Infraspecific Variation in Silene Seed Tubercles.

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.

Quantitative Analysis of Seed Surface Tubercles in Silene Species.

In the Caryophyllaceae, seed surfaces contain cell protrusions, of varying sizes and shapes, called tubercles. Tubercles have long been described in many species, but quantitative analyses with measurements of size and shape are lacking in the literature. Based on optical photography, the seeds of Silene were classified into four types: smooth, rugose, echinate and papillose. Seeds in each of these groups have characteristic geometrical properties: smooth seeds lack tubercles and have the highest values of circularity and solidity in their lateral views, while papillose seeds have the largest tubercles and lowest values of circularity and solidity both in lateral and dorsal views. Here, tubercle width, height and slope, maximum and mean curvature values and maximum to mean curvature ratio were obtained for representative seeds of a total of 31 species, 12 belonging to Silene subg. Behenantha and 19 to S. subg. Silene. The seeds of the rugose type had lower values of curvature. Additionally, lower values of curvature were found in species of S. subg. Silene in comparison with S. subg. Behenantha. The seeds of S. subg. Behenantha had higher values of tubercle height and slope and higher values of maximum and average curvature and maximum to mean curvature ratio.

Curvature Analysis of Seed Silhouettes in Silene L.

The application of seed morphology to descriptive systematics requires methods for shape analysis and quantification. The complexity of lateral and dorsal views of seeds of Silene species is investigated here by the application of the Elliptic Fourier Transform (EFT) to representative seeds of four morphological types: smooth, rugose, echinate and papillose. The silhouettes of seed images in the lateral and dorsal views are converted to trigonometric functions, whose graphical representations reproduce them with different levels of accuracy depending on the number of harmonics. A general definition of seed shape in Silene species is obtained by equations based on 40 points and 20 harmonics, while the detailed representation of individual tubercles in each seed image requires between 100 and 200 points and 60-80 harmonics depending on their number and complexity. Smooth-type seeds are accurately represented with a low number of harmonics, while rugose, echinate and papillose seeds require a higher number. Fourier equations provide information about tubercle number and distribution and allow the analysis of curvature. Further estimation of curvature values in individual tubercles reveals differences between seeds, with higher values of curvature in S. latifolia, representative of echinate seeds, and lower in S. chlorifolia with rugose seeds.

The Outline of Seed Silhouettes: A Morphological Approach to Silene (Caryophyllaceae).

Seed morphology is an important source of information for plant taxonomy. Nevertheless, the characters under study are diverse, and a simple, unified method is lacking in the literature. A new method for the classification of seeds of the genus Silene based on optical images and image analysis has recently been described on the basis of morphological measurements of the lateral seed views. According to the outline of their silhouettes, seeds from 52 species (49 of Silene and three related species) were classified in three groups: smooth, rugose and echinate, revealing remarkable differences between these groups. This methodology has been applied here to 51 new species, making a total of 100 species of Silene analyzed so far. According to our data, a new group was described, termed papillose. The results showed morphological differences between the four mentioned seed groups, with reduced values of circularity for dorsal and lateral seed views in the papillose and echinate groups and reduced values of solidity in the papillose seeds. The method was applied to the analysis of individual as well as to average seed silhouettes and some of the differences between groups were maintained in both cases.

Seed Morphology in Species from the Silene mollissima Aggregate (Caryophyllaceae) by Comparison with Geometric Models.

The description of seed shape by comparison with geometric models allows shape quantification, providing the means for an accurate comparison between different species or populations. Geometric models described for the lateral and dorsal views of the seeds of Silene species are applied to the quantification of the shape in the seeds belonging to twenty populations of the eleven taxa of S. mollissima aggregate. Cardioid models LM1, LM5 and LM6 adjust differentially to the lateral views of the seeds, while models DM1, DM5 and DM6 are applied to the dorsal views of the seeds. Quantification of the lateral view of seeds with LM5 results in two groups of species of different geographic origin. The seeds more resembling DM5 include S. andryalifolia, S. badaroi, S. gazulensis, S. hifacensis and S. tomentosa, i.e., the taxa with a continental distribution from southern Spain to northern Italy; in contrast, the group of seeds with lower similarity to DM5 includes those from species in northern Africa and the Mediterranean Tyrrhenian islands: S. auricolifolia, S. hicesiae, S. ichnusae, S. mollissima, S. oenotriae and S. velutina. The description of the seed shape based on geometric models contributes to investigating the relationships between related species and constitutes a promising technique for taxonomy.

New Geometric Models for Shape Quantification of the Dorsal View in Seeds of Silene Species.

The description of shape in Silene seeds is based on adjectives coined by naturalists in the 19th century. The expressions reniform, dorso plana, and dorso canaliculata were applied in reference to lateral or dorsal views of seeds, but the characters described can be submitted now to an analytical description by quantitative methods, allowing shape quantification and the comparison between species or populations. A quantitative morphological analysis is based on the comparison with geometric models that adjust to the shape of seeds. Morphological analysis of the dorsal view of Silene seeds based on geometric models is applied here to 26 seed populations belonging to 12 species. According to their dorsal views, the seeds are classified as convex and non-convex. New geometric models are presented for both types, including figures such as super-ellipses and modified ellipses. The values of J index (percent of similarity of a seed image with the model) are obtained in representative seed samples from diverse populations and species. The quantitative description of seed shape based on the comparison with geometric models allows the study of variation in shape between species and in populations, as well as the identification of seeds in Silene species. The method is of application to other plant species.

Geometric Models for Seed Shape Description and Quantification in the Cactaceae.

Seed shape in species of the Cactaceae is described by comparison with geometric models. Three new groups of models are presented, two for symmetric seeds, and a third group for asymmetric seeds. The first two groups correspond, respectively, to superellipses and the combined equations of two semi-ellipses. The third group contains models derived from the representation of polar equations of Archimedean spirals that define the shape of asymmetric seeds in genera of different subfamilies. Some of the new models are geometric curves, while others are composed with a part resulting from the average silhouettes of seeds. The application of models to seed shape quantification permits the analysis of variation in seed populations, as well as the comparison of shape between species. The embryos of the Cactaceae are of the peripheral type, strongly curved and in contact with the inner surface of the seed coat. A relationship is found between seed elongation and the models, in which the genera with elongated seeds are represented by models with longer trajectories of the spiral. The analysis of seed shape opens new opportunities for taxonomy and allows quantification of seed shape in species of the Cactaceae.

Seed Geometry in the Vitaceae.

The Vitaceae Juss., in the basal lineages of Rosids, contains sixteen genera and 950 species, mainly of tropical lianas. The family has been divided in five tribes: Ampelopsideae, Cisseae, Cayratieae, Parthenocisseae and Viteae. Seed shape is variable in this family. Based on new models derived from equations representing heart and water drop curves, we describe seed shape in species of the Vitaceae. According to their similarity to geometric models, the seeds of the Vitaceae have been classified in ten groups. Three of them correspond to models before described and shared with the Arecaceae (lenses, superellipses and elongated water drops), while in the seven groups remaining, four correspond to general models (waterdrops, heart curves, elongated heart curves and other elongated models) and three adjust to the silhouettes of seeds in particular genera (heart curves of Cayratia and Pseudocayratia, heart curves of the Squared Heart Curve (SqHC) type of Ampelocissus and Ampelopsis and Elongated Superellipse-Heart Curves (ESHCs), frequent in Tetrastigma species and observed also in Cissus species and Rhoicissus rhomboidea). The utilities of the application of geometric models for seed description and shape quantification in this family are discussed.

Seed Morphology in Silene Based on Geometric Models.

Seed description in morphology is often based on adjectives such as "spherical", "globular", or "reniform", but this does not provide a quantitative method. A new morphological approach based on the comparison of seed images with geometric models provides a seed description in Silene species on a quantitative basis. The novelty of the proposed method is based in the comparison of the seed images with geometric models according to a cardioid shape. The J index is a measurement that indicates the seed percentage of similarity with a cardioid or cardioid-derived figures used as models. The seeds of Silene species have high values of similarity with the cardioid and cardioid-derived models (J index superior to 90). The comparison with different figures allows species description and differentiation. The method is applied here to seeds of 21 species and models are proposed for some of them including S. diclinis, an endangered species. The method is discussed in the context of previous comparison with the measures used in traditional morphometric analysis. The similarity of seed images with geometric figures opens a new perspective for the automatized taxonomical evaluation of samples linking seed morphology to functional traits in endangered Silene species.