Flower and fruit micromorphology and anatomy in Circaea lutetiana (Onagraceae)

Authors

Keywords:

epidermal trichomes, gynoecium, pyrenarium, seed coat, vascular anatomy

Abstract

Questions: In the genus Circaea, many important floral and fruit traits remain insufficiently studied which precludes congruence between molecular and non-molecular data in systematics and hampers understanding of morphological evolution within the Onagraceae family. Location: fixed specimens of Circaea lutetiana flowers and fruits from Lviv and Zakarpattia regions. Materials and methods: permanent and temporal preparations, light microscope technique, gynoecium micromorphology, vascular anatomy, fruit and seed coat anatomy. Nomenclature: Wagner et al. 2007, POWO 2026. Results: The investigation revealed that sepals, petals, stamens, nectary disc, and the style are all inserted at the top end of the ovary neck, previously termed the floral tube or hypanthium. The bilocular ovary is composed of synascidiate and symplicate zones with apical septa in the latter. Each ovule is attached to the top of synascidiate zone and is provided by trans-septal vascular bundles from both septal veins. Two dorsal and two septal veins supply the inferior ovary till the top of the ovary neck, where they branch into traces of floral organs and vascular bundles supplying the nectary. After anthesis, ovary neck separates from the locular part and falls away together with the attached floral organs. Fruit wall is largely parenchymatous. It is composed of unilayered exocarp, homogenous compact mesocarp and unlignified endocarp. During fruit development, the inner 1–2 subepidermal layers of the mesocarp becomes to lignify. Two types of one-celled epidermal trichomes were found on the ovary wall: lignified stiff uncinate (hooked) and glandular clavate-tipped hairs. Seed coat is composed of a multilayered testa and bilayered tegmen. The mesotesta consists of 5–6 layers; endotesta and exotegmen are lignified. The fruit type is defined as a two-stoned pyrenarium based on the ovary structure and fruit wall anatomy. Conclusions: Our study provides deeper insight into the internal structure of flower and fruit of Circaea lutetiana. The inner gynoecium structure, fruit wall and seed coat anatomy were examined for the first time in the genus Circaea. Newly revealed traits, such as lignified subepidermal layer of the mesocarp, trans-septal ovule supply, and glandular trichomes, also occur in Epilobium and Oenothera species, offering new perspectives on the morphological evolution within the Onagraceae.

References

Bojňanský, V. & Fargašová, A. (2007). Atlas of seeds and fruits of Central and East-European flora the Carpathian Mountains Region. Dordrecht: Springer, 1046 p. https://doi.org/10.1007/978-1-4020-5362-7

Boufford, D.E. (1982). The systematics and evolution of Circaea (Onagraceae). Annals of the Missouri Botanical Garden 69 (4): 804–994. https://doi.org/10.2307/2398997

Boufford, D.E. (1987). Pollination biology of Circaea (Onagraceae). Acta Phytotaxonomica et Geobotanica 38: 133–154. https://doi.org/10.18942/bunruichiri.KJ00002992247

Boufford, D.E., Crisci, J.V., Tobe, H. & Hoch, P.C. (1990). A cladistic analysis of Circaea (Onagraceae). Cladistics 6 (2): 171–182. https://doi.org/10.1111/j.1096-0031.1990.tb00534.x.

Braune, W., Leman, A. & Taubert, H. (1991). Pflanzenanatomisches Praktikum I: Zur Einführung in die Anatomie der Vegetationsorgane der Samenpflanzen, 6 Aufl. Jena: Gustav Fischer, 283 S.

Conti, E., Fischbach, A. & Sytsma, K.J. (1993). Tribal relationships in Onagraceae: implications from rbcL sequence data. Annals of the Missouri Botanical Garden 80 (3): 672–685. https://doi.org/10.2307/2399853

Corner, E.J.H. (1976). The seeds of dicotyledons 1. Cambridge: University Press, 312 p.

Eichler, A.W. (1878). Blüthendiagramme 2. Leipzig: Engelmann, 575 p. https://doi.org/10.5962/bhl.title.12323

Endress, P.K. (2011). Evolutionary diversification of the flowers in angiosperms. American Journal of Botany 98 (3): 370–396. https://doi.org/10.3732/ajb.1000299

Eyde, R.H. (1978). Reproductive structures and evolution in Ludwigia (Onagraceae). II. Fruit and Seed. Annals of the Missouri Botanical Garden 65 (2): 656–675. https://doi.org/10.2307/2398865

Eyde, R.H. (1982). Evolution and systematics of the Onagraceae: Floral Anatomy. Annals of the Missouri Botanical Garden 69 (4): 735–747. https://doi.org/10.2307/2398993

Fedoronchuk, M.M. (2023). Ukrainian flora checklist. 7: family Caprifoliaceae s. l. (incl. Dipsacaceae, Linnaeaceae, Valerianaceae), Viburnaceae s. l. (incl. Adoxaceae, Sambucaceae) (Dipsacales, Angiosperms), and Lythraceae (incl. Punicaceae, Trapaceae), Onagraceae, Myrtaceae (Myrtales, Angiosperms). Chornomorski Botanical Journal 19 (3): 243–271. https://doi.org/10.32999/ksu1990-553X/2023-19-3-1 (in Ukrainian)

Gorb, E. & Gorb, S. (2002). Contact separation force of the fruit burrs in four plant species adapted to dispersal by mechanical interlocking. Plant Physiology and Biochemistry 40 (4): 373–381. https://doi.org/10.1016/S0981-9428(02)01381-5

Islam, M., Gottlieb, A. Maher, M., Scher, J. & Redford A.J. (2024). Fruit and Seed Family ID, Edition 2. USDA APHIS PPQ Identification Technology Program. Fort Collins, CO. [30/04/2026] https://idtools.org/seed_families/index.cfm?packageID=2246&entityID=57965

Kubitzki, K. (2007). Introduction to the groups treated in this volume. In: Kubitzki, K. (ed.) The Families and Genera of Vascular Plants, Vol. 9. Flowering Plants, Eudicots: 1–20. Berlin-Heidelberg: Springer. https://doi.org/10.1007/978-3-540-32219-1_1

Levin, R.A., Wagner, W.L., Hoch, P.C., Nepokroeff, M., Pires, J.C., Zimmer, E.A. & Sytsma, K.J. (2003). Family-level relationships of Onagraceae based on chloroplast rbcL and ndhL data. American Journal of Botany 90 (1): 107–115. https://doi.org/10.3732/ajb.90.1.107

Luo, Y., He, J., Lyu, R., Xiao, J., Li, W., Yao, M., Pei, L., Cheng, J., Li, J. & Xie, L. (2021). Comparative analysis of complete chloroplast genomes of 13 species in Epilobium, Circaea, and Chamaenerion and insights into phylogenetic relationships of Onagraceae. Frontiers in Genetics 12: 730495. https://doi.org/10.3389/fgene.2021.730495

Novikov, A. & Sup-Novikova, M. (2021). Modified staining protocol with Safranin O and Astra Blue for the plant histology. Plant Introduction 89/90: 110–113. https://doi.org/10.46341/PI2021005

Odintsova, A.V. (2022). Morphogenesis of fruit as a subject matter for the carpological studies. Ukrainian Botanical Journal 79 (3): 169–183 (in Ukrainian). https://doi.org/10.15407/ukrbotj79.03.169

Odintsova, A. (2023). Method for drafting a morpho-anatomical description of the fruit using light microscopy Visnyk of the Lviv University. Series Biology 89: 3–19. (іn Ukrainian). https://doi.org/10.30970/vlubs.2023.89.01

Odintsova, A.V. & Klimovych, N.B. (2017). Anatomical and morphological fruit structure in Epilobium hirsutum and E. angustifolium (Onagraceae). Ukrainian Botanical Journal 74 (6): 582–593. (in Ukrainian). https://doi.org/10.15407/ukrbotj74.06.582

Overson, R.P., Johnson, M.G., Bechen, L.L., Kinosian, S.P., Douglas, N.A., Fant, J.B., Hoch, P.C., Levin, R.A., Moore, M.J., Raguso, R.A., Wagner, W.L., Skogen, K.A. & Wickett, N.J. (2023). A phylogeny of the evening primrose family (Onagraceae) using a target enrichment approach with 303 nuclear loci. BMC Ecology and Evolution 23: 66. https://doi.org/10.1186/s12862-023-02151-9

Ou, C.-Y., Chang, C.-H., Yeh, T.-Y., Chung, K.-F., Hoch, P.C. & Liu, S.-H. (2025). Novel perspectives on plastome evolution in Onagraceae. AoB PLANTS 17 (3): plaf025. https://doi.org/10.1093/aobpla/plaf025

POWO (2026). Plants of the World Online. Facilitated by the Royal Botanic Gardens, Kew. URL: https://powo.science.kew.org/ [02/03/2026].

Pradhan Mitra, P. & Loqué, D. (2014). Histochemical staining of Arabidopsis thaliana secondary cell wall elements. Journal of Visualized Experiments 87: e51381. https://doi.org/10.3791/51381

Raven, P.H. (1963). Circaea in the British Isles. Watsonia 5 (5): 262–272.

Rostanski, K., Rostanski, A., Gerold-Śmietaсska, I. & Włsowicz, P. (2010). Evening-Primroses (Oenothera) occurring in Europe. Katowice-Krakуw: W.Szafer Institute of Botany, Polish Academy of Sciences. 157 p.

Soukup, A. & Tylová, E. (2019). Essential methods of plant sample preparation for light microscopy. In: Cvrčková, F. & Žárský, V. (eds.) Plant Cell Morphogenesis. Methods in Molecular Biology, Vol. 1992: 1–26. New York, NY: Humana. https://doi.org/10.1007/978-1-4939-9469-4_1.

Spjut, R.W. (1994). A systematic treatment of fruit types. Memoirs of the New York Botanical Garden 70: 1‒182. http://www.worldbotanical.com/fruit_types.htm

Strgulc Krajšek, S., Kreft, S., Klandnik, A., Draslar, K., Jogan, N. & Dermastia, M. (2011). Morphology and glandular activity of unicellular trichomes of Epilobium hirsutum. Biologia Plantarum 55 (1): 149–152. https://doi.org/10.1007/s10535-011-0020-z

Tan, Y.-Y., Li, D.-S., Hua, J., Luo, S.-H., Liu, Y. & Li, S.-H. (2017). Localization of a defensive volatile 4-hydroxy-4-methylpentan-2-one in the capitate glandular trichomes of Oenothera glazioviana. Plant Diversity 39 (3): https://doi.org/10.1016/j.pld.2017.05.004

Tobe, H., Wagner, W.L. & Chin, H.-C. (1987). A systematic and evolutionary study of Oenothera (Onagraceae): seed coat anatomy. Botanical Gazette (Crawfordsville) 148 (2): 235–257.

Tobe, H., Raven, P.H. & Peng C.-I. (1988). Seed coat anatomy and relationships of Ludwigia sects. Microcarpium, Dantia, and Miquelia (Onagraceae) and notes on fossil seeds of Ludwigia from Europe. Botanical Gazette (Crawfordsville) 149 (4): 450–457.

Wagner, W.L. & Hoch, P.C. (2005). Onagraceae. The Evening Primrose Family website. URL: https://naturalhistory2.si.edu/botany/onagraceae/ [27/02/2026] Wagner, W.L., Hoch, P.C. & Raven, P.H. (2007). Revised classification of the Onagraceae. Systematic Botany Monographs 83: 1–240. https://repository.si.edu/handle/10088/7611

Xie, L., Wagner, W., Ree, R., Berry, P. & Wen, J. (2009). Molecular phylogeny, divergence time estimates, and historical biogeography of Circaea (Onagraceae) in the Northern Hemisphere. Molecular Phylogenetics and Evolution 53 (3): 995–1009. https://doi.org/10.1016/j.ympev.2009.09.009

Published

2026-06-30

How to Cite

Odintsova, A. (2026). Flower and fruit micromorphology and anatomy in Circaea lutetiana (Onagraceae). CHORNOMORSKI BOTANICAL JOURNAL, 22(2), 186–198. Retrieved from https://cbj.kspu.edu/index.php/cbj/article/view/855