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Seed Shape Quantification in the Order Cucurbitales

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Seed Shape Quantification in the Order Cucurbitales ISSN 2226-3063 e-ISSN 2227-9555 Modern Phytomorphology 12: 1–13, 2018 https://doi.org/10.5281/zenodo.1174871 RESEARCH ARTICLE Seed shape quantification in the order Cucurbitales Emilio Cervantes 1, 2*, José Javier Martín Gómez 1 1 Instituto de Recursos Naturales y Agrobiología de Salamanca-Consejo Superior de Investigaciones Científicas (IRNASA–CSIC), Cordel de Merinas 40, 37008 Salamanca, Spain; * [email protected] 2 Grupo de Investigación Reconocido Bases Moleculares del Desarrollo, Universidad de Salamanca (GIR BMD-USAL), Edificio Departamental, Campus Miguel de Unamuno, 37007 Salamanca, Spain Received: 03.10.2017 | Accepted: 23.01.2018 | Published: 17.02.2018 Abstract Seed shape quantification in diverse species of the families belonging to the order Cucurbitales is done based on the comparison of seed images with geometric figures. Quantification of seed shape is a useful tool in plant description for phenotypic characterization and taxonomic analysis. J index gives the percent of similarity of the image of a seed with a geometric figure and it is useful in taxonomy for the study of relationships between plant groups. Geometric figures used as models in the Cucurbitales are the ovoid, two ellipses with different x/y ratios and the outline of the Fibonacci spiral. The images of seeds have been compared with these figures and values of J index obtained. The results obtained for 29 species in the family Cucurbitaceae support a relationship between seed shape and species ecology. Simple seed shape, with images resembling simple geometric figures like the ovoid, ellipse or the Fibonacci spiral, may be a feature in the basal clades of taxonomic groups. Keywords: Cucurbitales, seed, morphology, geometry Introduction shrubs and trees in 111 genera with a total of 2600 species belonging to eight families: The Angiosperm Phylogeny Group (APG Anisophylleaceae Ridl., Apodanthaceae 2016) includes the order Cucurbitales Tiegh. ex Takht, Begoniaceae C. Agardh, Dumort. in the Fabids, inside the Eurosid Coriariaceae Mirb., Corynocarpaceae Engl., group. The order Cucurbitales Juss. ex Cucurbitaceae Juss. and Datiscaceae L. and Bercht. & J. Presl comprises herbs, climbers, Tetramelaceae Airy Shaw (Airy Shaw 1964). © The Author(s) 2018. Published by Andriy Novikov, State Natural History Museum NAS of Ukraine on behalf of Modern Phytomorphology. This is an open access article under the Creative Commons BY-NC-ND license (http://creativecommons.org/ licenses/by-nc-nd/4.0/) freely available on https://phytomorphology.org/ . 2 Cervantes & Martín Gómez The number of genera is unevenly distributed Kubitzki 2011). Based on molecular studies, among families, and the most of the genera the Tetramelaceae was before classified belong in the Cucurbitaceae (98 genera). within the Datiscaceae (Swensen & Kubitzki Two genera belonging to the Tetramelaceae 2011), but recent studies place them were previously included in the Datiscaceae, separately. Two genera Octomeles Miq. and but recent studies have separated these Tetrameles R. Br. have one buttressed tree two families (Thorne & Reveal 2007; APG species each: Octomeles sumatrana Miq. and 2009; Reveal & Chase 2011; Christenhusz Tetrameles nudiflora R. Br. & Byng 2016). The Tetramelaceae shares The family Anisophylleaceae comprise with Datiscaceae and Begoniaceae several four genera: Anisophyllea R. Br. ex Sabine; features such as the presence of numerous Combretocarpus Hook. f.; Poga Pierre, and small ovules and seeds with a large-celled Polygonanthus Ducke, with 71 species of surface, two-cell-layered integuments, and shrubs and medium to fairly large trees. The a collar around the funicle by an extension of family Coriariaceae includes 14 species of the outer integument (Matthews & Endress shrubs, subshrubs, or rarely perennial herbs 2004). Three families (Anisophylleaceae, in a single genus Coriaria L. (Kubitzki 2011a). Coriariaceae, Corynocarpaceae) were added The family Corynocarpaceae includes six lately on the basis of molecular studies species of evergreen trees in another unique (Kubitzki 2011b). Considering the number genus Corynocarpus J.R. Forst. & G. Forst of species, the order is formed by two large (Kubitzki 2011c). families – Begoniaceae and Cucurbitaceae; Seed shape is an important characteristic one family of intermediate size – in plants. It is related with their ecology and Anisophylleaceae, and five small families – reproduction (especially seed dispersal), Apodanthaceae, Datiscaceae, Coriariaceae, but the use of shape as a taxonomic criteria Corynocarpaceae and Tetramelaceae. requires special methods of quantification. The family Apodanthaceae with 10 Quantification of seed shape may be species of endoparasitic herbs belonging based on automated programs that give to the genera Apodanthes Poit. and Pilostyles magnitudes such as the diameter or axial Guill. was recently included in the order distances of a figure, perimeter, circularity Cucurbitales in APG IV system (APG 2016). index or roundness (Sonka et al. 2008). But The family Begoniaceae comprises two giving the similarity of seed images to a genera of herbs: Begonia L. with at least range of geometric figures in our analysis is 1500 species and Hillebrandia Oliv. with the based on the comparison of surface shared only species H. sandwicensis Oliv., endemic in bi-dimensional images. The comparison to Hawaiian islands (de Wilde 2011). The is by means of the J index, a magnitude that family Cucurbitaceae contains 97 genera gives the percent of similarity between the with 940–980 species (Schaefer & Renner seed image and a chosen geometric figure 2011). Most of them are annual vines, but (Cervantes et al. 2012; also see Cervantes some are woody lianas, thorny shrubs, et al. 2016 for a recent review). The method or exceptionally, trees (e.g., Dendrosicyos was first applied to seeds of the model plant Balf. f.). The family Datiscaceae contains Arabidopsis thaliana (L.) Heynh., comparing two species of robust perennial herbs of their images to a modified cardioid the genus Datisca L.: D. cannabina L. and (elongated in parallel to the symmetry D. glomerata (C. Presl) Baill. (Swensen & axis by a factor of Phi, the Golden Ratio Modern Phytomorphology 12, 2018 Seed shape quantification in the order Cucurbitales 3 = 1.618, approximately). This was used in tripartita (Ucria) Grande in the complex of the identification of differences in seed family Anacardiaceae R. Br. (Saadaoui et al. shape between genotypes (Cervantes et al. 2017). The analysis of seed shape at the level 2010) as well as during seed germination, of order may reveal the validity of these showing that J index reaches a maximum assumptions. value early upon imbibition (Martín et al. This work is a first approach to seed 2014). A cardioid was also the geometric shape quantification in species of the figure used in the quantification of the order Cucurbitales. The hypothesis is that model legume Lotus japonicus L. (Cervantes seeds having single, well defined shapes et al. 2012) as well as for Capparis spinosa L. resembling well defined geometrical figures (Saadaoui et al. 2013). A modified cardioid will appear more frequently in herbs than (elongated in perpendicular to the symmetry in trees. axis by a factor of Phi) was the geometric model for the quantification of seeds in other model legume Medicago truncatula Material and methods Gaertn. In the family Euphorbiaceae Juss., seed shape has been quantified using as Seed image observation the geometric model an ellipse in Ricinus Seed images belonging to six of the eight communis L. (Martín et al. 2016) and Jatropha families (Anisophylleaceae, Begoniaceae, curcas L. (Saadaoui et al. 2015), and variation Coriariaceae, Corynocarpaceae, in values of J index was evaluated in diverse Cucurbitaceae and Datiscaceae) in the cultivars. Cucurbitales were analysed. Seeds from A clear relation exists between seed shape Apodanthaceae and Tetramelaceae and ecological adaptations. In ecology the were not analysed. Seed images were organisms may be classified according to taken from sources indicated in Tab. 1. their structural complexities and life cycle Individual seeds of Ecballium elaterium (L.) in two types: r and K (Begon et al. 2005). The A. Rich. were placed over a flat surface former r type contains species with small with their micropile oriented to the right. individuals having rapid life cycles and not Photographs of orthogonal views of the much structural complexities. The K type, seeds were taken with a digital camera on the contrary, contains species with large Leica C Type 112. individuals, slow cycles and specialised organs (structural complexities). Plants Image analysis that follow the r type strategy tend to have Composed images containing the geometric simple seeds resembling geometric figures. model and each seed were elaborated with For example we found seeds whose images Corel Photo-Paint X7. Quantification of adjust well to the cardioid in model small areas was done with Image J. Individual plants with rapid life cycles and without seed images were used for determination specialised structures (Cervantes et al. of J index. To obtain the J index areas in 2010, 2012). In addition, in the context of two regions were compared: the regions complex plant families we also found seeds shared by the model and the seed image adjusting well to cardioids in the species (i.e. common regions C) and the regions D that have shorter life cycles and simplest not shared between both areas (Fig. 1). The structures, such as in
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