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1 Taxonomy, Botany and Physiology 1 Taxonomy, Botany and Physiology Rafel Socias i Company*, José M. Ansón and María T. Espiau Centro de Investigación y Tecnología Agroalimentaria de Aragón, Zaragoza, Spain 1.1 Almond Taxonomy almond (Amygdalus), peach (Persica), cherry (Cerasus), apricot (Armeniaca) and plum (Prunus Almond belongs to the genus Prunus, which in- sensu stricto). This division seems excessive, par- cludes all stone fruits, and belongs to the Rosace- ticularly if the possibilities of hybridization ae family. Almond is a diploid species with n = 8 among different stone fruit species are con- (Darlington, 1930), the basic ploidy number of sidered. At present, most of these groups are con- Prunus. In contrast to the other species of this sidered subgenera of the genus Prunus, a single genus, whose commercial interest lies in their genus including all the stone fruit tree species, as juicy flesh or mesocarp, almond is the only Prunus the genus Prunus sensu lato. species grown for its seeds, whereas the corky According to Linnaeus’ criteria, under mesocarp is only utilized in animal feeding which the taxa well known to him were exces- (Aguilar et al., 1984) or as a manure (Alonso sively subdivided, almond was designated as et al., 2012). Thus, almond is often classified Amygdalus communis L. (1753, Sp. Pl. 1: 473). more as a nut than as a stone fruit, despite its This same denomination was also adopted by very close genetic similarities with the other A.A. von Bunge as A. communis Bunge (1833, stone fruits, and mainly with peach. Enum. Pl. Chin. Bor. 21), although this name is The botanical name of almond has been a invalid because it is recurrent. The first change subject of diverse controversies and has been was made by P. Miller, who adopted the denomin- changing continuously since Linnaeus began bo- ation A. dulcis Mill. (1768, Gard. Dict., ed. 8. n. 2), tanical systematics. Although taxonomy aims at without taking into account the presence of bitter unambiguous identification of species, different kernels in almond and of sweet kernels in other subjective criteria have led to different classifica- Prunus species, thus making the dulcis designa- tions, despite the application of objective param- tion equivocal. eters for species description. Consequently, fre- The first inclusion of almond in the genus quent cases of synonyms for almond and other Prunus was made by A.J.G.K. Batsch with the de- related species have been – and continue to be – nomination P. amygdalus Batsch (1801, Beytr. reported. Linnaeus considered Amygdalus as an Entw. Gewächsreich 1: 30). This name was independent genus, classifying the stone fruits in widely accepted for a long time and was also different genera, corresponding to the main crops: adopted by J.S. Stokes as P. amygdalus Stokes * Corresponding author e-mail: [email protected] © CAB International 2017. Almonds: Botany, Production and Uses. R. Socias i Company and T.M. Gradziel (eds) 1 2 Rafel Socias i Company et al. (1812, Bot. Mat. Med. 3: 101), although this is hybridizations involving several of these wild also an invalid name because of recurrence. species due to the ease of crossing (Kovalyov and H.G.L. Reichenbach recovered the wrong de- Kostina, 1935), following the theory of species nomination of dulcis, but in the genus Prunus evolution put forward by Vavilov (1930). Hy- under the name P. dulcis (Mill.) Rchb. (1832, Fl. bridizations are frequent among all species of Germ. Excurs. 644), a denomination reactivated this subgenus, giving rise to hybrid forms receiv- more than a century later by D.A. Webb as P. dulcis ing a botanical name and creating some confu- (Mill.) D.A. Webb (1967, Feddes Repert. 74: 24). sion about their real classification (Rehder, Finally, another illegal denomination was pro- 1940; Browicz and Zohary, 1996). Among posed by G. Arcangeli as P. communis (L.) Arcang. these species, P. fenzliana Fritsch, P. bucharica (1882, Comp. Fl. Ital. 209). (Korsh.) Fedtsch, P. kuramica (Korsh.) Kitam. The same diversity of names, mainly due to and P. triloba Lindl. may have been involved in consideration of whether or not Amygdalus is a natural hybridizations, giving rise to the cur- genus or a subgenus, is found for the wild species rent cultivated almond (Grasselly and Crossa- related to almond (Grasselly, 1976; Browicz and Raynaud, 1980; Kester et al., 1990). Recent Zohary, 1996). The most rational classification research with molecular markers has reinforced is that of Rehder (1940), which considered this previous hypothesis (Zeinalabedini et al., Amygdalus (L.) Focke as a subgenus under the 2010). genus Prunus L., although this classification Wild almond species are mostly distributed was not fully adopted. Browicz and Zohary in western and central Asia and eastern Europe (1996) proposed a classification of the species (Fig. 1.1), although some isolated populations related to almond considering Amygdalus as a are found in other locations such as southern genus. Subsequently, Socias i Company (1998) Mongolia and central China (Denisov, 1988) proposed a classification of all almond species and central Spain (Felipe and Socias i Company, under the subgenus Amygdalus (Box 1.1). Unity 1977). In these peripheral regions new hybrid- of the genus Prunus has been confirmed at the izations may have occurred, as could be the molecular level by the Genome Database for case of the wild Mediterranean species P. webbii Rosaceae (2016) and the synteny of traits (Spach) Vierh. (Godini, 1979; Socias i Com- among all species (Arús et al., 2006). pany, 2004) when almond cultivation spread The most recent phylogenetic studies con- into the Mediterranean region, resulting in sider that all almond and peach species must be unique self-compatible populations found considered as forming the subgenus Amygdalus along the northern shore of the Mediterranean of the genus Prunus (Yazbek and Oh, 2013), des- Sea from Greece and the Balkans to Spain and pite the preference of some botanists to main- Portugal. tain Amygdalus as a genus, mostly based on mor- Interest in the wild almond species in scion phological traits and the inconsistency of and rootstock breeding emerged later when they considering mesocarp dehiscence as a differential were considered as possible sources for some trait (Zohary, 1998). interesting traits, such as self-compatibility, early ripening, vigour control, drought resistance, dis- ease resistance and kernel composition (Kostina and Ryabov, 1959; Grasselly, 1977; Vlasic, 1.2 Wild Almond Species 1977; Socias i Company, 1978; Denisov, 1988; Gradziel et al., 2001; Gradziel, 2003). As a con- The interest in wild almond species emerged in sequence, interest developed in maintaining the 19th century and was pursued mainly by some representatives of these species in research botanists (Grasselly, 1976). However, observa- centres, not only as botanical samples, but also tion of the diversity of fruit species in central as a source of genetic variability and as possible Asia in the early 20th century, where not only donors of some traits (Grasselly, 1976). Seeds wild almond species but also wild almond trees of some populations of these wild species had could be found (Popov et al., 1929), led to a new been introduced into the arboretum in Davis, interest in these species (see Chapter 2). The ori- California (Kester et al., 1990) and in the col- gin of the cultivated almond was attributed to lections of INRA, France (Grasselly, 1975); Taxonomy, Botany and Physiology 3 Box 1.1. Classification of almond species (Socias i Company, 1998) Genus: Prunus L. (1735) Subgenus: Amygdalus (L.) Focke Series: Icosandrae Spach (1843) (Syn.: Subgen. Amygdalus Browicz et Zohary) (i) Section: Euamygdalus Spach (1843) (Syn.: Sect. Amygdalus Browicz et Zohary) Group Amygdalus 1. P. amygdalus Batsch (1801) (Syn.: A. communis L., A. dulcis Mill., P. communis (L.) Arcang., P. dulcis (Mill.) D.A. Webb). It includes P. korshinskyi Hand.-Mazz.) 2. P. trichamygdalus Hand.-Mazz. (1913) (Syn.: A. trichamygdalus (Hand.-Mazz.) Woronow) 3. P. fenzliana Fritsch (1892) (Syn.: A. fenzliana (Fritsch) Lipsky) 4. P. webbii (Spach) Vierh. (1915) (Syn.: A. webbii Spach, A. salicifolia Boiss. et Bal.) 5. P. haussknechtii C. Schneider (1905) (Syn.: A. haussknechtii (C. Schneider) Bornm.) 6. P. zabulica (Seraf.) SIC, comb. nov. (1998) (Syn.: A. zabulica Seraf.). It includes A. browiczii Freitag) 7. P. kuramica (Korsh.) Kitam. (1960) (Syn.: A. kuramica Korsh.) 8. P. bucharica (Korsh.) Hand.-Mazz. (1913) (Syn.: A. bucharica Korsh.) 9. P. tangutica (Batal.) Koehne (1912) (Syn.: A. tangutica (Batal.) Korsh.) Group Orientalis 10. P. argentea (Lam.) Rehd. (1922) (Syn.: A. orientalis Duhamel, A. argentea Lam.) 11. P. discolor (Spach) C. Schneider (1905) (Syn.: A. graeca Lindl., A. discolor (Spach) Roemer) 12. P. elaeagnifolia (Spach) E. Murray (1969) (Syn.: A. elaeagnifolia Spach, A. kermanensis Bornm.) 13. P. kotschyi (Boiss. et Hohen.) Náb. (1923) (Syn.: A. kotschyi Boiss. et Hohen.) 14. P. carduchorum (Bornm.) Meikle (1965) (Syn.: A. carduchorum Bornm.) 15. P. mongolica Maxim. (1879) (Syn.: A. mongolica (Maxim.) Ricker) (ii) Section: Chamaeamygdalus Spach (1843) 16. P. nana (L.) Stokes (1812) (Syn.: A. nana L., P. tenella Batsch). It includes P. georgica (Desf.) and P. ledebouriana (Schlecht.) 1 7. P. petunnikovii (Litv.) Rehd. (1926) (Syn.: A. petunnikovii Litv.) (iii) Section: Spartioides Spach (1843) 18. P. arabica (Olivier) Meikle (1967) (Syn.: A. arabica Olivier, A. spartioides Spach, P. spartioides (Spach) C. Schneider) Continued 4 Rafel Socias i Company et al. Box 1.1. Continued. 19. P. scoparia (Spach) C. Schneider (1905) (Syn.: A. scoparia Spach) Series: Dodecandra Spach (1843) (Syn.: Lycioides Spach) 20. P. lycioides (Spach) C. Schneider (1906) (Syn.: A. lycioides Spach) 2 1. P. spinosissima (Bge.) Franch. (1883) (Syn.: A. spinosissima Bge.) 22. P. eburnea (Spach) Aitch. et Hemsley (1886) (Syn.: A. eburnea Spach, A. spathulata Boiss.) 23. P. brahuica (Boiss.) Aitch. et Hemsley (1886) (Syn.: A. brahuica Boiss.) 24. P. erioclada (Born.) SIC, comb.
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