Pollen morphology of 33 species belonging to 12 genera of Orobanchaceae tribe Rhinantheae was studied using light and scanning electron microscopy. Pollen grains of the studied species are 3(4)-colpate, prolate, oval, spheroidal and oblate-spheroidal. Outline in equatorial view is elliptical or circular, outline in polar view is slightly-3-lobed, 3(4)-lobed, rounded, rounded-triangular or triangular; pollen grains are mainly medium- and sometimes small-sized. Colpi are long, sometimes medium or short. Pollen grains in Rhinantheae have different patterns of their exine sculpture: retipilate, retipilate-rugulate, rugulate, rugulate-foveolate, implecto-microreticulate, implecto-microreticulate-tuberculate, rugulate-tuberculate, unequally-tuberculate, tuberculate, granulate-tuberculate-perforate, and perforate-tuberculate. Pollen grains in the studied taxa can be subdivided into 10 basic types, based on their exine sculpture. Within some types, 6 additional subtypes are distinguished, mainly according to their exine sculpture peculiarities, size, shape, outline, colpi, colpus membrane and exine thickness. Pollen grains in members of the tribe are distinguishable at generic and species levels. At the generic level, pollen grains of Pterygiella, Melampyrum, Rhynchocorys and Tozzia are well distinguished. Pollen grains of Odontites and Rhinanthus are similar in their implecto-microreticulate and rugulate-foveolate exine and differ in other characteristics (shape, outline, size, peculiarities of colpi structure) at the species level. Pollen grains of Bellardia and Lathraea are similar in their retipilate and retipilate-rugulate exine sculpture and differ in shape, outline, structure of colpi, and sculpture membranes. Pollen grains of Bartsia, Parentucellia and Macrosyringion are similar in having retipilate exine sculpture and differ in size, outline, of colpi structure and characters of exine sculpture. Palynomorphological data correspond well to the plylogenetic pattern of genera of Rhinantheae according to molecular phylogenetic data, and partly correspond to traditional systems. Possible trends of evolution of exine sculpture in pollen grains of Rhinantheae are revealed.

Keywords: pollen grains, morphology, sculpture, taxonomy, Rhinantheae, Orobanchaceae

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1. Aleshina L.A. Scrophulariaceae. In: Pyltsa dvudolnykh rasteniy flory evropeyskoy chasti SSSR. Lamiaceae – Zygophyllaceae, Leningrad: Nauka, 1978, vol. 2, pp. 125– 132.
2. Bennett J.R., Mathews S. Phylogeny of the parasitic plant family Orobanchaceae inferred from phytochrome A. Amer. J. Bot., 2006, 93(7): 1039–1051. https://doi.org/10.3732/ajb.93.7.1039 https://www.ncbi.nlm.nih.gov/pubmed/21642169
3. Bolliger M., Wick L. The pollen morphology of Odontites (Scrophulariaceae) and its taxonomic significance. Plant Syst. Evol., 1990, 173: 159–178. https://doi.org/10.1007/BF00940860
4. Erdtman G. Pollen morphology and plant taxonomy. Angiosperms, Stockholm: Almqvist & Wiksell, 1952, 539 pp.
5. Faegri K., Iversen J. Textbook of pollen analysis, Oxford: Blackwell, 1964, 237 pp.
6. Fischer E. Scrophulariaceae. In: The families and genera of vascular plants. Ed. K. Kubitzki, Berlin: Springer, 2004, pp. 333–432.
7. Inceoğlu Ö. Pollen grains in some Turkish Rhinantheae (Scrophulariaceae). Grana, 1982, 21: 83–96. https://doi.org/10.1080/00173138209427684
8. Kupriyanova L.A., Aleshina L.A. Pyltsa i spory rasteniy flory evropeyskoy chasti SSSR, Leningrad: Nauka, 1972, vol. 1, 170 pp.
9. Lu L., Wang H., Blackmore S., Li D.-Z., Dong L.-N. Pollen morphology of the tribe Rhinantheae (Orobanchaceae) and its systematic significances. Plant Syst. Evol., 2007, 268: 177–198. https://doi.org/10.1007/s00606-007-0562-x
10. Minkin J.P., Eshbaugh W.H. Pollen morphology of the Orobanchaceae and rhinanthoid Scrophulariaceae. Grana, 1989, 28: 1–18. https://doi.org/10.1080/00173138909431007
11. Moore P.D., Webb J.A. An illustrated guide to pollen analysis, London; Sydney; Auckland; Toronto: Hodder and Stoughton, 1983, 133 pp.
12. McNeal J.R., Bennett J.R., Wolfe A.D., Mathews S. Phylogeny and origins of holoparasitism in Orobanchaceae. Amer. J. Bot., 2013, 100: 971–983. https://doi.org/10.3732/ajb.1200448 https://www.ncbi.nlm.nih.gov/pubmed/23608647
13. Olmstead R.G., DePamphilis C.W., Wolfe A.D., Young N.D., Elisons W.J., Reeves P.A. Disintegration of the Scrophulariaceae. Amer. J. Bot., 2001, 88(2): 348–361. https://doi.org/10.2307/2657024 https://www.ncbi.nlm.nih.gov/pubmed/11222255
14. Olmstead R. A synoptical classification of the Lamiales. Version 2.6.2 (updated 12 April, 2016), available at: http://depts.washington.edu/phylo/Classification.pdf
15. Oxelman B., Kornhall P., Olmstead R., Bremer B. Further disintegration of the Scrophulariaceae. Taxon, 2005, 54: 411–425. https://doi.org/10.2307/25065369
16. Peregrym O.M., Tsymbalyuk Z.M. Nauk. visnyk Chernivets. univ., 2009, 455: 107–110.
17. Peregrym O.M., Tsymbalyuk Z.M., Mosyakin S.L. Pollen morphology of Ukrainian species of the genus Pedicularis L. (Orobanchaceae Vent.). Biodiv. Res. Conserv., 2011, 24: 5–12.
18. Punt W., Blackmore S., Nilsson S., Thomas A. Le. Glossary of pollen and spore terminology, Utrecht: LPP Found., 1994, 71 pp.
19. Reveal J.L. An outline of a classification scheme for extant flowering plants. Phytoneuron, 2012, 2012-37: 1–221.
20. Saeidi-Mehrvarz S., Panah S.P., Faghir M.B. A palynological study of the genus Pedicularis (Orobanchaceae) in Iran. Willdenowia, 2013, 43: 279–285. https://doi.org/10.3372/wi.43.43207
21. Saeidi-Mehrvarz S., Roohi S., Mehrgan I., Roudi E. Comparative morphological, anatomical and palynological investigations of the genus Euphrasia L. (Orobanchaceae) in Iran. Wulfenia, 2012, 19: 23–37.
22. Severova E.E. In: Materialy X Moskovskogo soveshchaniya po filogenii rasteniy, Moscow: Izd-vo MGU, 1999a, pp. 149–151.
23. Severova E.E. In: Aktualnye problemy palinologii na rubezhe tretego tysyacheletiya: tezisy dokl. IX Vseros. palinol. konf., Moscow: IGiRGI, 1999b, pp. 263–264.
24. Takhtajan A.L. Sistema magnoliofitov, Leningrad: Nauka, 1987, 439 pp.
25. Takhtajan A.L. Diversity and classification of flowering plants, New York: Columbia Univ. Press, 1997, 663 pp.
26. Takhtajan A. Flowering Plants, Berlin: Springer Verlag, 2009, 871 pp. https://doi.org/10.1007/978-1-4020-9609-9
27. Tank D.C., Beardsley P.M., Kelchner S.A., Olmstead R.G. Review of the systematics of Scrophulariaceae s. l. and their current disposition. Austral. Syst. Bot., 2006, 19: 289–307. https://doi.org/10.1071/SB05009
28. Tokarev P.I. Morfologiya i ultrastruktura pyltsevykh zeren, Moscow: T-vo nauchn. izd. KMK, 2002, 51 pp.
29. Tsymbalyuk Z.N. Palinomorfologicheskie osobennosti roda Rhinanthus L. In: Evolyutsiya zhizni na Zemle: mat. IV Mezhdunar. simp., 10–12 noyabrya 2010 g., Tomsk: TML-Press, 2010, pp. 117–120.
30. Tsymbalyuk Z.M. Ukr. Bot. J., 2011, 68(1): 45–57.
31. Tsymbalyuk Z.N. Palinomorfologiya predstaviteley triby Rhinantheae Wettst. flory Ukrainy. In: Problemy sovremennoy palinologii: mat. XIII Ros. palinol. konf., Syktyvkar: IG Komi NTs UrO RAN, 2011, vol. 1, pp. 68–73.
32. Tsymbalyuk Z.M. Palinomorfolohichni osoblyvosti predstavnykiv poryadku Lamiales s. l.: filohenetychne znachennya ta napryamky evolyutsii: Dr. Sci. Diss. Abstract, Kyiv, 2016, 48 pp.
33. Tsymbalyuk Z.M., Mosyakin S.L. Ukr. Bot. J., 2012, 69(6): 818–831.
34. Tsymbalyuk Z.M., Mosyakin S.L. Atlas pylkovykh zeren predstavnykiv rodyn Plantaginaceae ta Scrophulariaceae (Atlas of pollen grains of representatives of Plantaginaceae and Scrophulariaceae), Kyiv: Nash format, 2013a, 276 pp.
35. Tsymbalyuk Z.M., Mosyakin S.L. Ukr. Bot. J., 2013b, 70(5): 600–609. https://doi.org/10.15407/ukrbotj70.05.600
36. Tsymbalyuk Z.M., Mosyakin S.L. Ukr. Bot. J., 2013c, 70(6): 723–731. https://doi.org/10.15407/ukrbotj70.06.723
37. Uribe-Convers S., Tank D.C. Phylogenetic revision of the genus Bartsia (Orobanchaceae): disjunct distributions correlate to independent lineages. Syst. Bot., 2016, 41(3): 672–684. https://doi.org/10.1600/036364416X692299
38. Wang H., Mill R.R., Blackmore S. Pollen morphology and infrageneric evolutionary relationships in some Chinese species of Pedicularis (Scrophulariaceae). Plant Syst. Evol., 2003, 237: 1–17. https://doi.org/10.1007/s00606-002-0188-y
39. Wang H., Yu W.B., Chen J.Q., Blackmore S. Pollen morphology in relation to floral types and pollination syndromes in Pedicularis (Orobanchaceae). Plant Syst. Evol., 2009, 277: 153–162. https://doi.org/10.1007/s00606-008-0112-1
40. Young N.D., Steiner K.E., dePamphilis C.W. The evolution of parasitism in Scrophulariaceae/Orobanchaceae: plastid gene sequences refute an evolutionary transition series. Ann. Missouri Bot. Gard., 1999, 86(4): 876–893. https://doi.org/10.2307/2666173%0A