Comparative morphometric and morphological study of the pollen of Beta trigyna, B. vulgaris and B. vulgaris subsp. maritima (Chenopodiaceae/Amaranthaceae sensu APG IV)


  • Zoya M. Tsymbalyuk Department of Systematics and Floristics of Vascular Plants, M.G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine, Kyiv
  • Daniella Ivanova Department of Plant and Fungal Diversity and Resources, Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, Sofia
  • Lyudmila M. Nitsenko Department of Systematics and Floristics of Vascular Plants, M.G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine, Kyiv


Beteae, LM, palynomorphology, sculpture, SEM, structure, UPGMA


Pollen morphology of Beta trigyna, B. vulgaris and B. vulgaris subsp. maritima, last of those studied for the first time, was investigated using light and scanning electron microscopy, based on 10 herbarium specimens. The aim of the study was to provide detailed data on the pollen characteristics of these taxa to identify similarities and differences between them. Pollen grains are pantoporate, spheroidal, circular in outline; small- and medium-sized. Exine sculpture is nanoechinate, tectum is psilate or psilate-perforate. Pore membranes are nanoechinate. Diagnostic relevance of the characters of pollen grains is discussed (pollen and pore diameters, distance between pores and between pore centres, nanoechini size and density, number of nanoechini on pore membranes, structure of columellae). UPGMA dendrograms based on palynological data support the differentiation of B. trigyna (section Corollinae), B. vulgaris and B. vulgaris subsp. maritima (section Beta). The obtained characteristics of pollen grains of Beta species can be used in spore-pollen analysis, especially in identifying the impact of human economic activity in the past.


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Albach, D. C., Tsymbalyuk, Z. M., & Mosyakin, S. L. (2021). Pollen morphology of Ellisiophyllum and Sibthorpia (Plantaginaceae, tribe Sibthorpieae) and phylogenetics of the tribe. Plant Systematics and Evolution, 307, art. 66.

Angelini, P., Bricchi, E., Gigante, D., Poponessi, S., Spina, A., & Venanzoni, R. (2014). Pollen morphology of some species of Amaranthaceae s. lat. common in Italy. Flora Mediterranea, 24, 247–272.

Ball, P. W., & Akeroyd, J. R. (1993). Beta L. In T. G. Tutin, N. A. Burges, A. O. Chater, J. R. Edmondson, V. H. Heywood, D. M. Moore, D. H. Valentine, S. M. Walters & D. A. Webb (Eds.), Flora Europaea. 2nd edition, Vol. 1. Psilotaceae to Platanaceae (pp. 110–111). Cambridge University Press, Cambridge.

Bassett, I. J., Crompton, C. W., McNeill, J., & Taschereau, P. M. (1983). The genus Atriplex (Chenopodiaceae) in Canada. Communications Branch, Agriculture Canada, Ottawa. Monograph No. 31.

Bezusko, L. G., Mosyakin, S. L., & Tsymbalyuk, Z. M. (2003). Pylok rodyny Chenopodiaceae Vent. – indykator pryrodnykh ta antropohennykh zmin roslynnoho pokryvu Ukrayiny v Holotseni [Pollen of the family Chenopodiaceae Vent. as an indicator of natural and anthropogenic changes of vegetation of Ukraine in the Holocene]. Naukovi zapysky NaUKMA. Pryrodnychi nauky, 22(3), 392–395 [in Ukrainian].

Bezusko, L. G., & Tsymbalyuk, Z. M. (2011). Palinoteka Instytutu botaniky im. M. H. Kholodnoho NAN Ukrainy [Palynotheca of the M. G. Kholodny Institute of Botany, NAS of Ukraine]. In N. M. Shiyan (Ed.), Herbaria of Ukraine (pp. 138–141). Index Herbariorum Ucrainicum. Alterpress, Kyiv [in Ukrainian].

Bezusko, L. G., Tsymbalyuk, Z. M., & Nitsenko, L. M. (2023). Participation of cultivated plants in the palynofloras of Late Holocene deposits of the Left Bank Forest-Steppe zone of Ukraine. In Ethnobotanical traditions in agronomy, pharmacy and garden design. Proceedings of the Sixth International Scientific Conference, dedicated to the Year of Indomitability of Ukraine (July 5–8, 2023) (pp. 17–24). Uman [in Ukrainian].

Biancardi, E., Panella, L. W., & Lewellen, R. T. (2012). Range of Distribution. In Beta maritima: The Origin of Beets (pp. 75–84). Springer, New York.

Brummitt, R. K., & Powell, C. E. (Eds.) (1992). Authors of plant names. A list of authors of scientific names of plants, with recommended standard forms of their names, including abbreviations. Royal Botanic Gardens, Kew.

Celenk, S., Dirmenci, T., Malyer, H., & Bicakci, A. (2008). A palynological study of the genus Nepeta L. (Lamiaceae). Plant Systematics and Evolution, 276(1), 105–123.

Česchmedjiev, I. V. (1976). Reports. In Á. Löve (Ed.), IOPB Chromosome number reports LIV. Taxon, 25(5/6): 642–643.

Dempsey, R. E., Gornall, R. J., & Bailey, J. P. (1994). Contributions to a cytological catalogue of the British and Irish flora, 4. Watsonia, 20(1), 63–66.

El Ghazali, G. E. B. (2022). Pollen morphological studies in Amaranthaceae s. lat. (incl. Chenopodiaceae) and their taxonomic significance: A review. Grana, 61(1), 1–7.

Erdtman, G. (1952). Pollen morphology and plant taxonomy. Angiosperms. Almqvist & Wiksell, Stockholm.

Ford-Lloyd, B. V., & Williams, J. T. (1975). A revision of Beta section Vulgares (Chenopodiaceae), with new light on the origin of cultivated beets. Botanical Journal of the Linnean Society, 71(2), 89–102.

Halbritter, H., Ulrich, S., Grímsson, F., Weber, M., Zetter, R., Hesse, M., Buchner, R., Svojtka, M., & Frosch-Radivo, A. (2018). Illustrated pollen terminology. 2nd edition. Springer International Publishing, Cham, XVIII.

Hammer, Ø., Harper, D. A. T., & Ryan, P. D. (2001). Past: Paleontological statistics software package for education and data analysis. Palaeontologica Electronica, 4(1), 19.

Hohmann, S., Kadereit, J. W., & Kadereit, G. (2006). Understanding Mediterranean-Californian disjunctions: molecular evidence from Chenopodiaceae–Betoideae. Taxon, 55(1), 67–78.

Iamonico, D. (2019). Remarks on the subfam. Betoideae (Chenopodiaceae/Amaranthaceae sensu APG IV). Phytotaxa, 400(3), 145–164.

Iljin, M. M. (1952). Chenopodiaceae. In M. I. Kotov (Ed.), Flora of the Ukrainian SSR [Flora URSR], Vol. 4 (pp. 260–400). Vydavnytstvo AN URSR, Kyiv [in Ukrainian].

IPNI. International Plant Names Index. (2023). The Royal Botanic Gardens, Kew, Harvard University Herbaria & Libraries and Australian National Botanic Gardens. Available at: [accessed 14 July 2023].

Jordanov, D., & Kuzmanov, B. (1966). Beta L. In D. Jordanov (Ed.), Florae Reipublicae Popularis Bulgaricae, Vol. 3 (pp. 551–553). Aedibus Academiae Scientiarum Bulgaricae, Serdicae [in Bulgarian].

Kadereit, G., Hohmann, S., & Kadereit, J. W. (2006). A synopsis of Chenopodiaceae subfam. Betoideae and notes on the taxonomy of Beta. Willdenowia, 36(1), 9–19.

Kupriyanova, L. A., & Alyoshina, L. A. (1972). Pyltsa i spory rasteniy flory evropeyskoy chasti SSSR [Pollen and spores of plants from the Flora of the European part of the USSR], Vol. 1. Nauka, Leningrad [in Russian].

Letschert, J. P. W., Lange, W., Frese, L., & Van Den Berg, R. G. (1994). Taxonomy of Beta section Beta. Journal of Sugar Beet Research, 31(1–2), 69–85.

Lewandowska, A., Milecka, K., Niedzielski, P., Czerwiński, S., & Gałka, M. (2023). Late Glacial development of lakes and wetland vegetation in a dune area in Central Poland. Acta Palaeobotanica, 63(1), 1–20.

McAndrews, J. H., & Swanson, A. R. (1967). The pore number of periporate pollen with special references to Chenopodium. Review of Palaeobotany and Palynology, 3, 105–117.

Monoszon, M. Kh. (1973). Opredelitel pyltsy vidov semeystva marevykh (posobie po sporovo-pyltsevomu analizu) [Identification key of pollen of species of the family Chenopodiaceae (manual on spore-pollen analysis)]. Nauka, Moscow [in Russian].

Mosyakin, S. L., & Fedoronchuk, M. M. (1999). Vascular plants of Ukraine. A nomenclatural checklist. M. G. Kholodny Institute of Botany, Kyiv.

Nowіcke, J. W. (1975). Pollen morphology in the order Centrospermae. Grana, 15(1–3), 51–77.

Nowіcke, J. W., & Skvarla, J. J. (1979). Pollen morphology: the potential influence in higher order systematics. Annals of the Missouri Botanical Garden, 66(4), 633–700.

Olvera, H. F., Fuentes-Soriano, S., & Hernández, E. M. (2006). Pollen morphology and systematics of Atripliceae (Chenopodiaceae). Grana, 45(3), 175–194.

Pastor, J., Fernández, I., & Díez, M. J. (1988). Números cromosómicos para la flora española, 528–543. Lagascalia, 15(1), 124–129.

POWO (2023). Plants of the World Online. Facilitated by the Royal Botanic Gardens, Kew. Available at: [accessed 14 July 2023].

Punt, W., Hoen, P. P., Blackmore, S., Nilsson, S., & Le Thomas, A. (2007). Glossary of pollen and spore terminology. Review of Palaeobotany and Palynology, 143(1–2), 1–81.

Queirós, M. (1975). Contribuiçao para o conhecimento citotaxonómico das Spermatophyta de Portugal. X. Chenopodiaceae. Boletim da Sociedade Broteriana, 49(2), 121–142.

Richards, C. M., Reeves, P. A., Fenwick, A. L., & Panella, L. (2014). Genetic structure and gene flow in Beta vulgaris subspecies maritima along the Atlantic coast of France. Genetic Resources Crop Evolution, 61, 651–662.

Runemark, H. (1996). Reports (590–678). In G. Kamari, F. Felber & F. Garbari (Eds.), Mediterranean chromosome number reports – 6. Flora Mediterranea 6, 223–243.

Shen, Y., Ford-Lloyd, B., & Newbury, H. (1998). Genetic relationships within the genus Beta determined using both PCR-based marker and DNA sequencing techniques. Heredity, 80(5), 624–632.

Skvarla, J. J., & Nowicke, J. W. (1976). Ultrastructure of pollen exine in Centrospermous families. Plant Systematics and Evolution, 126(1), 55–78.

Stevenato, P., De Biaggi, M., Skaracis, G. N., Colombo, C., Mandolino, G., & Biancardi, E. (2001). The Sea Beet (Beta vulgaris L. ssp. maritima) of the Adriatic Coast as Source of Resistance for Sugar Beet. Sugar Tech, 3(3), 77–82.

Thiers, B. (2023 [continuously updated]). Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. Available at: [accessed 14 July 2023].

Tsukada, M. (1967). Chenopod and amaranth pollen: Electron-microscopic identification. Science, 157(3784), 80–82.

Tsymbalyuk, Z. M. (2005). Palinomorfolohiya predstavnykiv rodyny Chenopodiaceae Vent. (dlya tsiley systematyky y sporovo-pylkovoho analizu) [Pollen morphology of representatives of the family Сhenopodiaceae Vent. (for taxonomy and spore-pollen analysis]. Ph.D. Thesis, M. G. Kholodny Institute of Botany, NAS of Ukraine: Kyiv [in Ukrainian].

Tsymbalyuk, Z. M. (2008). Palinomorfologicheskie osobennosti predstaviteley semeystva Chenopodiaceae [Palynomorphological peculiarities of the family Chenopodiaceae members]. Botanicheskiy Zhurnal, 93(3), 430–438 [in Russian].

Tsymbalyuk, Z. M., Celenk, S., Bell, C. D., Nitsenko, L. M., & Mosyakin, S. L. (2022a). Comparative palynomorphological study of the genus Symphoricarpos (Caprifoliaceae): exine sculpture and implications for evolution. Palynology, 46(4), 1–14.

Tsymbalyuk, Z. М., Ivanova, D., Bell, C. D., & Nitsenko, L. M. (2022b). Pollen morphology of selected taxa of Valeriana (Caprifoliaceae): taxonomic and evolutionary implications. Phytotaxa, 577(1), 57–78.

Tsymbalyuk, Z. M., Ivanova, D., & Nitsenko, L. M. (2021). Taxonomic significance of palynomorphological characteristics of selected Centranthus (Caprifoliaceae) species. Hacquetia, 20(2), 243–256.

Tsymbalyuk, Z. M., Mosyakin, S. L., & Bezusko, L. G. (2005). Novi pidkhody u rozrobtsi vyznachnyka pylku Chenopodiaceae dlya tsiley pylkovoho analizu (taksony flory Ukrayiny) [New approaches to developing an identification manual of pollen of Chenopodiaceae for pollen analysis (taxa occurring in Ukraine)]. Naukovi zapysky NaUKMA. Biolohiya ta ekolohiya, 43, 19–25 [in Ukrainian].

Tsymbalyuk, Z. M., Mosyakin, S. L., & Nitsenko, L. M. (2023a). New data on pollen morphology of the genus Camphorosma (Chenopodiaceae). Acta Palaeobotanica, 63(1), 21–31.

Tsymbalyuk, Z. M., Nitsenko, L. M., & Mosyakin, S. L. (2023b). Taxonomic significance of pollen morphology of selected taxa of Bassia, Sedobassia, Spirobassia and Neokochia (Chenopodiaceae/Amaranthaceae sensu APG IV). Palynology, 47(4), 1–16.

Van Dijk, H., & Hautekèete, N. C. (2014). Evidence of genetic change in the flowering phenology of sea beets along a latitudinal cline within two decades. Journal of Evolutionary Biology, 27(8), 1572–1581.

Yena, A. V., & Yevseyenkov, P. Ye. (2010). Beta vulgaris L. subsp. maritima (L.) Arcang. (Chenopodiaceae) – novyy takson dlya flory Kryma [Beta vulgaris L. subsp. maritima (L.) Arcang. (Chenopodiaceae) – a new taxon in the Crimean flora]. Ukrainian Botanical Journal, 67(5), 700–703 [in Russian].




How to Cite

Tsymbalyuk, Z. M. ., Ivanova, D., & Nitsenko, L. M. . (2024). Comparative morphometric and morphological study of the pollen of Beta trigyna, B. vulgaris and B. vulgaris subsp. maritima (Chenopodiaceae/Amaranthaceae sensu APG IV). Hacquetia. Retrieved from