Skip to main content Skip to main navigation menu Skip to site footer
Published: 2022-09-22

Taxonomic significance of seed and leaf micromorphology in Saponaria (Caryophyllaceae)

Department of Biology, Faculty of Science, Eskişehir Osmangazi University, Eskişehir, Turkey
Department of Biology, Faculty of Science, Eskişehir Osmangazi University, Eskişehir, Turkey
endemic mesophyll symmetry Saponaria seeds SEM taxonomy Eudicots


The seed and leaf structures of some species belonging to the genus Saponaria (Caryophyllaceae) were examined using SEM. These species were: S. calabrica, S. dalmasii (endemic), S. glutinosa, S. halophila (endemic), S. karapinarensis (endemic), S. kotschyi (endemic), S. mesogitana, S. officinalis, S. orientalis, S. pamphylica (endemic), S. picta (endemic), S. pinetorum var. elatior (endemic), S. prostrata subsp. anatolica (endemic), S. prostrata subsp. calvertii, S. prostrata subsp. prostrata (endemic), S. pumilio, and S. tridentata. The results obtained show that the micromorphology of the seed are useful to distinguish the studied species. Four groups can be distinguished using the number of seed per capsule. In addition, the shape of the seed, microsculpture, shape and margin of testa cells were found as important characteristics to identify the various species. Leaf anatomy reveals that trichome type and mesophyll symmetry define significant differences between the studied species.


  1. Amini, E., Zarre, S. & Assadi, M. (2011) Seed micro-morphology and its systematic significance in Gypsophila (Caryophyllaceae) and allied genera. Nordic Journal of Botany 29: 660–669.

  2. Arman, M. & Gholipour, A. (2013) Seed morphology diversity in some Iranian endemic Silene (Caryophyllaceae) species and their taxonomic significance. Acta Biologica Szegediensis 57 (1): 31–37.

  3. Ataşlar, E. (2004) Morphological and anatomical investigations on the Saponaria kotschyi Boiss. (Caryophyllaceae). Turkish Journal of Botany 28: 193–199.

  4. Ataşlar, E. & Ocak, A. (2017) Anatomy and micromorphology of the endemic Gypsophila osmangaziensis (Caryophyllaceae) and taxonomic contributions for the genus Gypsophila. Phytotaxa 331 (1): 84–92.

  5. Barthlott, W. (1981) Epidermal and seed surface characters of plants. Nordic Journal of Botany 1: 345–355.

  6. Barthlott, W. (1984) Microstructural features of seed surface. In: Heywood, V.H. & Moore, D.M. (Eds.) Current concepts in plant taxonomy. Academic Press, London, pp. 95–105.

  7. Bittrich, V. (1993) Caryophyllaceae. In: Kubitzki, K., Rohwer, J.G. & Bittrich, V. (Eds.) The families and genera of vascular plants, Flowering plants, Dicotyledons, Magnoliid, Hamamelid and Caryophyllid families, vol. 2. Springer-Verlag, Berlin, pp. 206–236.

  8. Bozchaloyi, S.E. & Keshavarz, M. (2014) Micro- and macromorphological study of Stellaria (Caryophyllaceae) and its closest relatives in Iran. Phytologia Balcanica 20 (2): 179–197.

  9. Bülbül, A.S., Armağan, M. & Varlık, K. (2017) Seed micromorphology of Acanthophyllum C.A.Mey. (Caryophyllaceae) Genus in Turkey. Kastamonu University, Journal of Forestry Faculty 17 (1): 215–224.

  10. Chandra, S., Rawat, D.S., Verma, S.R. & Uniyal, P. (2019) Trichome diversity of the family Caryophyllaceae from Western Himalaya and their taxonomic implication. Tropical Plant Research 6 (3): 397–407.

  11. Conti, F., Bartolucci, F., Iamonico, D. & Del Guacchio, E. (2014) A new combination in Mcneillia (Caryophyllaceae) for the Italian flora. Phytotaxa 170 (2): 139–140.

  12. Crow, G.E. (1979) The systematic significance of seed morphology in Sagina (Caryophyllaceae) under scanning electron microscopy. Brittonia 31: 52–63.

  13. Çelebioğlu, T., Favarger, C. & Huynh, K.L. (1983) Contribution a la micromorphologie de la testa des graines du genre Minuartia (Caryophyllaceae) I. Sect. Minuartia. Adansonia 4: 415–435.

  14. Çinbilgel, İ., Karadeniz, A. & Gökçeoğlu, M. (2007) Morphological and anatomical study on endemic Saponaria pamphylica Boiss. & Heldr. (Caryophyllaceae). Journal of Applied Biological Sciences 1 (2): 19–25.

  15. Dadandı, M.Y. & Yıldız, K. (2015) Seed morphology of some Silene L. (Caryophyllaceae) species collected from Turkey. Turkish Journal of Botany 39: 280–297.

  16. Dashti, A. (2018) Role of seed micro-morphology in the taxonomy of Saponaria (Caryophyllaceae). Iranian Journal of Botany 24 (2): 130–137.

  17. Dashti, A., Assadi, M. & Sharifinia, F. (2014) A new species of the genus Saponaria L. (Caryophyllaceae) in Iran. Iranian Journal of Botany 20 (2): 146–151.

  18. Davis, P.H. (1967) Flora of Turkey and the East Aegean Islands, vol. 2, Edinburgh University Press, Edinburgh, 580 pp.

  19. Davis, P.H., Mill, R.R. & Tan, K. (1988) Flora of Turkey and the East Aegean Islands, vol. 10. Edinburgh University Press, Edinburgh, 216 pp.

  20. Dillenberger, M.S. & Kadereit, J.W. (2014) Maximum polyphyly: Multiple origins and delimitation with plesiomorphic characters require a new circumscription of Minuartia (Caryophyllaceae). Taxon 63: 64–88.

  21. Echlin, P. (1968) The use of scanning reflection electron microscope in the study of plant and microbial material. Journal of the Royal Microscopical Society 88 (3): 407–418.

  22. El Naggar, S.M. (2004) The seed coat and pollen morphology of Gypsophila pilosa (Caryophyllaceae). Flora Mediterranea 14: 109–114.

  23. Erdir, M. (2009) Taxonomical, morphological and anatomical studies on Saponaria L. (Caryophyllaceae) in Turkey. PhD Thesis, Eskişehir Osmangazi University, Institute of Science and Technology, Eskişehir, 319 pp.

  24. Eröz-Poyraz, İ. & Ataşlar, E. (2010) Pollen and seed morphology of Velezia L. (Caryophyllaceae) genus in Turkey. Turkish Journal of Botany 34: 179–190.

  25. Esau, K. (1977) Anatomy of seed plants, 2nd Edition. John Wiley & Sons, Inc., 550 pp.

  26. Fawzi, N.M., Fawzy, A.M. & Mohamed, A.A. (2010) Seed morphological studies on some species of Silene L. (Caryophyllaceae). International Journal of Botany 6 (3): 287–292.

  27. Fedotova, T.A. & Ardjanova, R.R. (1992) Seed morphology in the genus Gypsophila (Caryophyllaceae). Russkii Botanicheskii Zhurnal (=Journal Russe de Botanique) 77 (5): 1–16.

  28. Gemici, Y., Tan, K., Yıldırım, H. & Pirhan, A.F. (2011) Saponaria emineana (Caryophyllaceae), a new species from inner Anatolia, Turkey. Phytotaxa 30: 45–52.

  29. Greenberg, A.K. & Donoghue, M.J. (2011) Molecular systematics and character in Caryophyllaceae. Taxon 60 (6): 1637–1652.

  30. Greuter, W. (1995) Silene (Caryophyllaceae) in Greece: a subgeneric and sectional classification. Taxon 44: 543–581.

  31. Güner, A., Özhatay, N., Ekim, T. & Başer, K.H.C. (2000) Flora of Turkey and the East Aegean Islands, vol. 11. Edinburgh University Press, Edinburgh, 680 pp.

  32. Güner, A., Aslan, S., Ekim, T., Vural, M. & Babaç, M.T. (2012) Türkiye Bitkileri Listesi (Damarlı Bitkiler). Nezahat Gökyiğit Botanik Bahçesi ve Flora Araştırmaları Derneği Yayını, Istanbul, 1290 pp.

  33. Hamzaoğlu, E., Koç, M. & Aksoy, A. (2014) A new pricking Carnation (Caryophyllaceae) grows on tuff from Turkey: Dianthus aculeatus sp. nov. Biological Diversity and Conservation 7 (2): 159–162.

  34. Hernández-Ledesma, P., Berendsohn, W.G., Borsch, T., von Mering, S., Akhani, H., Arias, S., Castañeda-Noa, I., Eggli, U., Eriksson, R., Flores-Olvera, H., Fuentes-Bazán, S., Kadereit, G., Klak, C., Korotkova, N., Nyffeler, R., Ocampo, G., Ochoterena, H., Oxelman, B., Rabeler, R.K., Sanchez, A., Schlumpberger, B.O. & Uotila, P. (2015) A taxonomic backbone for the global synthesis of species diversity in the angiosperm order Caryophyllales. Willdenowia 45: 281–383.

  35. Heywood, V.H. (1969) Scanning Electron Microscopy in the study of plant material. Micron 1: 1–14.

  36. Heywood, V.H. (1971) Scanning Electron Microscopy. Systematics Association Special, vol. 4. Academic Press, London and New York, 331 pp.

  37. Heywood, V.H. (1998) Flowering plants of the world. 3rd Edition. Oxford University Press, Oxford, 335 pp.

  38. Hong, S.P., Han, M.J. & Kim, K.J. (1999) Systematic significance of seed coat morphology in Silene L. s. str. (Sileneae-Caryophyllaceae) from Korea. Journal Plant Biology, Korea 42: 146–150.

  39. Iamonico, D. (2015) Augustea (Polycarpaeae, Caryophyllaceae), a new genus from South America. Phytotaxa 236 (1): 71–78.

  40. Iamonico, D. (2016) Nomenclatural notes on four Linnaean names in Arenaria (Caryophyllaceae). Taxon 65 (3): 610–616.

  41. Iamonico, D. (2021a) Engellaria (Caryophyllaceae), a new North American genus segregated from Stellaria. Acta Botanica Mexicana 128: e1846.

  42. Iamonico, D. (2021b) The taxonomic position of Silene multinervia (Caryophyllaceae), endemic to California and northernwestern Mexico, and description of section Californicae, sect. nov. Rhodora 122 (991): 202–213.

  43. Iamonico, D. & Domina, G. (2015) Nomenclatural notes on the Polycarpon tetraphyllum aggregate (Caryophyllaceae). Plant Biosystems 149 (4): 720–727.

  44. Kanwal, D., Abid, R. & Qaiser, M. (2012) The seed atlas of Pakistan–VI. Caryophyllaceae. Pakistan Journal of Botany 44 (1): 407–424.

  45. Kaplan, A., Çölgeçen, H. & Büyükkartal, H.N. (2009) Seed morphology and histology of some Paronychia taxa (Caryophyllaceae) from Turkey. Bangladesh Journal of Botany 38 (2): 171–176.

  46. Keshavarzi, M., Mahdavinejad, M., Sheidai, M. & Gholipour, M.A. (2015) Seed and pollen morphology of some Silene species in Iran. Phytologia Balcanica 21 (1): 7–12.

  47. Kovtonyuk, N.K. (1994) The structure of seed surface and the systematics of the Siberian Gypsophila species (Caryophyllaceae). Russkii Botanicheskii Zhurnal (=Journal Russe de Botanique) 79 (4): 48–51.

  48. Külköylüoğlu, G., Yıldız, K. & Minareci, E. (2009) Minuartia anatolica var. anatolica ve M. pestalozzae türleri üzerine morfolojik, karyolojik ve palinolojik bir çalışma. Biyoloji Bilimleri Araştırma Dergisi 2 (2): 49–57.

  49. Linnaeus, C. (1753) Species Plantarum, vol. 1. Impensis Laurentii Salvii, Stockholm, 560 pp.

  50. Mahdavi, M., Assadi, M., Fallahian, F. & Nejadsattari, T. (2012) The systematic significance of seed micro-morphology in Stellaria L. (Caryophyllaceae) and its closest relatives in Iran. Iranian Journal of Botany 18 (2): 302–310.

  51. Mantovani, A. & Vieira, R.C. (2000) Leaf micromorphology of Antarctic pearlwort Colobanthus quitensis (Kunth) Bartl. Polar Biology 23: 531–538.

  52. Metcalfe, C.R. & Chalk, L. (1950) Anatomy of the Dicotyledons 42: Caryophyllaceae, 2. Oxford University Press, London, pp. 147–152.

  53. Meyer, C.A. von (1831) Verzeichniss der Pflanzen welche während der, auf allerhöchsten Befehl, den Jahren 1829 und 1830 unternommenen Reise im Caucasus und indenProvinzen am westlichen Ufer descaspischenMeeres gefunden und engesammelt worden sind vom Dr. Carl Anton Meyer. Buchdruckerei der Kaiserlichen Akademie der Wissenschaften, St. Petersbourg, 241 pp.

  54. Miller, P. (1754) The Gardeners Dictionary 4th ed., vol. 3. Printed and Sold by John and James Rivington, London, 1019 pp.

  55. Minuto, L., Fior, S., Roccotiello, E. & Casazza, G. (2006) Seed morphology in Moehringia L. and its taxonomic significance in comparative studies within the Caryophyllaceae. Plant Systematics and Evolution 262: 189–208.

  56. Minuto, L., Roccotiello, E. & Casazza, G. (2011) New seed morphological features in Moehringia L. (Caryophyllaceae) and their taxonomic and ecological significance. Plant Biosystems 145 (1): 60–67.

  57. Mostafavi, G., Assadi, M., Nejadsattari, T., Sharifnia, F. & Mehregan, I. (2013) Seed micromorphological survey of the Minuartia species (Caryophyllaceae) in Iran. Turkish Journal of Botany 37: 446–454.

  58. Mutlu, B. (2006) Saponaria bargyliana Gombault (Caryophyllaceae): A new record for Turkey and analysis of its morphological characters with related species. Turkish Journal of Botany 30: 63–70.

  59. Ocana, M.E., Fernandez Gonzalez, I. & Pastor, J. (1997) Fruit and seed morphology in Paronychia Miller from South-West Spain. Lagascalia 19: 521–528.

  60. Pirani, A., Zarre, S., Rabeler, R., Assadi, M., Joharchi, M.R. & Oxelman, B. (2019) Systematic significance of seed morphology in Acanthophyllum (Caryophyllaceae: tribe Caryophylleae) in Iran. Phytotaxa 387 (2): 105–118.

  61. POWO (2022) Plants of the World Online | Kew Science. Available from: (accessed 9 August 2022).

  62. Rola, K., Jędrzejczak, E., Zalewska-Gałosz, J. & Nobis, M. (2019) Is the presence of simple or glandular hairs a good trait for distinguishing species in Caryophyllaceae? A case study of Arenaria serpyllifolia sensu lato in southern Poland. Acta Societatis Botanicorum Poloniae 88 (3): 3630.

  63. Sadeghi, S., Faghir, M.B., Attar, F. & Aalami, A. (2021) Achene micromorphology of the genus Potentilla L. (Rosaceae) in Iran and its systematic application. Turkish Journal of Botany 45: 15–42.

  64. Sadeghian, S., Zarre, S. & Heubl, G. (2014) Systematic implication of seed micromorphology in Arenaria (Caryophyllaceae) and allied genera. Flora 209: 513–529.

  65. Sharma, S. (2016) SEM (Scanning Electron Microscopy) studies on Dianthus (Caryophyllaceae). Journal Indian Botanical Society 95 (3–4): 288–292.

  66. Simmler, G. (1910) Monographie der Gattung Saponaria. Denkschriften der Kaiserlichen Akademie der Wissenschaften / Mathematisch-Naturwissenschaftliche Classe 85: 433–509.

  67. Stearn, W.T. (1992) Botanical Latin: History, grammar, syntax, terminology and vocabulary. 4th Edition. David and Charles Publishers, England, 546 pp.

  68. Thiers, B. (2022 [continuously updated]) Index Herbariorum: A global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. Available from: (accessed 9 August 2022).

  69. Ulukuş, D. (2018) Morphology, anatomy and palynology study on Turkish endemic species Saponaria karapinarensis (Caryophyllaceae). Phytotaxa 374 (1): 80–86.

  70. Ullah, F., Zafar, M., Ahmad, M., Shah, S.N., Razzaq, A., Sohail, A., Zaman, W., Çelik, A., Ayaz, A. & Sultana, S. (2018a) A systematic approach to the investigation of foliar epidermal anatomy of subfamily Caryophylloideae (Caryophyllaceae). Flora 246–247: 61–70.

  71. Ullah, F., Zafar, M., Amhad, M., Sultana, S., Ullah, A., Shah, S.N., Butt, M.A. & Mir, S. (2018b) Taxonomic implications of foliar epidermal characteristics in subfamily Alsinoideae (Caryophyllaceae). Flora 242: 31–44.

  72. Vural, M., Duman, H., Aytaç, Z. & Adigüzel, N. (2006) Saponaria karapinarensis, Senecio salsuginea and Centaurea tuzgoluensis, three new species from Central Anatolia, Turkey. Belgian Journal of Botany 139: 252–260.

  73. Wofford, B.E. (1981) External seed morphology of Arenaria (Caryophyllaceae) of the southeastern United States. Systematic Botany 6: 126–135.

  74. Wyatt, R. (1984) Intraspecific variation in seed morphology of Arenaria uniflora (Caryophyllaceae). Systematic Botany 9: 423–431.

  75. Yıldız, K. (2002) Seed morphology of Caryophyllaceae species from Turkey (North Anatolia). Pakistan Journal of Botany 34 (2): 161–171.

  76. Yıldız, K. & Çırpıcı, A. (1998) Seed morphological studies of Silene L. from Turkey. Pakistan Journal of Botany 30 (2): 173–188.