The seeds of many species in the order Caryophyllales exhibit surface protuberances called tubercles. While tubercle shape and distribution have often been proposed as taxonomic criteria, paradoxically, their description has primarily relied on adjectives, with quantitative data on tubercle width, height, and other measurements lacking in the literature. Recently, a quantitative analysis of seed surface tubercles based on tubercle width, height, and curvature values (maximum and average curvature, and maximum to average curvature ratio) was proposed and applied to individual populations of a total of 31 species, with 12 belonging to Silene subg. Behenantha and 19 to S. subg. Silene. Tubercles were classified into two categories: echinate and rugose. Echinate tubercles exhibited higher values of height and curvature, and lower width, and were more prevalent in species of S. subg. Behenantha, while the rugose type was more abundant in S. subg. Silene. This work explored infraspecific differences in tubercle size and shape. For this, measurements of tubercle width, height and curvature were applied to 31 populations of eight species of Silene. Significant differences between populations were observed for most of the species examined. A particular tubercle type, previously described as umbonate or mammillate, was identified in S. nocturna seeds, characterized by high curvature values.

Departamento de Ciencias Ambientales y Recursos Naturales University of Alicante San Vicente del Raspeig 03690 Alicante Spain

Departamento de Matemáticas Facultad de Ciencias Universidad de Salamanca Plaza de la Merced 1 4 37008 Salamanca Spain

Instituto de Recursos Naturales y Agrobiología Consejo Superior de Investigaciones Científicas Cordel de Merinas 40 37008 Salamanca Spain

Plant Developmental Genetics Institute of Biophysics v v i Academy of Sciences of the Czech Republic Královopolská 135 61265 Brno Czech Republic

See more in PubMed

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., et al. A taxonomic backbone for the global synthesis of species diversity in the angiosperm order Caryophyllales. Willdenowia. 2015;45:281–383. doi: 10.3372/wi.45.45301. DOI

Bittrich V. Caryophyllaceae. In: Kubitzki K., Rohwer J.C., Bittrich V., editors. The Families and Genera of Vascular Plants. Volume 2. Springer; Berlin/Heidelberg, Germany: 1993. pp. 206–236. (Magnoliid, Hamamelid, and Caryophyllid Families)

Johri B.M., Ambegaokar K.B., Srivastava P.S. Comparative Embryology of Angiosperms. Volume 1 Springer; Berlin/Heidelberg, Germany: 1992.

Martin A.C. The comparative internal morphology of seeds. Am. Midl. Nat. 1946;36:513–660. doi: 10.2307/2421457. DOI

Ullah F., Papini A., Shah S.N., Zaman W., Sohail A., Iqbal M. Seed micromorphology and its taxonomic evidence in subfamily Alsinoideae (Caryophyllaceae) Microsc. Res. Tech. 2019;82:250–259. doi: 10.1002/jemt.23167. PubMed DOI

Wofford B.E. External seed morphology of Arenaria (Caryophyllaceae) of the Southeastern United States. Syst. Bot. 1981;6:126. doi: 10.2307/2418543. DOI

Wyatt R. Intraspecific variation in seed morphology of Arenaria uniflora (Caryophyllaceae) Syst. Bot. 1984;9:423. doi: 10.2307/2418791. DOI

Arabi Z., Ghahremaninejad F., Rabeler R.K., Heubl G., Zarre S. Seed micromorphology and its systematic significance in tribe Alsineae (Caryophyllaceae) Flora. 2017;234:41–59. doi: 10.1016/j.flora.2017.07.004. DOI

Sadeghian S., Zarre S., Heubl G. Systematic implication of seed micromorphology in Arenaria (Caryophyllaceae) and allied genera. Flora. 2014;209:513–529. doi: 10.1016/j.flora.2014.07.004. DOI

Poyraz I.E., Ataşlar E. Pollen and seed morphology of Velezia L. (Caryophyllaceae) genus in Turkey. Turk. J. Bot. 2010;34:179–190. doi: 10.3906/bot-0910-185. DOI

Amini E., Zarre S., Assadi M. Seed micro-morphology and its systematic significance in Gypsophila (Caryophyllaceae) and allied genera. Nord. J. Bot. 2011;29:660–669. doi: 10.1111/j.1756-1051.2011.01208.x. DOI

Kovtonyuk N.K. The structure of seed surface and the systematics of the Siberian Gypsophila species (Caryophyllaceae) Bot Zhurn. 1994;79:48–51.

Mostafavi G., Assadi M., Nejadsattari T., Sharifnia F., Mehregan I. Seed micromorphological survey of the Minuartia species (Caryophyllaceae) in Iran. Turk. J. Bot. 2013;37:446–454. doi: 10.3906/bot-1201-33. DOI

Minuto L., Fior S., Roccotiello E., Casazza G. Seed morphology in Moehringia L. and its taxonomic significance in comparative studies within the Caryophyllaceae. Plant Syst. Evol. 2006;262:189–208. doi: 10.1007/s00606-006-0458-1. DOI

Kaplan A., Çölgeçen H., Büyükkartal H.N. Seed morphology and histology of some Paronychia taxa (Caryophyllaceae) from Turkey. Bangladesh J. Bot. 2009;38:171–176. doi: 10.3329/bjb.v38i2.5142. DOI

Crow G.E. The systematic significance of seed morphology in Sagina (Caryophyllaceae) Under Scanning Electron Microscopy. Brittonia. 1979;31:52–63. doi: 10.2307/2806673. DOI

Hong S.-P., Han M.-J., Kim K.-J. Systematic significance of seed coat morphology in Silene L. s. str. (Sileneae-Caryophyllaceae) from Korea. J. Plant Biol. 1999;42:146–150. doi: 10.1007/BF03031023. DOI

Alonso M.Á., Crespo M.B., Martínez-Azorín M., Mucina L. Spergularia hanoverensis (Caryophyllaceae): Validation and Recircumscription of a Misinterpreted Species from South Africa. Plants. 2023;12:2481. doi: 10.3390/plants12132481. PubMed DOI PMC

Volponi C.R. Contribución a la espermatología de especies argentinas de Stellaria (Caryophyllaceae) Bol. Soc. Argent. Bot. 1986;24:283–294.

Volponi C.R. Stellaria yungasensis (Caryophyllaceae), una nueva cita para Argentina. An. Soc. Cient. Argent. 1992;222:69–72.

Volponi C.R. Stellaria cuspidata (Caryophyllaceae) and some related species in the Andes. Willdenowia. 1993;23:193–209.

Mahdavi M., Asadi M., Fallahian F., Nejadsattari T. The systematic significance of seed micro-morphology in Stellaria L. (Caryophyllaceae) and its closest relatives in Iran. Iran. J. Bot. 2012;18:302–310.

Sukhorukov A.P., Nilova M.V., Kushunina M., Mazei Y., Klak C. Evolution of seed characters and of dispersal modes in Aizoaceae. Front. Plant Sci. 2023;14:1140069. doi: 10.3389/fpls.2023.1140069. PubMed DOI PMC

Sukhorukov A.P., Zhang M.-L., Kushunina M., Nilova M.V., Krinitsina A., Zaika M.A., Mazei Y. Seed characters in Molluginaceae (Caryophyllales): Implications for taxonomy and evolution. Bot. J. Linn. Soc. 2018;187:167–208. doi: 10.1093/botlinnean/boy021. DOI

Danin A., Domina G., Raimondo F.M. Microspecies of the Portulaca oleracea aggregate found on major Mediterranean islands (Sicily, Cyprus, Crete, Rhodes) Fl. Medit. 2008;18:89–107.

Ocampo G., Travis Columbus J. Molecular phylogenetics, historical biogeography, and chromosome number evolution of Portulaca (Portulacaceae) Mol. Phylogenet. Evol. 2012;63:97–112. doi: 10.1016/j.ympev.2011.12.017. PubMed DOI

Martín-Gómez J.J., Rodríguez-Lorenzo J.L., Juan A., Tocino Á., Janousek B., Cervantes E. Seed Morphological Properties Related to Taxonomy in Silene L. Species. Taxonomy. 2022;2:298–323. doi: 10.3390/taxonomy2030024. DOI

Martín-Gómez J.J., Rodríguez-Lorenzo J.L., Tocino Á., Janoušek B., Juan A., Cervantes E. The Outline of Seed Silhouettes: A Morphological Approach to Silene (Caryophyllaceae) Plants. 2022;11:3383. doi: 10.3390/plants11233383. PubMed DOI PMC

Cervantes E., Rodríguez-Lorenzo J.L., Martín-Gómez J.J., Tocino Á. Curvature Analysis of Seed Silhouettes in Silene L. Plants. 2023;12:2439. doi: 10.3390/plants12132439. PubMed DOI PMC

Rodríguez-Lorenzo J.L., Martín-Gómez J.J., Juan A., Tocino Á., Cervantes E. Quantitative Analysis of Seed Surface Tubercles in Silene Species. Plants. 2023;12:3444. doi: 10.3390/plants12193444. PubMed DOI PMC

Tabaripur R., Koohdar F., Sheidai M., Gholipour A. Intra-specific variations in Silene: Morphometry and micromorphometry analyses. Afr. J. Biotech. 2013;12:5208–5217.

Boissier E. In: Flora Orientalis. Georg H., editor. Volume 1. Bibliopolam; Basel, Switzerland: 1867. pp. 567–656.

Rohrbach P. Monographic der Gattung Silene. Verlag von Engelmann; Leipzig, Germany: 1869. [(accessed on 6 March 2024)]. pp. 1–249. Available online: https://www.biodiversitylibrary.org/bibliography/15462.

Jafari F., Zarre S., Gholipour A., Eggens F., Rabeler R.K., Oxelman B. A new taxonomic backbone for the infrageneric classification of the species-rich genus Silene (Caryophyllaceae) Taxon. 2020;69:337–368. doi: 10.1002/tax.12230. DOI

Oxelman B., Lidén M. Generic boundaries in the tribe Sileneae (Caryophyllaceae) as inferred from nuclear rDNA sequences. Taxon. 1995;44:525–542. doi: 10.2307/1223498. DOI

Eggens F., Popp M., Nepokroeff M., Wagner W.L., Oxelman B. The origin and number of introductions of the Hawaiian endemic Silene species (Caryophyllaceae) Amer. J. Bot. 2007;94:210–218. doi: 10.3732/ajb.94.2.210. PubMed DOI

Naciri Y., Du Pasquier P.-E., Lundberg M., Jeanmonod D., Oxelman B. A phylogenetic circumscription of Silene sect. Siphonomorpha (Caryophyllaceae) in the Mediterranean Basin. Taxon. 2017;66:91–108. doi: 10.12705/661.5. DOI

Rodríguez-Lorenzo J.L., Martín-Gómez J.J., Tocino Á., Juan A., Janoušek B., Cervantes E. New Geometric Models for Shape Quantification of the Dorsal View in Seeds of Silene Species. Plants. 2022;11:958. doi: 10.3390/plants11070958. PubMed DOI PMC

Martín-Gómez J.J., Rodríguez-Lorenzo J.L., Janoušek B., Juan A., Cervantes E. Comparison of Seed Images with Geometric Models, an Approach to the Morphology of Silene (Caryophyllaceae) Taxonomy. 2023;3:109–132. doi: 10.3390/taxonomy3010010. DOI

Greuter W. Silene (Caryophyllaceae) in Greece: A Subgeneric and Sectional Classification. Taxon. 1995;44:543–581. doi: 10.2307/1223499. DOI

Runyeon H., Prentice H.C. Patterns of seed polymorphism and allozyme variation in the bladder campions, Silene vulgaris and Silene uniflora (Caryophyllaceae) Can. J. Bot. 1977;75:1868–1886. doi: 10.1139/b97-899. DOI

Prentice H.C. Numerical analysis of infraspecific variation in European Silene alba and S. dioica (Caryophyllaceae) Bot. J. Linn. Soc. 1979;78:181–212. doi: 10.1111/j.1095-8339.1979.tb02193.x. DOI

Prentice H.C. Climate and clinal variation in seed morphology of the white campion, Silene latifolia (Caryophyllaceae) Bot. J. Linn. Soc. 1986;27:179–189. doi: 10.1111/j.1095-8312.1986.tb01732.x. DOI

Dadandi M.Y., Yildiz K. Seed morphology of some Silene L. (Caryophyllaceae) species collected from Turkey. Turk. J. Bot. 2015;39:280–297. doi: 10.3906/bot-1307-35. DOI

Ocampo G. Morphological characterization of seeds in Portulacaceae. Phytotaxa. 2013;141:1–24. doi: 10.11646/phytotaxa.141.1.1. DOI

Rasband W.S. ImageJ. U.S. National Institutes of Health: Bethesda, MD, USA, 2018. [(accessed on 19 April 2024)]; Available online: http://imagej.nih.gov/ij/

Cervantes E., Tocino A. Geometric analysis of Arabidopsis root apex reveals a new aspect of the ethylene signal transduction pathway in development. J. Plant Physiol. 2005;162:1038–1045. doi: 10.1016/j.jplph.2004.10.013. PubMed DOI

Noriega A., Tocino A., Cervantes E. Hydrogen peroxide treatment results in reduced curvature values in the Arabidopsis root apex. J. Plant Physiol. 2009;166:554–558. doi: 10.1016/j.jplph.2008.07.009. PubMed DOI

Martín-Gómez J.J., Rewicz A., Goriewa-Duba K., Wiwart M., Tocino Á., Cervantes E. Morphological description and classification of wheat kernels based on geometric models. Agronomy. 2019;9:399. doi: 10.3390/agronomy9070399. DOI

Cervantes E., Martín-Gómez J.J., Espinosa-Roldán F.E., Muñoz-Organero G., Tocino Á., Cabello Sáenz de Santamaría F. Seed apex curvature in key Spanish grapevine cultivars. Vitic. Data J. 2021;3:e66478. doi: 10.3897/vdj.3.e66478. DOI

R Core Team . R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing; Vienna, Austria: 2020.

Campbell G., Skillings J.H. Nonparametric Stepwise Multiple Comparison Procedures. J. Am. Stat. Assoc. 1985;80:998–1003. doi: 10.1080/01621459.1985.10478216. DOI

Sokal R.R., Braumann C.A. Significance Tests for Coefficients of Variation and Variability Profiles. Syst. Zool. 1980;29:50. doi: 10.2307/2412626. DOI