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