Abstract
Bromus (Poaceae) includes more than 160 species of mesophytic and xerophytic grasses from temperate regions of the world, of which 39 occur in Iran. The genus is considered taxonomically problematic due to the occurrence of hybridization, variation in ploidy levels, and phenotypic plasticity. Moreover, morphological similarities make species delimitation difficult. It resulted in different taxonomic classifications proposed along the time. Thus, in order to contribute to a better understanding of the species of Bromus, 21 species from six different sections of Bromus occurring in Iran were evaluated micromorphologically. The abaxial surface of the lemma and palea of the lowermost florets were examined using Scanning Electron Microscopy (SEM). Twenty-three quantitative and qualitative characters were analysed using PAST software. Lemma characters such as shape of silica cells, cork cells, crown cells, outline of long cell wall, and long cell length, and palea characters such as shape of cork cells, silica cells, and outline of long cell wall were informative among species studied. In UPGMA clustering most species were aggregated according to the previously delimited sections demonstrating the taxonomic value of micromorphological features in the species studied. Two sections Bromus and Genea showed a close relationship whereas two sections Ceratochloa and Pnigma were placed in one cluster close to the cluster including section Boissiera. The only exception was B. japonicus which was resolved within cluster of sect. Genea, not in cluster of sect. Bromus.
References
Acedo, C. & Llamas, F. (2001) Variation of micromorphological characters of lemma and palea in the genus Bromus (Poaceae). Annales Botanici Fennici 38: 1–14.
Ainouche, M.L. & Bayer, R.J. (1997) On the origins of the tetraploid Bromus species (section Bromus, Poaceae): insights from internal transcribed spacer sequences of nuclear ribosomal DNA. Genome 40: 730–743. http://doi.org/10.1139/g97-796
Armstrong, K.C. (1981) Evolution of Bromus inermis and related species of Bromus sect. Pnigma. Botanische Jahrbucher fur Systematik, Pflanzengeschichte und Pflanzengeographie 102: 427–443.
Barthlott, W., Neinhuis, Ch., Cutler, D., Ditsch, F., Meusel, I., Theisen, I. & Wilhelmi, H. (1998) Classification and terminology of plant epicuticular waxes. Botanical Journal of the Linnean Society 126: 237–260. https://doi.org/10.1111/j.1095-8339.1998.tb02529.x
Boissier, E. (1846) Diagnoses Plantarum Orientalium novarum. ser 1, vol. 7. Typographia Ferd. Ramboz, Genevae, 781 pp.
Boissier, E. (1884) Bromus. In: Boissier, E (Eds.) Flora Orientalis: sive, Enumeratio plantarum in Oriente a Graecia et Aegypto ad Indiae fines hucusque observatarum. vol. 5. H. Georg, Geneva and Basel, pp. 640–656. https://doi.org/10.5962/bhl.title.20323
Bor, N.L (1970) Bromeae. In: Rechinger, K.H. (Ed.) Flora Iranica, vol. 70. Akademische Druck- u. Verlagsanstalt, Graz, pp. 105–141.
Bor, N.L. (1968) Bromeae. In: Townsend, C.C., Guest, E. & Al-Rawi A (Eds.) Flora of Iraq, vol 9. Ministry of Agriculture and Agrarian Reform, Baghdad, pp. 126–166.
Bunge, A.A. von (1851) Beitrag zur Kenntniss der Flora Russlands und der Steppen Central-Asiens. Mémoires Presentes a l’Académie Impériale des Sciences de St.-Pétersbourg par Divers Savans et lus dans ses Assemblées 7: 177–535.
Clayton, W.D. & Renvoize, S.A. (1986) Genera Graminum: Grasses of the World. Her Majesty’s Stationary Office, London, 389 pp.
Cope, Th.A. (1982) Poaceae. In: Nasir, E. & Ali, S.I. (Eds.) Flora of Pakistan, vol 143. Pakistan Agricultural Research Council, Islamabad, pp. 1–678.
de Candolle, A.P. & Palisot de Beauvois, A.M.F.J. (1812) Essai d’une nouvelle agrostographie; ou nouveaux genres des graminées; avec figures représentant les caractéres de tous le genres. Imprimerie de Fain, Paris, 182 pp. http://dx.doi.org/10.5962/bhl.title.474.
Drobow, V. P. (1925) Gramineae novae turkestanicae.I. Repertorium specierum novarum regni vegetabilis 21: 37–46. https://doi.org/10.1002/fedr.19250210106
Dumortier, B.C.J. (1823 [1824]) Observations sur les Graminées de la Flora Belgique. J. Casterman, Tournay, 116 pp.
Ellis, R.P. (1979) A procedure for standardizing comparative leaf anatomy in the Poaceae. II. The epidermis as seen in surface view. Bothalia 12: 641–671. https://doi.org/10.4102/abc.v12i4.1441
Figari, A. B., De Notaris, J. (1852) Agrostographiae Aegyptiacae fragmenta. Memorie della Reale Accademia delle Scienze di Torino 2 (12): 245–262.
Fischer, F.E.L., Meyer, C.A., Trautvetter, E.R. & Avé-Lallemant, J.L.E. (1837) Animadversiones botanicae. Index seminum, quae Hortus Botanicus imperialis petropolitanus pro mutua commutatione offert. Accedunt animadversiones botanicae nonnullae 3: 1–19.
Fortune, P.M., Pourtau, N., Viron, N. & Ainouche, M.L. (2008) Molecular phylogeny and reticulate origins of the polyploidy Bromus species from section Genea (Poaceae). American Journal of Botany 95: 456–464.
Ghahreman, A., Alemi, M., Attar, F., Hamzehee, B. & Columbus, J.T. (2006) Anatomical studies in some species of Bromus (Poaceae) in Iran. Iranian Journal of Botany 12: 1–14.
Hackel, E. (1887) Gramineae. In: Engler, A. & Prantel, K. A. (Eds.) Naturlichen Panzenfamilien. (Abt. 2, Lief. 12). W. Engelmann, Leipzig, pp. 1–130. https://doi.org/10.5962/bhl.title.4635
Hackel, E. (1903) Gramineae. Videnskabelige Meddelelser fra den Naturhistoriske Forening i Kjøbenhavn 161–178.
Hammer, Ø., Harper, D. & Ryan, P.D. (2001) PAST: Paleontological Statistics software package for education and data analysis. Palaeontologia Electronica 4 (1): 1–9.
Hemsley, W.B. (1896) Hooker’s Icones Plantarum; or figures, with brief descriptive characters and remarks of new or rare plants, 25: t. London, 210 pp. https://doi.org/10.5962/bhl.title.16059
Hitchcock, A.S. (1935) Manual of the grasses of the United States. United States Department of Agriculture Miscellaneous Publication no. 200. United States Government Printing Office, Washington, D.C., 1040 pp.
Hochstetter, Ch.F.F. & Steudel, E. G. (1838) Einige nähere Bestimmungen der ägyptische arabischen Pflanzen. Flora oder Allgemeine Botanische Zeitung 21 (1): 23–27.
Huttanus, T.D., Mack, R.N., Novak, S.J. (2011) Propagule pressure and introduction pathways of Bromus tectorum (Cheat grass; Poaceae) in the Central United States. International Journal of Plant Sciences 172: 783–794. https://doi.org/10.1086/660107
Keshavarzi, M., Direkvandi, S. & Khoshnood, N. (2015) Chromosome number variation in Bromus L. (Poaceae) in Khorassan and Lorestan. Journal of Plant Research 28: 826–834.
Klimko, M. & Wysakowska, I. (2015) Epidermal features of glumes and florets in Aegilops geniculata Roth and Aegilops peregrina (Hack.) Maire et weiller × Secale cereal L. hybrids, amphiploids and parental forms. Steciana 19 (1): 13–24. https://doi.org/10.12657/STECIANA.019.003
Kosina, R. (1999) Patterns of flower microstructural variation within the genus Bromus. Acta Societatis Botanicorum Poloniae 68 (3): 221–226.
Leandro, T.D., Shirasuna, R.T., Filgueiras, T.S. & Scatena, V.L. (2016) The utility of Bambusoideae (Poaceae, Poales) leaf blade anatomy for identification and systematics. Brazilian Journal of Biology 76: 708–717. http://dx.doi.org/10.1590/1519-6984.01715
Ledebour, C.F.von. (1852) Flora rossica sive enumeratio plantarum in totius imperii rossici provinciis europaeis, asiaticis, et americanis hucusque observatarum 4, part 13. Sumptibus librariae E. Schweizerbart, Stuttgartiae, pp. 241–464. http://dx.doi.org/10.5962/bhl.title.6606
Liang, L., Shouliang, Ch., Guanghu, Zh., Ammann, K.H. & Phillips, S.M. (2006) Bromeae. In: Zhengyi, W., Raven, P.H. & Deyuan, H. (Eds.) Flora of China, vol. 22. Science Press and Missouri Botanical Garden, Beijing and St. Louis, pp 370–386.
Linnaeus, C. (1753) Species plantarum: exhibentes plants rite cognitas, ad genera relates, cum differentiis specifics, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas, 2 vols. Impensis Laurentii Salvii, Holmiæ [Stockholm], 1200 pp. http://dx.doi.org/10.5962/bhl.title.669
Linnaeus, C. (1755) A. D. J. Centuria I. plantarum, quam, ex consens. nobil. et exper. Fac. med. in Regia Academia Upsaliensi, sub praesidio, Caroli Linnaei, orbi botanico examinandam sistit, Abraham. D. Juslenius, V-Gothus. Exc. L.M. Höjer, Reg.Acad. Typogr., Upsalla, 35 pp.
Liu, H., Hu, X.Y., Liu, Y.X. & Liu, Q. (2015) Caryopsis micromorphological survey of sorghum (Poaceae)- Taxonomic implications. South African Journal of Botany 99: 1–11. https://doi.org/10.1016/j.sajb.2015.02.015
Liu, Q., Zhang, D.X. & Peterson, P.M. (2010) Lemma micromorphological characters in the Chloridoideae (Poaceae) optimized on a molecular phylogeny. South African Journal of Botany 76: 196–209. https://doi.org/10.1016/j.sajb.2009.10.006
Matias Sales Machado, M.F. (1991) Contributiones selectae ad Floram et Vegetationem Orientis. Proceedings of the Third Plant Life of Southwest Asia Symposium Berlin 1990. In: Engel, T., Frey, W. & Kuerschner, H. (Eds.) Flora et Vegetation mundi, vol. 9. Borntraeger, Stuttgart, pp. 32.
Mejía-Saules, F. & Bisby, F.A. (2003) Silica bodies and hooked papillae in lemmas Melica species (Gramineae: Pooideae). Botanical Journal of the Linnean Society 141: 447–463. https://doi.org/10.1046/j.1095-8339.2003.00152.x
Meyer, C.A. (1831) Verzeichniss der Pflanzen, welche während der, auf Allerhöchsten Befehl, in den Jahren 1829 und 1830 unternommenen Reise im Caucasus und in den Provinzen am westlichen Ufer des Caspischen Meeres gefunden und eingesammelt worden sind. Bericht, abgestattet an die kaiserliche Academie der Wissenschaften in St. Petersburg, in ihrer Sitzung vom 30 März (11 April) 1831 vom Dr. Carl Anton Meyer. Gedruckt in der Buchdruckerei der kaiserl, Akademie der Wissenschaften, St.-Petersburg, 241 pp.
Naderi, R. & Rahiminejad, M.R. (2015) A taxonomic revision of the genus Bromus (Poaceae) and a new key to the tribe Bromeae in Iran. Annales Botanici Fennici 52: 233–248. http://dx.doi.org/10.5735/085.052.0316
Naderi, R., Rahiminejad, M.R., Assadi, M. & Vitek, E. (2016) A new taxonomic concept for Bromus danthoniae including comments on Bromus sect. Bromus and Triniusia (Poaceae). Annalen des Naturhistorischen Museums in Wien. Serie B 118: 167–180.
Nevski, S.A. (1934) Schedae ad Herbarium Florae Asiae Mediae ab Universitate Asiae Mediae editum. Trudy Sredne-Aziatskogo Gosudarstvennogo Universiteta. Seriya 8b, Botanika 17: 1–94.
Oja, T. & Jaaska, V. (1998) Allozyme diversity and phylogenetic relationships among diploid annual bromes (Bromus, Poaceae). Annales Botanici Fennici 35: 123–130.
Oja, T., Jaaska, V. & Vislap, V. (2003) Breeding system, evolution and taxonomy of Bromus arvensis, B. japonicus and B. squarrosus (Poaceae). Plant Systematics and Evolution 242: 101–117. https://doi.org/10.1007/s00606-003-0024-z
Olonova, M.V., Barkworth, M.E. & Gudkova, P.D. (2016) Lemma micromorphology and the systematics of Siberian species of Stipa (Poaceae). Nordic Journal of Botany 34 (3): 322–334.
Ortúňez, E. & Cano-Ruiz, J. (2013) Epidermal micromorphology of the genus Festuca L. subgenus Festuca (Poaceae). Plant Systematics and Evolution 299: 1471–1483. https://doi.org/10.1007/s00606-013-0809-7
Otfinowski, R., Kenkel, N.C. & Catling, P.M. (2007) The biology of Canadian weeds. 134. Bromus inermis Leyss. Canadian Journal of Plant Science 87: 183–198. https://doi.org/10.4141/P06-071
Pénzes, A. (1936) Rozsnok (Bromus) tanulmányok. Botanikai Közlemenyek 33: 111.
Pourmoshir, Z., Amirahmadi, A. & Naderi, R. (2017) Cladistic analysis of the tribe Bromeae (Poaceae) based on morphological characters in Iran. Taxonomy and Biosystematics 9: 1–14.
Presl, C.B. (1820) Cyperaceae et Gramineae siculae. apud D. Hartmann, Pragae, 58 pp.
Saarela, J.M., Burke, S.V., Wysocki, W.P., Barrett, M.D., Clark, L.G., Craine, J. M., Peterson, P.M., Soreng, R.J., Vorontsova, M.S. & Duvall, M. R. (2018) A 250 plastome phylogeny of the grass family (Poaceae): topological support under different data partitions. PeerJ 6: e4299. https://doi.org/10.7717/peerj.4299
Saarela, J.M., Wysocki, W.P., Barrett, C.F., Soreng, R.J., Davis, J.I., Clark, L.G., Kelchner, S.A., Pires, J.C., Edger, P.P., Mayfield, D.R. & Duvall, M.R. (2015) Plastid phylogenomics of the cool-season grass subfamily: clarification of relationships among early-diverging tribes. AoB Plants 7:plv046 https://doi.org/10.1093/aobpla/plv046.
Saarela, J.M., Peterson, P.M. & Valdés-Reyna, J. (2014) A taxonomic revision of Bromus (Poaceae: Pooideae: Bromeae) in Mexico and Central America. Phytotaxa 185: 1–147. http://dx.doi.org/10.11646/phytotaxa.185.1.1
Saarela, J.M., Peterson, P.M., Keane, R.M., Cayouette, J. & Graham, S.W. (2007) Molecular phylogenetics of Bromus (Poaceae: Pooideae) based on chloroplast and nuclear DNA sequence data. Aliso 23: 450–467. https://doi.org/10.5642/aliso.20072301.35
Sadeghian, S., Jafari, E. & Hatami, A. (2010) Cytogenetic studies in some species of Bromus L. in Iran. Asian Journal of Biological Sciences 3: 188–194. http://dx.doi.org/10.3923/ajbs.2010.188.194
Schneider, J., Döring, E., Hilu, K.W. & Röser, M. (2009) Phylogenetic structure of the grass subfamily Pooideae based on comparison of plastid matK gene–3′trnK exon and nuclear ITS sequences. Taxon 58: 405–424.
Schrenk, D.A. (1842) Novae Plantarum Species Nuperrime. Bulletin Scientifique Publie par l’Académie Imperiale des Sciences de Saint-Pétersbourg. St. Petersburg & Leipzig 10: 355.
Scholz, H. (1998) Notes on Bromus danthoniae and relatives (Gramineae). Willdenowia 28: 143–150. https://doi.org/10.3372/wi.28.2813
Shaw, R.B. & Montgomery, P. (2011) Guide to Texas grasses. Texas A & M University Press, College Station, 1096 pp.
Shear, C.L. (1900) A revision of the North American species of Bromus occurring north of Mexico. no. 23. United States Department of Agriculture, Division of Agrostology, Washington, D. C. pp. 5–66.
Sheidai, M. & Fadaei, F. (2005) Cytogenetic studies in some species of Bromus L., section Genea Dum. Journal of Genetics 84 (2): 189–194. https://doi.org/10.1007/BF02715845
Smith, P.M. (1985) Observations on Turkish brome- grasses. I. Some new taxa, new combinations and notes on typification. Notes from the Royal Botanic Garden, Edinburgh 42: 491–501.
Smith, P.M. & Sales, F. (1993) Bromus L. sect. Bromus: taxonomy and relationships of some species with small spikletes. Edinburgh Journal of Botany 50: 149–171. https://doi.org/10.1017/S0960428600002535
Smith, P.M. (1969) Serological relationship of Bromus L. and Boissiera Hochst. ex Steud Feddes Repertorium 79: 337–345 https://doi.org/10.1002/fedr.19690790602
Smith, P.M. (1970) Taxonomy and nomenclature of the brome grasses (Bromus L. s.l.). Notes from the Royal Botanic Garden, Edinburgh 30: 361–375.
Smith, P.M. (1972) Serology and species relationships in annual bromes (Bromus L. sect. Bromus). Annals of Botany 36: 1–30. https://doi.org/10.1093/oxfordjournals.aob.a084562
Snow, N. (1996) The phylogenetic utility of lemmatal micromorphology in Leptochloa s.l. and related genera in subtribe Eleusininae (Poaceae, Chloridoideae, Eragrostideae). Annals of the Missouri Botanical Garden 83: 504–529. https://doi.org/10.2307/2399991
Soreng, R.J. & Davis, J.I. (1998) Phylogenetics and character evolution in the grass family (Poaceae): simultaneous analysis of morphological and chloroplast DNA restriction site character sets. The Botanical Review 64: 1–85. https://doi.org/10.1007/BF02868851
Soreng, R.J., Peterson, P.M., Romaschenko, K., Davidse, G., Zuloaga, F.O., Judziewicz, E.J., Filgueiras, T.S., Davis, J.I. & Morrone, O. (2015) A worldwide phylogenetic classification of the Poaceae (Gramineae). Journal of Systematics and Evolution 53: 117–137. https://doi.org/10.1111/jse.12150
Stebbins, G.L. (1981) Chromosome and evolution in the genus Bromus (Gramineae). Botanische Jahrbücher fur Systematik, Pflanzengeschichte und Pflanzengeographie 102: 359–379.
Thunberg, C.P. (1784) Flora Japonica sistens plantas insularum japonicarum. Bibliopolio I.G. Mülleriano, Lipsiae, 419 pp. https://doi.org/10.5962/bhl.title.120525
Tournay, R. (1961) La nomenclature des sections du genre Bromus L. (Gramineae). Bulletin du Jardin botanique de l’État à Bruxelles 31: 289–299. https://doi.org/10.2307/3667334
Trinius, C.B. (1831) Geraminum genera quaedam speciesque complures definitionibus novis. Mémoires de l’Academie Imperiale des Sciences de St.-Petersbourg. Sixieme Serie. Sciences Mathematiques, Physiques et Naturelles t.1: 54–93.
Tzvelev, N.N. (1976) Zlaki SSSR. Nauka Publishers, Leningrad [in Russian]. [English translation: (1983) Grasses of the Soviet Union 1 & 2. Amerind Publishing Co, New Delhi, 778 pp.
Vahl, M. (1791) Symbolae botanicae, sive plantarum, tam earum, quas in itinere, imprimis orientali, collegit Petrus Forskål, quam aliarum, recentius detectarum, exactiores descriptiones, nec non observationes circa quasdam plantas dudum cognitas. Part 2. Impensis auctoris, excudebant Nicolaus Möller et filius, Hauniae [København], 105 pp.
Wagnon, H.K. (1952) A revision of the genus Bromus, section Bromopsis, of North America. Brittonia 7: 415–480. https://doi.org/10.2307/2804968
Weber, W.A. & Wittmann, R.C. (2011) Colorado Flora: Western Slope. Fourth edition. University Press of Colorado, Boulder, 608 pp.
Williams, W.M., Stewart, A.V. & Williamson, M.L. (2011) Bromus. In: Kole, C. (Ed.) Wild crop relatives: genomic and breeding resources, millets and grasses. Springer-Verlag, Heidelberg, pp 15–30.
Wróbel, A., Nobis, M., Nowak, A. (2017) Patterns of the lemma micromorphology: a useful tool in taxonomy of the Middle Asian Eragrostis species (Poaceae). Botany letters 164: 253–262. https://doi.org/10.1080/23818107.2017.1339293
Zhang, Y., Hu, X., Liu, Y. & Liu, Q. (2014) Caryopsis micromorphological survey of the genus Themeda (Poaceae) and allied spathaceous genera in the Andropogoneae. Turkish Journal of Botany 38: 665–676.
Ainouche, M.L. & Bayer, R.J. (1997) On the origins of the tetraploid Bromus species (section Bromus, Poaceae): insights from internal transcribed spacer sequences of nuclear ribosomal DNA. Genome 40: 730–743. http://doi.org/10.1139/g97-796
Armstrong, K.C. (1981) Evolution of Bromus inermis and related species of Bromus sect. Pnigma. Botanische Jahrbucher fur Systematik, Pflanzengeschichte und Pflanzengeographie 102: 427–443.
Barthlott, W., Neinhuis, Ch., Cutler, D., Ditsch, F., Meusel, I., Theisen, I. & Wilhelmi, H. (1998) Classification and terminology of plant epicuticular waxes. Botanical Journal of the Linnean Society 126: 237–260. https://doi.org/10.1111/j.1095-8339.1998.tb02529.x
Boissier, E. (1846) Diagnoses Plantarum Orientalium novarum. ser 1, vol. 7. Typographia Ferd. Ramboz, Genevae, 781 pp.
Boissier, E. (1884) Bromus. In: Boissier, E (Eds.) Flora Orientalis: sive, Enumeratio plantarum in Oriente a Graecia et Aegypto ad Indiae fines hucusque observatarum. vol. 5. H. Georg, Geneva and Basel, pp. 640–656. https://doi.org/10.5962/bhl.title.20323
Bor, N.L (1970) Bromeae. In: Rechinger, K.H. (Ed.) Flora Iranica, vol. 70. Akademische Druck- u. Verlagsanstalt, Graz, pp. 105–141.
Bor, N.L. (1968) Bromeae. In: Townsend, C.C., Guest, E. & Al-Rawi A (Eds.) Flora of Iraq, vol 9. Ministry of Agriculture and Agrarian Reform, Baghdad, pp. 126–166.
Bunge, A.A. von (1851) Beitrag zur Kenntniss der Flora Russlands und der Steppen Central-Asiens. Mémoires Presentes a l’Académie Impériale des Sciences de St.-Pétersbourg par Divers Savans et lus dans ses Assemblées 7: 177–535.
Clayton, W.D. & Renvoize, S.A. (1986) Genera Graminum: Grasses of the World. Her Majesty’s Stationary Office, London, 389 pp.
Cope, Th.A. (1982) Poaceae. In: Nasir, E. & Ali, S.I. (Eds.) Flora of Pakistan, vol 143. Pakistan Agricultural Research Council, Islamabad, pp. 1–678.
de Candolle, A.P. & Palisot de Beauvois, A.M.F.J. (1812) Essai d’une nouvelle agrostographie; ou nouveaux genres des graminées; avec figures représentant les caractéres de tous le genres. Imprimerie de Fain, Paris, 182 pp. http://dx.doi.org/10.5962/bhl.title.474.
Drobow, V. P. (1925) Gramineae novae turkestanicae.I. Repertorium specierum novarum regni vegetabilis 21: 37–46. https://doi.org/10.1002/fedr.19250210106
Dumortier, B.C.J. (1823 [1824]) Observations sur les Graminées de la Flora Belgique. J. Casterman, Tournay, 116 pp.
Ellis, R.P. (1979) A procedure for standardizing comparative leaf anatomy in the Poaceae. II. The epidermis as seen in surface view. Bothalia 12: 641–671. https://doi.org/10.4102/abc.v12i4.1441
Figari, A. B., De Notaris, J. (1852) Agrostographiae Aegyptiacae fragmenta. Memorie della Reale Accademia delle Scienze di Torino 2 (12): 245–262.
Fischer, F.E.L., Meyer, C.A., Trautvetter, E.R. & Avé-Lallemant, J.L.E. (1837) Animadversiones botanicae. Index seminum, quae Hortus Botanicus imperialis petropolitanus pro mutua commutatione offert. Accedunt animadversiones botanicae nonnullae 3: 1–19.
Fortune, P.M., Pourtau, N., Viron, N. & Ainouche, M.L. (2008) Molecular phylogeny and reticulate origins of the polyploidy Bromus species from section Genea (Poaceae). American Journal of Botany 95: 456–464.
Ghahreman, A., Alemi, M., Attar, F., Hamzehee, B. & Columbus, J.T. (2006) Anatomical studies in some species of Bromus (Poaceae) in Iran. Iranian Journal of Botany 12: 1–14.
Hackel, E. (1887) Gramineae. In: Engler, A. & Prantel, K. A. (Eds.) Naturlichen Panzenfamilien. (Abt. 2, Lief. 12). W. Engelmann, Leipzig, pp. 1–130. https://doi.org/10.5962/bhl.title.4635
Hackel, E. (1903) Gramineae. Videnskabelige Meddelelser fra den Naturhistoriske Forening i Kjøbenhavn 161–178.
Hammer, Ø., Harper, D. & Ryan, P.D. (2001) PAST: Paleontological Statistics software package for education and data analysis. Palaeontologia Electronica 4 (1): 1–9.
Hemsley, W.B. (1896) Hooker’s Icones Plantarum; or figures, with brief descriptive characters and remarks of new or rare plants, 25: t. London, 210 pp. https://doi.org/10.5962/bhl.title.16059
Hitchcock, A.S. (1935) Manual of the grasses of the United States. United States Department of Agriculture Miscellaneous Publication no. 200. United States Government Printing Office, Washington, D.C., 1040 pp.
Hochstetter, Ch.F.F. & Steudel, E. G. (1838) Einige nähere Bestimmungen der ägyptische arabischen Pflanzen. Flora oder Allgemeine Botanische Zeitung 21 (1): 23–27.
Huttanus, T.D., Mack, R.N., Novak, S.J. (2011) Propagule pressure and introduction pathways of Bromus tectorum (Cheat grass; Poaceae) in the Central United States. International Journal of Plant Sciences 172: 783–794. https://doi.org/10.1086/660107
Keshavarzi, M., Direkvandi, S. & Khoshnood, N. (2015) Chromosome number variation in Bromus L. (Poaceae) in Khorassan and Lorestan. Journal of Plant Research 28: 826–834.
Klimko, M. & Wysakowska, I. (2015) Epidermal features of glumes and florets in Aegilops geniculata Roth and Aegilops peregrina (Hack.) Maire et weiller × Secale cereal L. hybrids, amphiploids and parental forms. Steciana 19 (1): 13–24. https://doi.org/10.12657/STECIANA.019.003
Kosina, R. (1999) Patterns of flower microstructural variation within the genus Bromus. Acta Societatis Botanicorum Poloniae 68 (3): 221–226.
Leandro, T.D., Shirasuna, R.T., Filgueiras, T.S. & Scatena, V.L. (2016) The utility of Bambusoideae (Poaceae, Poales) leaf blade anatomy for identification and systematics. Brazilian Journal of Biology 76: 708–717. http://dx.doi.org/10.1590/1519-6984.01715
Ledebour, C.F.von. (1852) Flora rossica sive enumeratio plantarum in totius imperii rossici provinciis europaeis, asiaticis, et americanis hucusque observatarum 4, part 13. Sumptibus librariae E. Schweizerbart, Stuttgartiae, pp. 241–464. http://dx.doi.org/10.5962/bhl.title.6606
Liang, L., Shouliang, Ch., Guanghu, Zh., Ammann, K.H. & Phillips, S.M. (2006) Bromeae. In: Zhengyi, W., Raven, P.H. & Deyuan, H. (Eds.) Flora of China, vol. 22. Science Press and Missouri Botanical Garden, Beijing and St. Louis, pp 370–386.
Linnaeus, C. (1753) Species plantarum: exhibentes plants rite cognitas, ad genera relates, cum differentiis specifics, nominibus trivialibus, synonymis selectis, locis natalibus, secundum systema sexuale digestas, 2 vols. Impensis Laurentii Salvii, Holmiæ [Stockholm], 1200 pp. http://dx.doi.org/10.5962/bhl.title.669
Linnaeus, C. (1755) A. D. J. Centuria I. plantarum, quam, ex consens. nobil. et exper. Fac. med. in Regia Academia Upsaliensi, sub praesidio, Caroli Linnaei, orbi botanico examinandam sistit, Abraham. D. Juslenius, V-Gothus. Exc. L.M. Höjer, Reg.Acad. Typogr., Upsalla, 35 pp.
Liu, H., Hu, X.Y., Liu, Y.X. & Liu, Q. (2015) Caryopsis micromorphological survey of sorghum (Poaceae)- Taxonomic implications. South African Journal of Botany 99: 1–11. https://doi.org/10.1016/j.sajb.2015.02.015
Liu, Q., Zhang, D.X. & Peterson, P.M. (2010) Lemma micromorphological characters in the Chloridoideae (Poaceae) optimized on a molecular phylogeny. South African Journal of Botany 76: 196–209. https://doi.org/10.1016/j.sajb.2009.10.006
Matias Sales Machado, M.F. (1991) Contributiones selectae ad Floram et Vegetationem Orientis. Proceedings of the Third Plant Life of Southwest Asia Symposium Berlin 1990. In: Engel, T., Frey, W. & Kuerschner, H. (Eds.) Flora et Vegetation mundi, vol. 9. Borntraeger, Stuttgart, pp. 32.
Mejía-Saules, F. & Bisby, F.A. (2003) Silica bodies and hooked papillae in lemmas Melica species (Gramineae: Pooideae). Botanical Journal of the Linnean Society 141: 447–463. https://doi.org/10.1046/j.1095-8339.2003.00152.x
Meyer, C.A. (1831) Verzeichniss der Pflanzen, welche während der, auf Allerhöchsten Befehl, in den Jahren 1829 und 1830 unternommenen Reise im Caucasus und in den Provinzen am westlichen Ufer des Caspischen Meeres gefunden und eingesammelt worden sind. Bericht, abgestattet an die kaiserliche Academie der Wissenschaften in St. Petersburg, in ihrer Sitzung vom 30 März (11 April) 1831 vom Dr. Carl Anton Meyer. Gedruckt in der Buchdruckerei der kaiserl, Akademie der Wissenschaften, St.-Petersburg, 241 pp.
Naderi, R. & Rahiminejad, M.R. (2015) A taxonomic revision of the genus Bromus (Poaceae) and a new key to the tribe Bromeae in Iran. Annales Botanici Fennici 52: 233–248. http://dx.doi.org/10.5735/085.052.0316
Naderi, R., Rahiminejad, M.R., Assadi, M. & Vitek, E. (2016) A new taxonomic concept for Bromus danthoniae including comments on Bromus sect. Bromus and Triniusia (Poaceae). Annalen des Naturhistorischen Museums in Wien. Serie B 118: 167–180.
Nevski, S.A. (1934) Schedae ad Herbarium Florae Asiae Mediae ab Universitate Asiae Mediae editum. Trudy Sredne-Aziatskogo Gosudarstvennogo Universiteta. Seriya 8b, Botanika 17: 1–94.
Oja, T. & Jaaska, V. (1998) Allozyme diversity and phylogenetic relationships among diploid annual bromes (Bromus, Poaceae). Annales Botanici Fennici 35: 123–130.
Oja, T., Jaaska, V. & Vislap, V. (2003) Breeding system, evolution and taxonomy of Bromus arvensis, B. japonicus and B. squarrosus (Poaceae). Plant Systematics and Evolution 242: 101–117. https://doi.org/10.1007/s00606-003-0024-z
Olonova, M.V., Barkworth, M.E. & Gudkova, P.D. (2016) Lemma micromorphology and the systematics of Siberian species of Stipa (Poaceae). Nordic Journal of Botany 34 (3): 322–334.
Ortúňez, E. & Cano-Ruiz, J. (2013) Epidermal micromorphology of the genus Festuca L. subgenus Festuca (Poaceae). Plant Systematics and Evolution 299: 1471–1483. https://doi.org/10.1007/s00606-013-0809-7
Otfinowski, R., Kenkel, N.C. & Catling, P.M. (2007) The biology of Canadian weeds. 134. Bromus inermis Leyss. Canadian Journal of Plant Science 87: 183–198. https://doi.org/10.4141/P06-071
Pénzes, A. (1936) Rozsnok (Bromus) tanulmányok. Botanikai Közlemenyek 33: 111.
Pourmoshir, Z., Amirahmadi, A. & Naderi, R. (2017) Cladistic analysis of the tribe Bromeae (Poaceae) based on morphological characters in Iran. Taxonomy and Biosystematics 9: 1–14.
Presl, C.B. (1820) Cyperaceae et Gramineae siculae. apud D. Hartmann, Pragae, 58 pp.
Saarela, J.M., Burke, S.V., Wysocki, W.P., Barrett, M.D., Clark, L.G., Craine, J. M., Peterson, P.M., Soreng, R.J., Vorontsova, M.S. & Duvall, M. R. (2018) A 250 plastome phylogeny of the grass family (Poaceae): topological support under different data partitions. PeerJ 6: e4299. https://doi.org/10.7717/peerj.4299
Saarela, J.M., Wysocki, W.P., Barrett, C.F., Soreng, R.J., Davis, J.I., Clark, L.G., Kelchner, S.A., Pires, J.C., Edger, P.P., Mayfield, D.R. & Duvall, M.R. (2015) Plastid phylogenomics of the cool-season grass subfamily: clarification of relationships among early-diverging tribes. AoB Plants 7:plv046 https://doi.org/10.1093/aobpla/plv046.
Saarela, J.M., Peterson, P.M. & Valdés-Reyna, J. (2014) A taxonomic revision of Bromus (Poaceae: Pooideae: Bromeae) in Mexico and Central America. Phytotaxa 185: 1–147. http://dx.doi.org/10.11646/phytotaxa.185.1.1
Saarela, J.M., Peterson, P.M., Keane, R.M., Cayouette, J. & Graham, S.W. (2007) Molecular phylogenetics of Bromus (Poaceae: Pooideae) based on chloroplast and nuclear DNA sequence data. Aliso 23: 450–467. https://doi.org/10.5642/aliso.20072301.35
Sadeghian, S., Jafari, E. & Hatami, A. (2010) Cytogenetic studies in some species of Bromus L. in Iran. Asian Journal of Biological Sciences 3: 188–194. http://dx.doi.org/10.3923/ajbs.2010.188.194
Schneider, J., Döring, E., Hilu, K.W. & Röser, M. (2009) Phylogenetic structure of the grass subfamily Pooideae based on comparison of plastid matK gene–3′trnK exon and nuclear ITS sequences. Taxon 58: 405–424.
Schrenk, D.A. (1842) Novae Plantarum Species Nuperrime. Bulletin Scientifique Publie par l’Académie Imperiale des Sciences de Saint-Pétersbourg. St. Petersburg & Leipzig 10: 355.
Scholz, H. (1998) Notes on Bromus danthoniae and relatives (Gramineae). Willdenowia 28: 143–150. https://doi.org/10.3372/wi.28.2813
Shaw, R.B. & Montgomery, P. (2011) Guide to Texas grasses. Texas A & M University Press, College Station, 1096 pp.
Shear, C.L. (1900) A revision of the North American species of Bromus occurring north of Mexico. no. 23. United States Department of Agriculture, Division of Agrostology, Washington, D. C. pp. 5–66.
Sheidai, M. & Fadaei, F. (2005) Cytogenetic studies in some species of Bromus L., section Genea Dum. Journal of Genetics 84 (2): 189–194. https://doi.org/10.1007/BF02715845
Smith, P.M. (1985) Observations on Turkish brome- grasses. I. Some new taxa, new combinations and notes on typification. Notes from the Royal Botanic Garden, Edinburgh 42: 491–501.
Smith, P.M. & Sales, F. (1993) Bromus L. sect. Bromus: taxonomy and relationships of some species with small spikletes. Edinburgh Journal of Botany 50: 149–171. https://doi.org/10.1017/S0960428600002535
Smith, P.M. (1969) Serological relationship of Bromus L. and Boissiera Hochst. ex Steud Feddes Repertorium 79: 337–345 https://doi.org/10.1002/fedr.19690790602
Smith, P.M. (1970) Taxonomy and nomenclature of the brome grasses (Bromus L. s.l.). Notes from the Royal Botanic Garden, Edinburgh 30: 361–375.
Smith, P.M. (1972) Serology and species relationships in annual bromes (Bromus L. sect. Bromus). Annals of Botany 36: 1–30. https://doi.org/10.1093/oxfordjournals.aob.a084562
Snow, N. (1996) The phylogenetic utility of lemmatal micromorphology in Leptochloa s.l. and related genera in subtribe Eleusininae (Poaceae, Chloridoideae, Eragrostideae). Annals of the Missouri Botanical Garden 83: 504–529. https://doi.org/10.2307/2399991
Soreng, R.J. & Davis, J.I. (1998) Phylogenetics and character evolution in the grass family (Poaceae): simultaneous analysis of morphological and chloroplast DNA restriction site character sets. The Botanical Review 64: 1–85. https://doi.org/10.1007/BF02868851
Soreng, R.J., Peterson, P.M., Romaschenko, K., Davidse, G., Zuloaga, F.O., Judziewicz, E.J., Filgueiras, T.S., Davis, J.I. & Morrone, O. (2015) A worldwide phylogenetic classification of the Poaceae (Gramineae). Journal of Systematics and Evolution 53: 117–137. https://doi.org/10.1111/jse.12150
Stebbins, G.L. (1981) Chromosome and evolution in the genus Bromus (Gramineae). Botanische Jahrbücher fur Systematik, Pflanzengeschichte und Pflanzengeographie 102: 359–379.
Thunberg, C.P. (1784) Flora Japonica sistens plantas insularum japonicarum. Bibliopolio I.G. Mülleriano, Lipsiae, 419 pp. https://doi.org/10.5962/bhl.title.120525
Tournay, R. (1961) La nomenclature des sections du genre Bromus L. (Gramineae). Bulletin du Jardin botanique de l’État à Bruxelles 31: 289–299. https://doi.org/10.2307/3667334
Trinius, C.B. (1831) Geraminum genera quaedam speciesque complures definitionibus novis. Mémoires de l’Academie Imperiale des Sciences de St.-Petersbourg. Sixieme Serie. Sciences Mathematiques, Physiques et Naturelles t.1: 54–93.
Tzvelev, N.N. (1976) Zlaki SSSR. Nauka Publishers, Leningrad [in Russian]. [English translation: (1983) Grasses of the Soviet Union 1 & 2. Amerind Publishing Co, New Delhi, 778 pp.
Vahl, M. (1791) Symbolae botanicae, sive plantarum, tam earum, quas in itinere, imprimis orientali, collegit Petrus Forskål, quam aliarum, recentius detectarum, exactiores descriptiones, nec non observationes circa quasdam plantas dudum cognitas. Part 2. Impensis auctoris, excudebant Nicolaus Möller et filius, Hauniae [København], 105 pp.
Wagnon, H.K. (1952) A revision of the genus Bromus, section Bromopsis, of North America. Brittonia 7: 415–480. https://doi.org/10.2307/2804968
Weber, W.A. & Wittmann, R.C. (2011) Colorado Flora: Western Slope. Fourth edition. University Press of Colorado, Boulder, 608 pp.
Williams, W.M., Stewart, A.V. & Williamson, M.L. (2011) Bromus. In: Kole, C. (Ed.) Wild crop relatives: genomic and breeding resources, millets and grasses. Springer-Verlag, Heidelberg, pp 15–30.
Wróbel, A., Nobis, M., Nowak, A. (2017) Patterns of the lemma micromorphology: a useful tool in taxonomy of the Middle Asian Eragrostis species (Poaceae). Botany letters 164: 253–262. https://doi.org/10.1080/23818107.2017.1339293
Zhang, Y., Hu, X., Liu, Y. & Liu, Q. (2014) Caryopsis micromorphological survey of the genus Themeda (Poaceae) and allied spathaceous genera in the Andropogoneae. Turkish Journal of Botany 38: 665–676.