biologica Reperta</em> 8 (1): 56–67.</p>
<p>Knapp, H.D. (1987) On the distribution of the genus <em>Cousinia </em>(Compositae). <em>Plant Systematics and Evolution</em> 155: 15–25. https://doi.org/10.1007/BF00936283</p>
<p>Köie, M.E. & Rechinger, K.H. (1955) <em>Symbolae Afghanicae</em>: Enumeration and Descriptions of the Plants Collected By L. Edelberg & M. Köie. <em>Biologiske Skrifter</em> 8 (2).</p>
<p>Kuntze, O. (1891) <em>Revisio generum plantarum</em>, vol. 1. A. Felix, Leipzig.</p>
<p>Kultiassov, M.V. (1928) <em>Cousinia syrdariensis</em> M. Kult. sp. n. <em>In</em>: Schedae ad Herbarium florae Asiae Mediae ab Universitate Asiae Mediae editum. Fasc. XIV–XX. Addenda et corrigenda. Index alphabeticus. <em>Trudy Sredne-Aziatskogo Gosudarstvennogo Universiteta</em>, ser. 8b, 3: 106.</p>
<p>Ledebour, C. von. (1829) <em>Icones plantarum novarum vel imperfecte cognitarum floram Rossicam, imprimis Altaicam, illustrantes</em>, vol. 1. I. Deubner, Riga. https://doi.org/10.5962/bhl.title.46626</p>
<p>Linnaeus, C. (1753) <em>Species Plantarum</em>, vol. 2. L. Salvius, Stockholm.</p>
<p>López-Vinyallonga, S., Mehregan, I., Garcia-Jacas, N., Tscherneva, O., Susanna, A. & Kadereit, J.W. (2009) Phylogeny and evolution of the <em>Arctium-Cousinia</em> complex (Compositae, Cardueae-Carduinae). <em>Taxon</em> 58: 153–171. https://doi.org/10.1002/tax.581016</p>
<p>López-Vinyallonga, S., Romaschenko, K., Susanna, A. & Garcia-Jacas, N. (2011) Systematics of the Arctioid group: Disentangling <em>Arctium </em>and <em>Cousinia </em>(Cardueae, Carduinae). <em>Taxon</em> 60: 539–554. https://doi.org/10.1002/tax.602020</p>
<p>Mehregan, I. & Kadereit, J.W. (2008) Taxonomic revision of <em>Cousinia</em> sect. <em>Cynaroideae</em> (Asteraceae, Cardueae). <em>Willdenowia</em> 38 (2): 293–362. https://doi.org/10.3372/wi.38.38201</p>
<p>Mehregan, I. & Kadereit, J.W. (2009) The role of hybridization in the evolution of <em>Cousinia </em>s.str. (Asteraceae, Cardueae). <em>Willdenowia</em> 39 (1): 35–47. https://doi.org/10.3372/wi.39.39102</p>
<p>Minaeifar, A.M., Sheidai, M., Attar, F., Noormohammadi, Z. & Ghasemzadeh-Baraki, B. (2015a) Genetic morphological diversity in <em>Cousinia tabrisiana</em> (Asteraceae) populations. <em>Biologia</em> 70 (3): 328–338. https://doi.org/10.1515/biolog-2015-0042</p>
<p>Minaeifar, A.M., Sheidai, M. & Attar, F. (2015b) Genetic and morphological diversity in <em>Cousinia cylindracea </em>(Asteraceae) populations: Identification of gene pools. <em>Biodiversitas</em> 16: 288–294. https://doi.org/10.13057/biodiv/d160225</p>
<p>Minaeifar, A., Sheidai, M., Attar, F., Noormohammadi, Z. & Ghasemzadeh-Baraki, S. (2016) Biosystematic study in the genus <em>Cousinia</em> Cass. (Asteraceae), section <em>Cousinia</em>. <em>Biochemical Systematics and Ecology</em> 69: 252–260. https://doi.org/10.1016/j.bse.2016.10.008</p>
<p>Noroozi, J., Akhani, H. & Breckle, S. (2007) Biodiversity and phytogeography of the alpine flora of Iran. <em>Biodiversity and Conservation</em> 17: 493–521. https://doi.org/10.1007/s10531-007-9246-7</p>
<p>Nylander, J.A.A. (2004) <em>MrModeltest</em> v2. Program distributed by the author. Evolutionary Biology Centre. Uppsala University, Uppsala.</p>
<p>Parsa, A. (1945) <em>Flore de l’Iran</em>, vol. 3. Tehran University Press, Tehran.</p>
<p>Posada, D. & Buckley, T.R. (2004) Model selection and model averaging in phylogenetics: advantages of AIC and Bayesian approaches over likelihood ratio tests. <em>Systematic Biology </em>53: 793–808. https://doi.org/10.1080/10635150490522304</p>
<p>Post, G.E. (1933) <em>Flora of Syria, Palestine and Sinai</em>, vol. 2. American University of Beirut, Beirut, pp. 84–87.</p>
<p>Rambaut, A. (2018) FigTree 1.4.4 Software. Institute of Evolutionary Biology, University of Edinburgh, Edinburgh.</p>
<p>Rastegar, A., Attar, F. & Mirtadzadini, M. (2018) Novelties in Iranian <em>Cousinia </em>sect. <em>Cynaroideae </em>(Asteraceae, Cardueae): new taxa and taxonomic notes. <em>Phytotaxa</em> 343: 227–239. https://doi.org/10.11646/phytotaxa.343.3.3</p>
<p>Rechinger, K.H. (1940) <em>Plantae novae iranicae</em> II. <em>Repertorium Specierum Novarum Regni Vegetabilis </em>48: 121–167.</p>
<p>Rechinger, K.H. (1950) <em>Cousiniae novae iranicae. (Rechingeri iter iranicum secundum</em><strong> -</strong> No. 11.). <em>Annalen des Naturhistorischen Museums in Wien</em> 57: 71–84.</p>
<p>Rechinger, K.H. (1972) Compositae-Cynareae І: <em>Cousinia</em>. <em>In</em>: Rechinger, K.H. (Ed.) <em>Flora Iranica,</em> vol. 90. Akad. Druck- und Verlagsanstalt, Graz, pp. 1–329. https://doi.org/10.2307/1218060</p>
<p>Rechinger, K.H. (1979) Compositae-Cynareae ІІІ:<em> Cousinia</em>. <em>In</em>: Rechinger, K.H. (Ed.) <em>Flora Iranica,</em> vol. 139a. Akad. Druck- und Verlagsanstalt, Graz, pp. 105–156.</p>
<p>Rechinger, K.H. (1986) <em>Cousinia</em>—morphology, taxonomy and phytogeographic implications. <em>Proceedings of the Royal Society of Edinburgh </em>89: 45–58. https://doi.org/10.1017/S0269727000008897</p>
<p>Ronquist, F., Teslenko, M., Mark, P., Ayres, D., Darling, D., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. <em>Systematic Biology</em> 61: 539–542. https://doi.org/10.1093/sysbio/sys029</p>
<p>Ruprecht, F.J. (1869) Sertum Tianschanicum. Botanische Ergebnisse einer Reise im mittleren Tian-Schan. <em>Mémoires de l’Académie Impériale des Sciences de Saint Pétersbourg</em>, sér. 7, 14 (4): 33–74.</p>
<p>Saber, A. (2005) <em>Biosystematical studies on the Cousinia section Stenocephalae (Asteraceae) in Iran.</em> M.Sc. Thesis, University of Tehran.</p>
<p>Saber, A., Attar, F. & Djavadi, S.B. (2009) Studies of pollen grains in the sect. <em>Stenocephalae</em> (<em>Cousinia</em> Cass., Asteraceae) in Iran. <em>Iranian Journal of Botany</em> 15 (1): 114–128.</p>
<p>Sajjadi, S.E., Ghanadian, M., Haghighi, M. & Mouhebat, L. (2015) Cytotoxic effect of <em>Cousinia verbascifolia</em> Bunge against OVCAR-3 and HT-29 cancer cells. <em>HerbMed Pharmacology</em> 4 (1): 15–19.</p>
<p>Sang, T., Crawford, D.J. & Stuessy, T.F. (1995) Documentation of reticulate evolution in peonies (<em>Paeonia</em>) using internal transcribed spacer sequences of nuclear ribosomal DNA: implications for biogeography and concerted evolution. <em>Proceedings of the</em> <em>National Academy of Sciences of the United States of America</em> 92 (15): 6813–6817. https://doi.org/10.1073/pnas.92.15.6813</p>
<p>Sennikov, A.N. (2010) A revision of <em>Cousinia </em>sections <em>Alpinae </em>(syn. <em>Carduncellus</em>), <em>Subappendiculatae </em>and <em>Tianschanicae </em>(Asteraceae) in the Kirghizian Tian-Shan and the neighbouring territories<em>. Phytotaxa</em> 5: 1–30. https://doi.org/10.11646/phytotaxa.5.1.1</p>
<p>Shahverdi, A.R., Khorramizadeh, M.R. & Attar, A. (2007a) Chemopreventive effect of <em>Cousinia shulabadensis</em> ethanol extract. <em>African Journal of Traditional, Complementary and Alternative Medicines</em> 4 (1): 12–16. https://doi.org/10.4314/ajtcam.v4i1.31186</p>
<p>Shahverdi, A.R., Khorramizadeh, M.R., Attar, F., Saadat, F., Vahid, S. & Ghahraman, A. (2007b) Concomitant chemopreventive and antibacterial effects of some Iranian plants from the genus <em>Cousinia</em> (Asteraceae). <em>Brazilian Journal of Pharmacognosy</em> 17: 325–330. https://doi.org/10.1590/S0102-695X2007000300004</p>
<p>Sheidai, M., Ahmad-Khanbeygi, Z. & Attar, F. (2012) New chromosome number reports in <em>Cousinia </em>species (Compositae). <em>Cytologia</em> 77 (1): 11–16. https://doi.org/10.1508/cytologia.77.11</p>
<p>Susanna, A. & Garcia-Jacas, N. (2007) Tribe Cardueae. <em>In</em>: Kadereit, J.W. & Jeffrey, C. (Eds.) <em>The families and genera of vascular plants</em>, vol. 8. Springer, Berlin & Heidelberg, pp. 123–146.</p>
<p>Susanna, A., Garcia-Jacas, N., Hidalgo, O., Vilatersana, R. & Garnatje, T. (2006) The Cardueae (Compositae) revisited: Insights from ITS, <em>trnL-trnF</em>, and <em>matK </em>nuclear and chloroplast DNA analysis. <em>Annals of the Missouri Botanical Garden </em>93: 150–171. https://doi.org/10.3417/0026-6493(2006)93[150:TCCRIF]2.0.CO;2</p>
<p>Susanna, A., Garcia-Jacas, N., Vilatersana, R. & Garnatje, T. (2003a) Generic boundaries and evolution of characters in the <em>Arctium </em>group: A nuclear and chloroplast DNA analysis. <em>Collectanea Botanica </em>26: 101–118. https://doi.org/10.3989/collectbot.2003.v26.17</p>
<p>Susanna, A., Garcia-Jacas, N., Vilatersana, R., Garnatje, T., Vallès, J. & Ghaffari, S.M. (2003b) New chromosome counts in the genus <em>Cousinia </em>and the related genus <em>Schmalhausenia </em>(Asteraceae, Cardueae). <em>Botanical Journal of the Linnean Society</em> 43: 411–418. https://doi.org/10.1111/j.1095-8339.2003.00231.x</p>
<p>Tareen, R.B., Bibi, T., Ajab Khan, M., Ahmad, M. & Zafar, M. (2010) Indigenous knowledge of folk medicine by the women of Kalat and Khuzdar regions of Balochistan, Pakistan. <em>Pakistan Journal of Botany</em> 42: 1456–1485.</p>
<p>Tscherneva, O.V. (1962) <em>Cousinia </em>Cass. <em>In</em>: Shishkin, B.K. (Ed.) <em>Flora of the USSR</em>, vol. 27. Academy of Sciences of the USSR, Moscow & Leningrad, pp. 108–357. [In Russian]</p>
<p>White, J.T., Bruns, T.D., Lee, S.T. & Taylor, J.W. (1990) Amplification and direct squencing of fungal ribosomal RNA genes for phylogenetics. <em>In</em>: Innis, D.H. (Ed.) <em>PCR protocols: a guide to methods and applications</em>. Academic Press, San Diego, pp. 315–322. https://doi.org/10.1016/B978-0-12-372180-8.50042-1</p>
<p>Zare, M., Khossravi, A.R. & Joharchi, M.R. (2013) Distribution patterns of the genus <em>Cousinia</em> (Asteraceae) in Iran. <em>Iranian Journal of Botany</em> 19 (1): 127–141.</p>
<p>Attar, F. (2011a) Taxonomic revision of <em>Cousinia</em> sect. <em>Albidae</em> Bunge (Asteraceae) in the flora of Iran. <em>Iranian Journal of Botany</em> 17 (2): 150–157.</p>
<p>Attar, F. (2011b) New taxa of <em>Cousinia </em>Cass. sect. <em>Stenocephalae </em>Bunge (Asteraceae) from Iran. <em>Taxonomy and Biosystematics</em> 6: 59–68.</p>
<p>Attar, F. & Djavadi, S.B. (2010) A taxonomic revision of<em> Cousinia</em> sect. <em>Cynaroides</em> (Asteraceae, Cardueae) in the flora of Iran. <em>Iranian Journal of Botany</em> 16 (1): 130–184.</p>
<p>Attar, F., Ghahreman, A., Saber, A. & Zarre, Sh. (2005) A new species of the genus <em>Cousinia </em>(Compositae, sect. <em>Stenocephalae</em>) from SW Iran. <em>Iranian Journal of Botany</em> 11 (1): 65– 69.</p>
<p>Attar, F. & Maroofi, H. (2010) <em>Cousinia</em> (sect. <em>Albidae</em>) <em>Iranshahriana</em> (Asteraceae, Cardueae), a new endemic species from Kurdistan province, Iran. <em>Iranian Journal of Botany</em> 16 (1): 197–199.</p>
<p>Bahraminejad, S., Amiri, R., Ghasemi, S. & Fathi, N. (2013) Inhibitory effect of some Iranian plant species against three plant pathogenic fungi. <em>International Journal of Agriculture and Crop Sciences</em> 5: 1002–1008.</p>
<p>Boissier, E. (1845) <em>Diagnosis plantarum orientalium novarum, ser. 1</em>, vol. 6. B. Hermann, Lipsiae [Leipzig].</p>
<p>Boissier, E. & Buhse, F.A. (1860) Aufzählung der auf einer Reise durch Transkaukasien und Persien gesammelten Pflanzen. <em>Nouveaux</em> <em>Meìmoires de la Socieìteì Impeìriale des Naturalistes de Moscou</em> 12: 1–246.</p>
<p>Boissier, E. (1875) <em>Flora Orientalis</em>. H. Georg, Geneva & Basilea [Basel].</p>
<p>Bordbar, F., Baqeri, M. & Mirtadzadini, M. (2017) New Chromosome Counts in <em>Cousinia</em> (Asteraceae)from Flora of Iran. <em>Cytologia</em>. 82 (5): 485–488.</p>
<p>Bornmüller, J.F.N. (1907) Beiträge zur Flora der Elbursgebirge Nord-Persiens. <em>Bulletin de l’Herbier Boissier</em>, ser. 2, 7: 209–224.</p>
<p>Bornmüller, J.F.N (1910) Bearbeitung der von J.A. Knapp im nordwestlichen Persien gesammelten Pflanzen. <em>Verhandlungen der Zoologisch-Botanischen Gesellschaft</em> <em>in Wien</em> 60: 1–512.</p>
<p>Bornmüller, J. (1911) Collectiones Straussianae novae. Weitere Beiträge zur Kenntnis der Flora West Persiens. <em>Beihefte zum Botanischen Centralblatt, Original Arbeiten</em> 28 (2): 225–267.</p>
<p>Bornmüller, J.F.N. (1912) Ein Beitrag zur Kenntnis der Gattung <em>Cousinia. Oesterreichische Botanische Zeitschrift</em> 62: 1–523. https://doi.org/10.1007/BF01632329</p>
<p>Bornmüller, J.F.N. (1916) Ein Beitrag zur Kenntnis der Gattung <em>Cousinia</em>. <em>Beihefte zum Botanischen Centralblatt</em>,<em> Zweite Abt.</em> 34: 131–203.</p>
<p>Bornmüller, J.F.N. (1934) Ein Beitrag zur Kenntnis der Gattung <em>Cousinia</em>. XII. Neue Arten und bemerkenswerte Formen aus Persien und Turkestan.<em> Repertorium Specierum Novarum Regni Vegetabilis</em> 36: 323–338.</p>
<p>Bornmüller, J.F.N. (1941) Neue Arten der Gattung <em>Cousinia</em> der Flora Persiens. <em>Mittheilungen des Thüringischen Botanischen Vereins</em> 47: 115–128.</p>
<p>Bunge, A. von (1841). Correspondenz.<em> Flora oder Botanische Zeitung </em>24 (1): 153–159.</p>
<p>Bunge, A. von (1865) Übersichtliche Zusammenstellung der Arten der Gattung <em>Cousinia</em> Cass. <em>Mémoires de l’Académie Impériale de Sciences de Saint Pétersbourg</em>, Ser. 7, 9: 1–56.</p>
<p>Cassini, A.H.G. (1827) <em>Cousinia</em>. <em>In</em>: Cuvier, G. (Ed.) <em>Dictionnaire des sciences naturelles</em>, vol. 47. Levrault, Strasbourg, pp. 503–505.</p>
<p>Candolle, A.P. de (1838) <em>Prodromus systematis naturalis regni vegetabilis,</em> vol. 6. Treuttel & Würtz, Paris.</p>
<p>Djamali, M., Baumel, A., Brewer, S., Jackson, S.T, Kadereit, J.W., López-Vinyallonga, S., Mehregan, I., Shabanian, E. & Simakova, A. (2012) Ecological implications of <em>Cousinia </em>Cass. (Asteraceae) persistence through the last two glacial–interglacial cycles in the continental Middle East for the Irano-Turanian flora. <em>Review of Palaeobotany and Palynology</em> 172: 10–20. https://doi.org/10.1016/j.revpalbo.2012.01.005</p>
<p>Djavadi, S.B. (2005) New or rare chromosome count in ten species of <em>Cousinia</em> from Iran. <em>Rostaniha </em>8 (1): 19–27.</p>
<p>Djavadi, S.B. (2012) New chromosome counts in <em>Cousinia</em> (Asteraceae, Cardueae) section <em>Cynaroideae </em>and <em>Stenocephalae</em> from Iran. <em>Iranian Journal of Botany</em> 18 (1): 86–93.</p>
<p>Djavadi, S.B. & Attar, F. (2006) Two new species of <em>Cousinia</em> Cass. of sect. <em>Stenocephalae</em> Bunge from Iran. <em>Feddes Repertorium</em> 117: 453–458. https://doi.org/10.1002/fedr.200611113</p>
<p>Djavadi, S.B. & Attar, F. (2010) New chromosome counts in the genus <em>Cousinia</em> (Asteraceae, Cardueae) from Iran. <em>Willdenowia</em> 40 (2): 351–357. https://doi.org/10.3372/wi.40.40214</p>
<p>Djavadi, S.B., Attar, F. & Najafi, Z. (2007) A new species of <em>Cousinia</em> sect. <em>Cousinia</em> from NW Iran. <em>Iranian Journal of Botany</em> 13 (1): 43–46.</p>
<p>Doyle, J.J. & Doyle, J.L. (1987) A rapid DNA isolation procedure for small quantities of fresh leaf tissue. <em>Phytochemical Bulletin</em> 19: 11–15.</p>
<p>Duistermaat, H. (1996) <em>Arctium</em> getting entangled to <em>Cou</em>s<em>inia</em> (Asteraceae: Cardueae). <em>Bocconea</em> 5: 685–689.</p>
<p>Edgar, R.C. (2004) Muscle: multiple sequence alignment with high accuracy and high throughput. <em>Nucleic Acids Research </em>32: 1792–1797. https://doi.org/10.1093/nar/gkh340</p>
<p>Ghaffari, S.M. & Chariat-Panahi, S. (1985) Chromosome count of some angiosperms from Iran. <em>Iranian Journal of Botany</em> 3 (1): 67–73.</p>
<p>Ghaffari, S.M. & Djavadi, S.B. (1998) Chromosome studies and distribution of nine species of <em>Cousinia</em> section <em>Stenocephalae</em> (Asteraceae) in Iran.<em> Bulletin de la Société Neuchâteloise des Sciences Naturelles</em> 121: 61–68.</p>
<p>Ghaffari, S.M., Susanna, A. & Garcia-Jacas, N. (2006) New chromosome counts in the genus <em>Cousinia </em>(Asteraceae) from Iran. <em>Botanical Journal of the Linnean Society </em>151: 411–419. https://doi.org/10.1111/j.1095-8339.2006.00506.x</p>
<p>Herder, F. von (1869) Plantae raddeanae monopetalae. <em>Bulletin de la Société Impériale des Naturalistes de Moscou</em> 41 (3): 1–53.</p>
<p>Herrando-Moraira, S., The Cardueae Radiations Group (Calleja, J.A., Galbany-Casals, M., Garcia-Jacas, N., Liu, J.-Q., López-Alvarado, J., López-Pujol, J., Mandel, J.R., Massó, S., Montes-Moreno, N., Roquet, C., Sáez, L., Sennikov, A., Susanna, A. & Vilatersana, R.) (2019) Nuclear and plastid DNA phylogeny of tribe Cardueae (Compositae) with Hyb-Seq data: A new subtribal classification and a temporal diversification framework. <em>Molecular Phylogenetics and Evolution</em> 137: 313–332. https://doi.org/10.1016/j.ympev.2019.05.001</p>
<p>Hiemeral, A. (1885) <em>Cousinia</em> Cass. <em>In: </em>Stapf, O., Die botanischen Ergebnisse der Polak’schen Expedition nach Persien im Jahre 1882. I. Theil. (Plantae collectae a Dr. J.E. Polak et. Th. Pichler). <em>Denkschriften der Kaiserlichen Akademie der Wissenschaften Mathematisch-Naturwissenschaftliche Classe</em> 50 (2): 1–355.</p>
<p>Huber-Morath, A. (1975) <em>Cousinia</em> Cass. <em>In</em>: Davis, P.H. (Ed.) <em>Flora of Turkey and the East Aegean Islands</em>, vol. 5. Edinburgh University Press, Edinburgh, pp. 329–353.</p>
<p>Jaubert, H.F. & Spach, E. (1850) <em>Illustrationes Plantarum Orientalium</em>, vol. 3. Roret, Paris. https://doi.org/10.1080/00378941.1856.10828752</p>
<p>Joudi, L., Habibi Bibalani, G.H. & Shadkami, H. (2011) Introduction to the medicinal species of Asteraceae family in Ilkhji and Sharafaldin regions of East Azarbaijan in Iran. <em>Journal of American Science</em> 7 (5): 455–458.</p>
<p>Kalouti, S., Kazempour Osaloo, S.H., Attar, F., Susanna, A. & Garcia-Jacas, N. (2021) Molecular phylogeny and taxonomy of species of <em>Cousinia </em>section <em>Stenocephalae</em> (Asteraceae) in NE of Iran. <em>Nova Biologica Reperta</em> 8 (1): 56–67.</p>
<p>Knapp, H.D. (1987) On the distribution of the genus <em>Cousinia </em>(Compositae). <em>Plant Systematics and Evolution</em> 155: 15–25. https://doi.org/10.1007/BF00936283</p>
<p>Köie, M.E. & Rechinger, K.H. (1955) <em>Symbolae Afghanicae</em>: Enumeration and Descriptions of the Plants Collected By L. Edelberg & M. Köie. <em>Biologiske Skrifter</em> 8 (2).</p>
<p>Kuntze, O. (1891) <em>Revisio generum plantarum</em>, vol. 1. A. Felix, Leipzig.</p>
<p>Kultiassov, M.V. (1928) <em>Cousinia syrdariensis</em> M. Kult. sp. n. <em>In</em>: Schedae ad Herbarium florae Asiae Mediae ab Universitate Asiae Mediae editum. Fasc. XIV–XX. Addenda et corrigenda. Index alphabeticus. <em>Trudy Sredne-Aziatskogo Gosudarstvennogo Universiteta</em>, ser. 8b, 3: 106.</p>
<p>Ledebour, C. von. (1829) <em>Icones plantarum novarum vel imperfecte cognitarum floram Rossicam, imprimis Altaicam, illustrantes</em>, vol. 1. I. Deubner, Riga. https://doi.org/10.5962/bhl.title.46626</p>
<p>Linnaeus, C. (1753) <em>Species Plantarum</em>, vol. 2. L. Salvius, Stockholm.</p>
<p>López-Vinyallonga, S., Mehregan, I., Garcia-Jacas, N., Tscherneva, O., Susanna, A. & Kadereit, J.W. (2009) Phylogeny and evolution of the <em>Arctium-Cousinia</em> complex (Compositae, Cardueae-Carduinae). <em>Taxon</em> 58: 153–171. https://doi.org/10.1002/tax.581016</p>
<p>López-Vinyallonga, S., Romaschenko, K., Susanna, A. & Garcia-Jacas, N. (2011) Systematics of the Arctioid group: Disentangling <em>Arctium </em>and <em>Cousinia </em>(Cardueae, Carduinae). <em>Taxon</em> 60: 539–554. https://doi.org/10.1002/tax.602020</p>
<p>Mehregan, I. & Kadereit, J.W. (2008) Taxonomic revision of <em>Cousinia</em> sect. <em>Cynaroideae</em> (Asteraceae, Cardueae). <em>Willdenowia</em> 38 (2): 293–362. https://doi.org/10.3372/wi.38.38201</p>
<p>Mehregan, I. & Kadereit, J.W. (2009) The role of hybridization in the evolution of <em>Cousinia </em>s.str. (Asteraceae, Cardueae). <em>Willdenowia</em> 39 (1): 35–47. https://doi.org/10.3372/wi.39.39102</p>
<p>Minaeifar, A.M., Sheidai, M., Attar, F., Noormohammadi, Z. & Ghasemzadeh-Baraki, B. (2015a) Genetic morphological diversity in <em>Cousinia tabrisiana</em> (Asteraceae) populations. <em>Biologia</em> 70 (3): 328–338. https://doi.org/10.1515/biolog-2015-0042</p>
<p>Minaeifar, A.M., Sheidai, M. & Attar, F. (2015b) Genetic and morphological diversity in <em>Cousinia cylindracea </em>(Asteraceae) populations: Identification of gene pools. <em>Biodiversitas</em> 16: 288–294. https://doi.org/10.13057/biodiv/d160225</p>
<p>Minaeifar, A., Sheidai, M., Attar, F., Noormohammadi, Z. & Ghasemzadeh-Baraki, S. (2016) Biosystematic study in the genus <em>Cousinia</em> Cass. (Asteraceae), section <em>Cousinia</em>. <em>Biochemical Systematics and Ecology</em> 69: 252–260. https://doi.org/10.1016/j.bse.2016.10.008</p>
<p>Noroozi, J., Akhani, H. & Breckle, S. (2007) Biodiversity and phytogeography of the alpine flora of Iran. <em>Biodiversity and Conservation</em> 17: 493–521. https://doi.org/10.1007/s10531-007-9246-7</p>
<p>Nylander, J.A.A. (2004) <em>MrModeltest</em> v2. Program distributed by the author. Evolutionary Biology Centre. Uppsala University, Uppsala.</p>
<p>Parsa, A. (1945) <em>Flore de l’Iran</em>, vol. 3. Tehran University Press, Tehran.</p>
<p>Posada, D. & Buckley, T.R. (2004) Model selection and model averaging in phylogenetics: advantages of AIC and Bayesian approaches over likelihood ratio tests. <em>Systematic Biology </em>53: 793–808. https://doi.org/10.1080/10635150490522304</p>
<p>Post, G.E. (1933) <em>Flora of Syria, Palestine and Sinai</em>, vol. 2. American University of Beirut, Beirut, pp. 84–87.</p>
<p>Rambaut, A. (2018) FigTree 1.4.4 Software. Institute of Evolutionary Biology, University of Edinburgh, Edinburgh.</p>
<p>Rastegar, A., Attar, F. & Mirtadzadini, M. (2018) Novelties in Iranian <em>Cousinia </em>sect. <em>Cynaroideae </em>(Asteraceae, Cardueae): new taxa and taxonomic notes. <em>Phytotaxa</em> 343: 227–239. https://doi.org/10.11646/phytotaxa.343.3.3</p>
<p>Rechinger, K.H. (1940) <em>Plantae novae iranicae</em> II. <em>Repertorium Specierum Novarum Regni Vegetabilis </em>48: 121–167.</p>
<p>Rechinger, K.H. (1950) <em>Cousiniae novae iranicae. (Rechingeri iter iranicum secundum</em><strong> -</strong> No. 11.). <em>Annalen des Naturhistorischen Museums in Wien</em> 57: 71–84.</p>
<p>Rechinger, K.H. (1972) Compositae-Cynareae І: <em>Cousinia</em>. <em>In</em>: Rechinger, K.H. (Ed.) <em>Flora Iranica,</em> vol. 90. Akad. Druck- und Verlagsanstalt, Graz, pp. 1–329. https://doi.org/10.2307/1218060</p>
<p>Rechinger, K.H. (1979) Compositae-Cynareae ІІІ:<em> Cousinia</em>. <em>In</em>: Rechinger, K.H. (Ed.) <em>Flora Iranica,</em> vol. 139a. Akad. Druck- und Verlagsanstalt, Graz, pp. 105–156.</p>
<p>Rechinger, K.H. (1986) <em>Cousinia</em>—morphology, taxonomy and phytogeographic implications. <em>Proceedings of the Royal Society of Edinburgh </em>89: 45–58. https://doi.org/10.1017/S0269727000008897</p>
<p>Ronquist, F., Teslenko, M., Mark, P., Ayres, D., Darling, D., Höhna, S., Larget, B., Liu, L., Suchard, M.A. & Huelsenbeck, J. (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. <em>Systematic Biology</em> 61: 539–542. https://doi.org/10.1093/sysbio/sys029</p>
<p>Ruprecht, F.J. (1869) Sertum Tianschanicum. Botanische Ergebnisse einer Reise im mittleren Tian-Schan. <em>Mémoires de l’Académie Impériale des Sciences de Saint Pétersbourg</em>, sér. 7, 14 (4): 33–74.</p>
<p>Saber, A. (2005) <em>Biosystematical studies on the Cousinia section Stenocephalae (Asteraceae) in Iran.</em> M.Sc. Thesis, University of Tehran.</p>
<p>Saber, A., Attar, F. & Djavadi, S.B. (2009) Studies of pollen grains in the sect. <em>Stenocephalae</em> (<em>Cousinia</em> Cass., Asteraceae) in Iran. <em>Iranian Journal of Botany</em> 15 (1): 114–128.</p>
<p>Sajjadi, S.E., Ghanadian, M., Haghighi, M. & Mouhebat, L. (2015) Cytotoxic effect of <em>Cousinia verbascifolia</em> Bunge against OVCAR-3 and HT-29 cancer cells. <em>HerbMed Pharmacology</em> 4 (1): 15–19.</p>
<p>Sang, T., Crawford, D.J. & Stuessy, T.F. (1995) Documentation of reticulate evolution in peonies (<em>Paeonia</em>) using internal transcribed spacer sequences of nuclear ribosomal DNA: implications for biogeography and concerted evolution. <em>Proceedings of the</em> <em>National Academy of Sciences of the United States of America</em> 92 (15): 6813–6817. https://doi.org/10.1073/pnas.92.15.6813</p>
<p>Sennikov, A.N. (2010) A revision of <em>Cousinia </em>sections <em>Alpinae </em>(syn. <em>Carduncellus</em>), <em>Subappendiculatae </em>and <em>Tianschanicae </em>(Asteraceae) in the Kirghizian Tian-Shan and the neighbouring territories<em>. Phytotaxa</em> 5: 1–30. https://doi.org/10.11646/phytotaxa.5.1.1</p>
<p>Shahverdi, A.R., Khorramizadeh, M.R. & Attar, A. (2007a) Chemopreventive effect of <em>Cousinia shulabadensis</em> ethanol extract. <em>African Journal of Traditional, Complementary and Alternative Medicines</em> 4 (1): 12–16. https://doi.org/10.4314/ajtcam.v4i1.31186</p>
<p>Shahverdi, A.R., Khorramizadeh, M.R., Attar, F., Saadat, F., Vahid, S. & Ghahraman, A. (2007b) Concomitant chemopreventive and antibacterial effects of some Iranian plants from the genus <em>Cousinia</em> (Asteraceae). <em>Brazilian Journal of Pharmacognosy</em> 17: 325–330. https://doi.org/10.1590/S0102-695X2007000300004</p>
<p>Sheidai, M., Ahmad-Khanbeygi, Z. & Attar, F. (2012) New chromosome number reports in <em>Cousinia </em>species (Compositae). <em>Cytologia</em> 77 (1): 11–16. https://doi.org/10.1508/cytologia.77.11</p>
<p>Susanna, A. & Garcia-Jacas, N. (2007) Tribe Cardueae. <em>In</em>: Kadereit, J.W. & Jeffrey, C. (Eds.) <em>The families and genera of vascular plants</em>, vol. 8. Springer, Berlin & Heidelberg, pp. 123–146.</p>
<p>Susanna, A., Garcia-Jacas, N., Hidalgo, O., Vilatersana, R. & Garnatje, T. (2006) The Cardueae (Compositae) revisited: Insights from ITS, <em>trnL-trnF</em>, and <em>matK </em>nuclear and chloroplast DNA analysis. <em>Annals of the Missouri Botanical Garden </em>93: 150–171. https://doi.org/10.3417/0026-6493(2006)93[150:TCCRIF]2.0.CO;2</p>
<p>Susanna, A., Garcia-Jacas, N., Vilatersana, R. & Garnatje, T. (2003a) Generic boundaries and evolution of characters in the <em>Arctium </em>group: A nuclear and chloroplast DNA analysis. <em>Collectanea Botanica </em>26: 101–118. https://doi.org/10.3989/collectbot.2003.v26.17</p>
<p>Susanna, A., Garcia-Jacas, N., Vilatersana, R., Garnatje, T., Vallès, J. & Ghaffari, S.M. (2003b) New chromosome counts in the genus <em>Cousinia </em>and the related genus <em>Schmalhausenia </em>(Asteraceae, Cardueae). <em>Botanical Journal of the Linnean Society</em> 43: 411–418. https://doi.org/10.1111/j.1095-8339.2003.00231.x</p>
<p>Tareen, R.B., Bibi, T., Ajab Khan, M., Ahmad, M. & Zafar, M. (2010) Indigenous knowledge of folk medicine by the women of Kalat and Khuzdar regions of Balochistan, Pakistan. <em>Pakistan Journal of Botany</em> 42: 1456–1485.</p>
<p>Tscherneva, O.V. (1962) <em>Cousinia </em>Cass. <em>In</em>: Shishkin, B.K. (Ed.) <em>Flora of the USSR</em>, vol. 27. Academy of Sciences of the USSR, Moscow & Leningrad, pp. 108–357. [In Russian]</p>
<p>White, J.T., Bruns, T.D., Lee, S.T. & Taylor, J.W. (1990) Amplification and direct squencing of fungal ribosomal RNA genes for phylogenetics. <em>In</em>: Innis, D.H. (Ed.) <em>PCR protocols: a guide to methods and applications</em>. Academic Press, San Diego, pp. 315–322. https://doi.org/10.1016/B978-0-12-372180-8.50042-1</p>
<p>Zare, M., Khossravi, A.R. & Joharchi, M.R. (2013) Distribution patterns of the genus <em>Cousinia</em> (Asteraceae) in Iran. <em>Iranian Journal of Botany</em> 19 (1): 127–141.</p>