plants
Article
Geometric Models for Seed Shape Description and
Quantification in the Cactaceae
José Javier Martín-Gómez
1
, Diego Gutiérrez del Pozo
2
, Ángel Tocino
3
and Emilio Cervantes
1,
*
Citation: Martín-Gómez, J.J.; del
Pozo, D.G.; Tocino, Á.; Cervantes, E.
Geometric Models for Seed Shape
Description and Quantification in the
Cactaceae. Plants 2021, 10, 2546.
https://doi.org/10.3390/
plants10112546
Academic Editor: Bertrand Dubreucq
Received: 10 November 2021
Accepted: 22 November 2021
Published: 22 November 2021
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1
Instituto de Recursos Naturales y Agrobiología del Consejo Superior de Investigaciones
Científicas (IRNASA-CSIC), Cordel de Merinas, 40, E-37008 Salamanca, Spain; jjavier.martin@irnasa.csic.es
2
Herbario Amazónico del Ecuador ECUAMZ, Universidad Estatal Amazónica UEA, Carretera Tena a Puyo
Km. 44, Puyo EC-150950, Ecuador; diego.gutierrez.pozo@gmail.com
3
Departamento de Matemáticas, Facultad de Ciencias, Universidad de Salamanca, Plaza de la Merced 1-4,
E-37008 Salamanca, Spain; bacon@usal.es
* Correspondence: emilio.cervantes@irnasa.csic.es
Abstract: 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.
Keywords: Archimedean spiral; cactaceae; geometry; models; morphology; seed shape; spiral
1. Introduction
The Cactaceae Juss., with about 2000 species distributed in ca. 120 genera, are char-
acterized by succulent photosynthetic stems. The family has been included in the order
Caryophyllales Juss. ex Bercht. & J.Presl [1], in the suborder Portulacineae Thorne [2] which
included Cactaceae, Portulacaceae, Didiereaceae, and Basellaceae, as well as other succulent
plant families (Anacampserotaceae, Halophytaceae, Montiaceae, and Talinaceae) [3–5].
Cacti are common in the arid ecosystems of America, in the southern States of USA
and Mexico, north-eastern Brazil, and along the slopes of the Andes from Ecuador to
Argentina and Chile. Many species also live in humid forests as epiphytes. All known
species are originally American with the only exception of Rhipsalis baccifera (J.S. Muell.)
Stearn, found also in local populations in tropical Africa and Sri Lanka.
The Cactaceae have been divided into four subfamilies: Cactoideae Eaton, Maihue-
nioideae P. Fearn, Opuntioideae Burnett, and Pereskioideae Engelm. The Cactoideae and
Opuntioideae contain, respectively, 100 and 9 genera [4,5]. Maihuenia (Phil.) Britton & Rose,
with two cushion-forming, mucilaginous species, is the only genus in the Maihuenioideae.
Pereskia Mill, together with Leuenbergeria Lodé in the Pereskioideae, are woody with large,
glossy leaves and long thorns on the stem, and not all are succulents. The Cactoideae,
with ten tribes, is the most diverse subfamily of globular and columnar cacti including all
the iconic species in American dessert biomes [5]. The Opuntioideae correspond to the
prickly pear cacti and relatives, with seeds enclosed by a bony aril that is considered a
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