The leaf blade epidermis of 20 taxa of Bromus s. l. (sections Genea, Bromus, Bromopsis and Ceratochloa; tribe Bromeae) occurring in Ukraine were studied using SEM. The aim of the study was assessment of the taxonomic value of micromorphological characters and resolving some taxonomical and phylogenetical issues in the group. The main epidermal characters of leaf blades were shape, size and distribution of long cells, silica cells, crown cells, prickles, macrohairs, and cells of stomatal complexes, as well as cell wall shape, for some of them. Almost all mentioned types of cells occur on both abaxial and adaxial surfaces. The results show that crown cells were not observed in species of sect. Ceratochloa and in species B. erectus, B. benekenii, B. ramosus; prickles were not found in species of sect. Genea and some species of sect. Bromopsis. By some micromorphological characters (crown cells, prickles, and length of long cells), species of sect. Bromopsis can be subdivided into two groups. The first group consists of Bromus erectus, B. benekenii, and B. ramosus, the second one, of B. riparius, B. cappadocicus (incl. Bromopsis heterophylla, B. calcarea, B. pseudocappadocica, B. cimmerica), and the hybrid known as Bromopsis × taurica (nom. illeg.). Bromus inermis have intermediate characters. Other micromorphological features (long cells, silica cells, macrohairs and cells of stomatal complexes) in the remaining taxa were common, their variability was high. It has been found that all studied characters of the leaf blade epidermis are not specifically useful for resolution of the main taxonomic issues within tribe Bromeae. According to the conducted analysis of micromorphological features, the genus Bromus s. l. in the flora of Ukraine comprises four section-rank taxa: sect. Bromus, sect. Genea (= genus Anisantha), sect. Bromopsis, and sect. Ceratochloa.

Supplementary Material. Electronic Supplement (Table E1, p. e1) is available in the online version of this article.

Keywords: Anisantha, Bromeae, Bromopsis, Bromus, Ceratochloa, epidermis, micromorphology, Poaceae, SEM, taxonomy

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1. Acedo C., Llamas F. 2001. Variation of morphological characters of lemma and palea in the genus Bromus (Poaceae). Annales Botanici Fennici, 38(1): 1–14.
2. Ainouche M.L., Bayer R.J. 1996. On the origins of two Mediterranean allotetraploid Bromus species: Bromus hordeaceus L. and B. lanceolatus Roth. (Poaceae). American Journal of Botany, 83: 135.
3. Ainouche M.L., Bayer R.J., Gourret J.-P., Defontaine A., Misset M.T. 1999. The allotetraploid invasive weed Bromus hordeaceus L. (Poaceae): genetic diversity, origin and molecular evolution. Folia Geobotanica, 34: 405–419. https://doi.org/10.1007/BF02914919
4. Ammann K. 1981. Bestimmungsschwierigkeiten bei europäischer Bromus-Arten. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie, 102(1–4): 459–469.
5. Armstrong K.C. 1981. The evolution of Bromus inermis and related species of Bromus sect. Pnigma. Botanische Jahrbücher fur Systematik, Pflanzengeschichte und Pflanzengeographie, 102(1–4): 427–443.
6. Barthlott W., Frölich D. 1983. Mikromorphologie und Orientierungsmuster epicuticularer Wachs-Kristalloide: Ein neues systematisches Merkmal bei Monokotylen. Plant Systematics and Evolution, 142: 171–185. https://doi.org/10.1007/BF00985897
7. Bor N.L. 1970. Gramineae. In: Flora Iranica, vol. 70. Ed. K.H. Rechinger. Graz (Austria): Natural History Museum Vienna, Akademische Druck- und Verlagsanstalt, 573 pp. + 72 tab.
8. Doğan M., Behçet L., Sinan A. 2015. Pseudophleum anatolicum, a new endemic species of Pseudophleum (Poaceae) from East Anatolia, Turkey. Systematic Botany, 40(2): 454–460. https://doi.org/10.1600/036364415X688349
9. Ellis R.P. 1979. A procedure for standardizing comparative leaf anatomy in the Poaceae. II. The epidermis as seen in surface view. Bothalia, 12(4): 641–671. https://doi.org/10.4102/abc.v12i4.1441
10. Fortune P.M, Pourta U.N, Viro N.N., Ainouche M.L. 2008. Molecular phylogeny and reticulate origins of the polyploid Bromus species from section Genea (Poaceae). American Journal of Botany, 95(4): 454–464. https://doi.org/10.3732/ajb.95.4.454
11. Garcke A. 1972. Illustrierte Flora. Deutschland und angrenzende Gebiete. Gefässkryptogamen und Blütenpflanzen, 23. Auflage. Berlin; Hamburg: Verlag Paul Parey, 1607 S.
12. Gutiérrez H.F., Pensiero J.F. 1998. Sinopsis de las especies argentinas del Género Bromus (Poaceae). Darwiniana 35(1–4): 75–114. https://doi.org/10.14522/darwiniana.2014.351-4.397
13. Ilvessalo-Pfäffli M.-S. 1995. Fiber atlas: identification of papermaking fibers. Berlin; Heidelberg: Springer, 407 pp. https://doi.org/10.1007/978-3-662-07212-7
14. Klokov M.V. 1950a. Notulae Systematicae ex Herbario Instituti Botanici nomine V.L. Komarovi Academiae Scientiarum URSS, 12: 35–60.
15. Klokov M.V. 1950b. Gramineae. In: Vyznachnyk Roslyn UkrRSR. Kyiv; Kharkiv: Derzhsilhospvydav, pp. 837–908.
16. Klokov M.V. 1976 (1977). In: Novitates Systematicae Plantarum Vascularium et Non Vascularium. Kyiv: Naukova Dumka, pp. 28–48.
17. Kožuharov S., Petrova A., Ehrendorfer F. 1981. Evolutionary patterns in some brome grass species (Bromus, Gramineae) of the Balkan Peninsula. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie, 102(1–4): 381–391.
18. Krasniak O.I. 2011. Chornomorski Botanical Journal, 7(1): 43–45. https://doi.org/10.14255/2308-9628/11.71/4
19. Krasniak O.I. 2012. In: Modern Phytomorphology: materials of the 1^st International scientific conference on plant morphology (Lviv, 4 April, 2012). Lviv, pp. 107–110.
20. Lavrenko E.M. 1940. Bromus. In: Flora of the Ukrainian RSR, vol. 2. Ed. E.M. Lavrenko. Kyiv: Academy of Sciences of Ukrainian RSR, pp. 293–315.
21. Massa A.N., Jensen K.B., Larson S.R., Hole D. 2011. Morphological variation in Bromus sect. Ceratochloa germplasm of Patagonia. Canadian Journal of Botany, 82(1): 136–144. https://doi.org/10.1139/b03-133
22. Mashau A.C., Fish L., Van Wyk A.E. 2015. Taxonomic significance of the abaxial lemma surface in southern African members of Helictotrichon (Poaceae). Bothalia, 45(1): 1–8. https://doi.org/10.4102/abc.v45i1.1393
23. Matthei O. 1986. The genus Bromus L. Poaceae in Chile. Gayana Botanica (Chile), 43(1–4): 47–110.
24. Meng L., Mao P. 2013. Micromorphological and anatomical features of four species of Elytrigia Desv. (Poaceae). Bangladesh Journal of Plant Taxonomy, 20(2): 135–144. https://doi.org/10.3329/bjpt.v20i2.17388
25. Metcalfe C.R. 1960. Anatomy of Monocotyledons, vol. 1. Gramineae. Oxford; New York: Oxford University Press, lxi + 731 pp.
26. Mosyakin S.L. 1990. Ukrainian Botanical Journal, 47(1): 101–102.
27. Mosyakin S.L. 1991. Ukrainian Botanical Journal, 48(3): 45–48.
28. Mosyakin S.L., Fedoronchuk M.M. 1999. Vascular plants of Ukraine. A nomenclatural Checklist. Kiev, xxiii + 345 pp. https://doi.org/10.13140/2.1.2985.0409
29. Naganowska B. 1993. Karyotypes of five Bromus species of Genea section. Genetica Polonica, 34(3): 197–213.
30. Oja T., Jaaska V. 1998. Allozyme diversity and phylogenetic relationships among diploid annual bromes (Bromus, Poaceae). Annales Botanici Fennici, 35(2): 123–130.
31. 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(1/4): 101–117. https://doi.org/10.1007/s00606-003-0024-z
32. Oja T., Paal J. 2007. Multivariate analysis of morphological variation among closely related species Bromus japonicus, B. squarrosus and B. arvensis (Poaceae) in comparison with isozyme evidences. Nordic Journal of Botany, 24(6): 691–702. https://doi.org/10.1111/j.1756-1051.2004.tb01939.x
33. Ortúñez E., de la Fuente V. 2010. Epidermal micromorphology of the genus Festuca L. (Poaceae) in the Iberian Peninsula. Plant Systematics and Evolution, 284(3–4): 201–218. https://doi.org/10.1007/s00606-009-0248-7
34. Palmer P.G., Tucker A.E. 1983. A Scanning Electron Microscope Survey of the Epidermis of East African Grasses, Series II. In: Smithsonian Contributions to Botany, pp. 1–72. https://doi.org/10.5479/si.0081024X.53
35. Pavlick L.E., Anderton L.K. 2007. Bromus L. In: Flora of North America, vol. 24. Eds M.E. Barkworth, K.M. Capels, S. Long, L.K. Anderton, M.B. Piep. New York; Oxford: Oxford University Press, pp. 193–237.
36. Peterson P.M., Planchuelo A.M. 1988. Bromus catharticus in South America (Poaceae: Bromeae). Novon, 8(1): 53–60. https://doi.org/10.2307/3391893
37. Petrova A., Kožuharov S., Ehrendorfer F. 1997. Karyosystematic notes on Bromus (Gramineae), and a new species of the B. riparius polyploid complex from Bulgaria. Bocconea, 5(2): 775–780.
38. Pillay M., Hilu K.W. 1990. Chloroplast DNA variation in diploid and polyploid species of Bromus (Poaceae) subgenera Festucaria and Ceratochloa. Theoretical and Applied Genetics, 80(3): 326–332. https://doi.org/10.1007/BF00210067
39. Pillay M., Hilu K.W. 1995. Chloroplast-DNA restriction siteanalysis in the genus Bromus (Poaceae). American Journal of Botany, 82(2): 239–249. https://doi.org/10.2307/2445531
40. Reimer E., Cota-Sánchez J.H. 2007. An SEM survey of the leaf epidermis in Danthonioid grasses (Poaceae). Danthonioideae. Systematic Botany, 32(1): 60–70. https://doi.org/10.1600/036364407780360139
41. Planchuelo A.M., Peterson P.M. 2000. The species of Bromus (Poaceae: Bromeae) in South America. In: Grasses systematics and evolution. Eds S .W.L. Jacobs, J. Everett. Collinwood, Australia: CSIRO Publishing, pp. 89–101.
42. 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(1): 379–396. https://doi.org/10.5642/aliso.20072301.35
43. 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): 1–147. https://doi.org/10.11646/phytotaxa.185.1.1
44. Sales F. 1993. Taxonomy and nomenclature of Bromus sect. Genea. Edinburgh Journal of Botany, 50(1): 1–31. https://doi.org/10.1017/S0960428600000627
45. Sales F. 1994. Evolutionary tendencies in some annual species of Bromus (Bromus L. sect. Genea Dum. (Poaceae)). Botanical Journal of the Linnean Society, 115(3): 197–210. https://doi.org/10.1111/j.1095-8339.1994.tb01778.x
46. Scholz H. 1981. Bemerkungen über Bromus madritensis und B. rubens (Gramineae). Willdenowia, 11(2): 249–258.
47. Schulz-Schaeffer J. 1960. Cytological investigations in the genus Bromus. III. The cytotaxonomic significance of the satellite chromosomes. Journal of Heredity, 51(6): 269–277. https://doi.org/10.1093/oxfordjournals.jhered.a107007
48. Shaw R.B., Montgomery P. 2011. Guide to Texas Grasses. Texas: Texas A&M University Press, College Station, 1096 pp.
49. Sljussarenko L.P. 1977. Bromeae. In: Grasses of Ukraine. Kyiv: Naukova Dumka, pp. 124–153.
50. Smith P.M. 1972. Serology and species relationship in annual bromes (Bromus L. sect. Bromus). Annals of Botany, 36(1): 1–30. https://doi.org/10.1093/oxfordjournals.aob.a084562
51. Smith P.M. 1985. Bromus. In: Flora of Turkey, vol. 9. Ed. P.H. David. Edinburgh: University of Edinburgh, pp. 272–302.
52. Smith P.M., Sales F. 1993. Bromus L. sect. Bromus: taxonomy and relationship of some species with small spikelets. Edinburgh Journal of Botany, 50(2): 149–171. https://doi.org/10.1017/S0960428600002535
53. Spalton L.M. 2002. An analysis of the characters of Bromus racemosus L., B. commutatus Schrad. and B. secalinus L. (Poaceae). Watsonia, 24(2): 193–202.
54. Stebbins G.L. 1981. Chromosomes and evolution in the genus Bromus. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie, 102(1–4): 359–379.
55. Terrell E.E., Peterson P.M., Wergin W.P. 2001. Epidermal Features and Spikelet Micromorphology in Oryza and Related Genera (Poaceae: Oryzeae). Smithsonian Institution Press Washington, D.C., 50 pp. https://doi.org/10.5479/si.0081024X.91
56. Tzvelev N.N. 1976. Grasses of the USSR. Leningrad: Nauka, 788 pp.
57. Tzvelev N.N. 2006. Novitates Systematicae Plantarum Vascularium, 38: 66–113.
58. Vvedensky A.I., Krechetovich V.I., Nevsky S.A., Sochava W.B. 1934. Bromus. In: Flora of the USSR, vol. 2. Leningrad: Academy of Sciences of the USSR, pp. 554–584.
59. Webb M.E., Almeida M.T. 1990. Micromorphology of the leaf epidermis in taxa of the Agropyron-Elymus-complex (Poaceae). Botanical Journal of the Linnean Society, 103(2):153–158. https://doi.org/10.1111/j.1095-8339.1990.tb00181.x
60. Weber W.A., Wittmann R.C. 2011. Colorado Flora: Western Slope. 4^th ed. University Press of Colorado, 608 pp. https://doi.org/10.5876/9781607321439.00
61. Yousaf Z., Shinwari Z.Kh., Asghar R., Parveen A. 2008. Leaf epidermal anatomy of selected Allium species, family Alliaceae from Pakistan. Pakistan Journal of Botany, 40(1): 77–90.