Concepts and nomenclature of the Farro wheats, with special reference to Emmer, Triticum turgidum subsp. dicoccum (Poaceae)

Summary

This paper discusses the different taxonomic concepts of the wheat group as exemplified by three species, commonly known together as ‘Farro’: diploid Einkorn, tetraploid Emmer and hexaploid Spelt. A narrow, morphology-based concept is contrasted with a much wider, genome-based one, leading to profound differences in the recognition of taxa at species level and below. The latter concept accepts far fewer taxa; it is advocated here and its effect illustrated for the Farro wheats. Considerations regarding the nomenclature of the accepted taxa are presented, applying both the International Code for the Nomenclature of Cultivated Plants (ICNCP) and the International Code of Nomenclature for algae, fungi and plants (ICN or the ‘Melbourne Code’). Within the genus Triticum L. we propose Farro wheats to be classified at subspecies rank, and to be defined as the total of their cultivars united under a botanical name. Einkorn is T. monococcum L. (the cultivars in subsp. monococcum) and Spelt T. aestivum L. subsp. spelta (L.) Thell.; if one chooses the species level the names are T. monococcum L. and T. spelta L., respectively. We show that, based on consideration of the original literature, the ICN and relevant type specimens, the correct name and authorship of Emmer at subspecies level should be T. turgidum L. subsp. dicoccum (Schrank ex Schübl.) Thell. At species level we recommend using T. farrum Bayle-Bar., published in 1809, over the more widely known T. dicoccum Schrank ex Schübl. from 1818, and provide arguments for rejecting T. dicoccon Schrank 1789, T. album Gaertn. 1790, T. spelta sensu Host 1809 (non Linnaeus) and (possibly) T. atratum Host 1809.

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References

  1. Alefeld, F. G. C. (1866). Landwirthschaftliche Flora. Wiegandt & Hempel, Berlin.

    Google Scholar 

  2. Ascherson, P. F. A. & Graebner, K. O. R. P. P. (1898 – 1899). Flora des Nordostdeutschen Flachlandes. Gebrüder Borntraeger, Berlin. (Triticum on pp. 122 – 125 from 16 July 1898).

  3. ____ & ____ (1901). Synopsis der Mitteleuropäischen Flora, Vol. 2(1). W. Engelmann, Leipzig. (pp. 545 – 741 from 10 Dec. 1901).

  4. Bailey, L. H. (1923). Gentes Herbarium 1(3). ‘Published (mailed at Ithaca Post Office to regular list)’.

  5. Bauhin, C. (1623). Pinax Theatri Botanici (1st ed.). Sumptibus & typis Ludovici Regis, Basel.

  6. Bayle-Barelle, G. (1809). Monografia Agronomica dei Cerealidel Formento. Trattato Diviso in Tre Parti con sei Tavole. Giovanni Silvestri (Stampatore-Librajo), Milano. (Parte prima from 1808, but the title page of the book cites 1809).

  7. Bluff, M. J., Nees von Esenbeck, C. G. D. & Schauer, J. C. (1836). Compendium Florae Germanicae. Sectio I. Tomus I. Pars I. J. L. Schrag, Nuremberg.

    Google Scholar 

  8. Bor, N. L. (1968). 29. Triticum L. In: C. C. Townsend et al., Flora of Iraq, Vol. 9, pp. 94 – 208. Ministry of Agriculture, Baghdad.

  9. ____ (1970). 6. Triticum L. In: K. H. Rechinger, Flora Iranica, No. 70/30, pp. 203 – 211. Akademische Druck- u. Verlagsanstalt, Graz.

    Google Scholar 

  10. Bowden, W. (1959). The taxonomy and nomenclature of the wheats, barleys, and ryes and their wild relatives. Canad. J. Bot. 37: 657 – 684.

    Google Scholar 

  11. Brickell, C. D., Alexander, C., David, J. C., Hetterscheid, W. L. A., Leslie, A. C., Malecot, V. & Xiaobai, J. (2009). International Code of Nomenclature for Cultivated Plants. 8th ed. (ICNCP or Cultivated Plant Code). International Society for Horticultural Science.

  12. Briggle, L. W. & Reitz, L. P. (1963). Classification of Triticum species and of wheat varieties grown in the United States. Technical Bulletin no. 1278, United States Department of Agriculture, Washington DC.

  13. Cai, L., Zhang, S. & Li, J. (1991). Studies on the classification of the genus Triticum L., Acta Bot. Boreal.-Occid. Sin. 11: 212 – 224.

    Google Scholar 

  14. De Vilmorin, H. (1880). Les Meillieurs Blés, Description et Culture des Principales Variétés de Froments d’Hiver et de Printemps par Vilmorin-Andrieux et C ie. Vilmorin-Andrieux et Cie, Paris.

    Google Scholar 

  15. De Vilmorin, Ph. L. (1905). Hortus Vilmorinianus. Catalogue des Plantes Ligneuses et Herbacées Existant en 1905 dans les Collections de m. Ph.L. de Vilmorin et dans les Cultures de MM. Vilmorin-Andrieux et Cie à Verrières-le-Buisson.

  16. De Wet, J. (1981). Species concepts and systematics of domesticated cereals. Kulturpflanze 29: 177 – 198.

    Article  Google Scholar 

  17. Döll, J. C. (1857). Flora des Grossherzogthums Baden, erster Band. G. Braun, Karlsruhe. (Vol. 1(2): 91 – 298 from Jan. – Dec. 1855).

  18. Dorofeev, V. F. & Korovina, O. N. (eds) (1979). Pshenitsa [Wheat]. Kulturnaya Flora SSSR, Vol. 1. Kolos, Leningrad.

  19. Dumont de Courset, G. L. M. [Baron] (1811). Le Botaniste Cultuvateur (2nd ed.), Vol. 2. Déterville, Goujon, Paris. (Author writes name as ‘du Mont de Courset’ on the title page; the different version of this entry follows Stafleu & Cowan, Taxonomic Literature 2(1)).

  20. Filatenko, A. A. & Kurkiev, U. K. (1975). Pshenitsa sinskoye [Sinskaja’s wheat]. Trudy Prikl. Bot., Genet. i Selekt. 54(1): 239 – 241.

    Google Scholar 

  21. Fiori, A. (1896). Triticum. In: A. Fiori & G. Paoletti (1896), Flora Analitica d'Italia, Vol. 1, pp. 107 – 108. Tipografia del Seminario, Padova.

  22. Flaksberger, C. A. (1915). [Determination of wheats.]. Trudy Byuro Prikl. Bot. 8: 9 – 198. (In Russian with English summary pp. 183 – 198).

  23. ____ (1935). Klebnyie zlaki — pshenitsa [Cereal grasses — wheat]. In: E. V. Wulff (ed.), Flora of Cultivated Plants of the USSR, Vol. 1. State Agricultural Publishing Company, Moskou-Leningrad. (In Russian).

  24. Gaertner, J. (1790). De Fructibus et Seminibus Plantarum, Volumen Alterum 2(1). G. H. Schramm, Tübingen.

  25. Gontcharov, N. P. (2011). Genus Triticum L. taxonomy: the present and the future. Pl. Syst. Evol. 295: 1 11.

    Article  Google Scholar 

  26. Hackel, E. (1887). Gramineae. In: H. G. A. Engler & K. A. E. Prantl (eds), Die Natürlichen Pflanzenfamilien 2(2), 80 – 86. W. Engelmann, Leipzig.

  27. Hammer, K., Filatenko, A. A. & Pistrick, K. (2011). Taxonomic remarks on Triticum L. and × Triticosecale Wittm. Genet. Resources Crop Evol. 58: 3 – 10.

    Article  Google Scholar 

  28. Harlan, J. R. & de Wet, J. M. J. (1971). Toward a rational classification of cultivated plants. Taxon 20: 509 – 517.

    Article  Google Scholar 

  29. Harz, C. O. (1885). Landwirthschaftliche Samenkunde, Vol. 2. Paul Parey, Berlin.

    Google Scholar 

  30. Host, N. T. (1805). Icones et descriptiones graminum austriacorum, Vol. 3. M. A. Schmidt, Vindobonae (Vienna).

  31. ____ (1809). Icones et Descriptiones Graminum Austriacorum, Vol. 4. M. A. Schmidt, Vindobonae (Vienna).

  32. Humphries, C. J. (1978). (297) Triticum L. In: V. H. Heywood (ed.), Flora Europaea Notulae Systematicae ad Floram Europaeam spectantes. No. 20. Bot. J. Linn. Soc. 76: 368 – 369.

  33. Husnot, P. T. (1899). Graminées 4: 80. T. Husnot, Cahan [Caen]. (Part 4, pp. 73 – 92 from Nov. 1899).

  34. Kerguélen, M. (1975). Les Gramineae (Poaceae) de la flore française. Essai de mise au point taxonomique et nomenclaturale. Lejeunia, Nouv. Sér. 75: 1 – 343.

    Google Scholar 

  35. Körnicke, F. A. (1885). Vol. 1: Die Arten und Varietäten des Getreides. In: F. A. Körnicke & H. Werner, Handbuch des Getreidebaues. Paul Parey, Berlin.

    Google Scholar 

  36. Lagasca y Segura, M. (1816). Elenchus Plantarum. Typographia Regia, Madrid.

  37. Linnaeus, C. (1753). Species Plantarum (1st ed.), Vol. 1. L. Salvius, Stockholm.

  38. ____ (1762). Species Plantarum (2nd ed.), Vol. 1. L. Salvius, Stockholm.

  39. ____ (1774). Systema Vegetabilium. Editio decima tertia (13th ed.). J. C. Dietrich, Göttingen & Gotha.

  40. Löve, Á. (1984). Conspectus of the Triticeae. Feddes Repert. 95: 425 – 521.

    Google Scholar 

  41. Lunell, J. (1915). Enumerantur plantae dakotae septentrionalis vasculares. — II (The vascular plants of North Dakota. — II). Amer. Midl. Naturalist 4: 211 – 228.

  42. Mac Key, J. (1954). The taxonomy of hexaploid wheat. Svensk Bot. Tidskr. 48: 579 – 590.

    Google Scholar 

  43. ____ (1966). Species relationship in Triticum. In: Proceedings of the Second International Wheat Genetics Symposium, Lund, Sweden. Hereditas, Suppl. Vol. 2: 237 – 276.

    Google Scholar 

  44. ____ (1981). Comments on the basic principles of crop taxonomy. Kulturpflanze 29: 199 – 207.

    Article  Google Scholar 

  45. ____ (2005). Wheat: its concept, evolution, and taxonomy. In: C. Rojo, M. M. Nachit, N. Di Fonzo, W. H. Pfeiffer & G. A. Slater (eds), Durum Wheat Breeding: Current Approaches and Future Strategies, pp. 3 – 61. Food Products Press, Binghamton, NY.

    Google Scholar 

  46. Malpighi, M. (1687). Opera Omnia. Petrum vander Aa, Lugduni Batavorum [Leiden].

  47. Mansfeld, R. (1958). Zur Nomenklatur einiger Nutz- und Kulturpflanzen II. Kulturpflanze 6: 237 – 242.

    Article  Google Scholar 

  48. McNeill, J., Barrie, F. R., Burdet, H. M., Demoulin, V., Hawksworth, D. L., Marhold, K., Nicholson, D. H., Prado, J., Silva, P. C., Skog, L. E., Wiersema, J. H. & Turland, N. J. (2006). International Code of Botanical Nomenclature (Vienna Code), Adopted by the Seventeenth International Botanical Congress, Vienna, Austria, July 2005. A. R. G. Gantner Verlag KG, Ruggell.

    Google Scholar 

  49. ____, ____, Buck, W. R., Demoulin, V., Greuter, W., Hawksworth, D. L., Herendeen, P. S., Knapp, S., Marhold, K., Prado, J., Prud’homme van Reine, W. F., Smith, G. F., Wiersema, J. H. & Turland, N. J. (2012). International Code of Nomenclature for Algae, Fungi, and Plants (Melbourne Code), Adopted by the Eighteenth International Botanical Congress, Melbourne, Australia, July 2011. Koeltz Scientific Books, Königstein.

    Google Scholar 

  50. Metzger, J. (1824). Europæische Cerealien. C. F. Winter, Heidelberg.

    Google Scholar 

  51. Morison, R. (1699). Plantarum Historiae Universalis, Vol. 3. Theathro Sheldoniano, Oxford.

    Google Scholar 

  52. Morris, R. & Sears, E. R. (1967). The cytogenetics of wheat and its relatives. In: K. S. Quisenberry & L. P. Reitz (eds), Wheat and Wheat Improvement, pp. 19 – 87. American Society of Agronomy, Wisconsin.

    Google Scholar 

  53. Morrison, L. A. (1998). Lectotypification of Triticum turgidum and T. spelta (Poaceae). Taxon 47: 705 – 710.

    Article  Google Scholar 

  54. ____ (2007). Triticum. In: M. E. Barkworth, K. M. Capels, S. Long, L. K. Anderton & M. B. Piep (eds), Flora of North America, North of Mexico, Vol. 24, pp. 268 – 277. Oxford University Press, New York, Oxford.

    Google Scholar 

  55. Percival, J. (1921). The Wheat Plant, a Monograph. Duckworth and Co., London.

    Google Scholar 

  56. Roemer, J. J. & Schultes, J. A. (1817). Systema Vegetabilium, Vol. 2. J. G. Cottae, Stuttgart.

    Google Scholar 

  57. Schlechtendahl, D. F. L. von (1838). Reliquiae Schraderianae. Linnaea 12: 360 – 476.

    Google Scholar 

  58. Schrank, F. von Paula von (1789). Bayerische Flora. Erster Band. J. B. Strobl, München.

  59. Schübler, G. (Nov. 1818). Dissertatio Inauguralis Botanica Sistens Characteristicen et Descriptiones Cerealium… Ludov. Frider. Fues., Tübingen. (The publication was the doctoral thesis of J. L. Rode, but the committee was presided over by Schübler to whom new names and combinations have to be ascribed).

  60. ____ (1820). Beschreibung und systematische Bestimmung der in Würtemberg unter dem Namen Ehmer (Emmer) gebauten Getreide – Art in Vergleichung mit Einkorn und einigen anderen zunächst damit verwandten Arten. Flora 3: 445 – 462, Plate 1, figs 1 – 4.

  61. ____ & von Martens, G. M. (1834). Flora von Würtemberg. C. F. Osiander, Tübingen.

    Google Scholar 

  62. Sears, E. R. (1959). The systematics, cytology and genetics of wheat. In: H. Kappert & W. Rudorf (eds), Manual of Plant Breeding (2nd revised ed.), Vol. 2: 164 – 187. Paul Parey, Berlin and Hamburg.

    Google Scholar 

  63. Seringe, N. C. (Sept. 1818). Mélanges Botaniques 1(2): Monographie des Céréales de la Suisse, pp. 65 – 244. chez l’Auteur, Berne.

  64. ____ (1841). Descriptions et figures des céréales européennes. Ann. Sci. Phys. Nat. Lyon 4: 321 – 384.

    Google Scholar 

  65. ____ (1842). Céréales européennes, second article. Ann. Sci. Phys. Nat. Lyon 5: 103 – 196.

    Google Scholar 

  66. Soreng, R. J. (2003). Triticum L. In: R. J. Soreng (ed.), Catalogue of New World Grasses (Poaceae): IV. Subfamily Pooideae, Contr. U.S. Natl. Herb. 48: 676 – 684.

  67. Stafleu, F. A. & Cowan, R. S. (1976). Taxonomic Literature. Vol. I: A – G. Bohn, Scheltema & Holkema, Utrecht.

    Google Scholar 

  68. ____ & ____ (1979). Taxonomic Literature. Vol. II: H – Le. Bohn, Scheltema & Holkema, Utrecht/Dr W. Junk Publishers, The Hague.

  69. ____ & Mennega, E. A. (1992). Taxonomic Literature. Supplement I: A – Ba. Koeltz Scientific Books, Königstein.

  70. Stallknecht, G. F., Gilbertson, K. M. & Ranney, J. E. (1996). Alternative wheat cereals as food grains: Einkorn, emmer, spelt, kamut and triticale. In: J. Janick (ed.), Progress in New Crops, pp. 156 – 170. ASHS Press, Alexandria, VA.

    Google Scholar 

  71. Szabó, A. T. & Hammer, K. (1996). Notes on the taxonomy of farro: Triticum monococcum, T. dicoccon and T. spelta. In: S. Padulosi, K. Hammer & J. Heller (eds), Hulled Wheats. Proceedings of the First International Workshop on Hulled Wheats, 21 – 22 July 1995, Castelvecchio, Tuscany, Italy, pp. 2 – 40. International Plant Genetic Resources Institute.

  72. Tan, K. (1985). Triticum. In: P. H. Davis (ed.), Flora of Turkey, Vol. 9. University Press, Edinburgh.

    Google Scholar 

  73. Thellung, A. (1912). La Flore Adventice de Montpellier. E. Le Maout, Cherbourg. [Pre-print from Mém. Soc. Natl. Sci. Nat. Cherbourg 38: 57 – 728 (1912).]

  74. ____ (1918a). Neuere Wege und Ziele der botanischen Systematik, erläutert am Beispiele unserer Getreidearten. Naturwiss. Wochenschr. N. F. 17: 465 – 474. (Appeared 18 Aug. 1918).

  75. ____ (1918b). Neuere Wege und Ziele der botanischen Systematik, erläutert am Beispiele unserer Getreidearten. Mitt. Naturwiss. Ges. Winterthur 12: 109 – 152. (Appeared Nov. 1918).

  76. Tournefort, J. Pitton de (1719). Institutiones Rei Herbariae (Editio Tertia), Tomus III [Vol. 3]. Typographia regia, Paris.

  77. Trehane, P., Brickell, C. D., Baum, B. R., Hetterscheid, W. L. A., Leslie, A. C., McNeill, J., Spongberg, S. A. & Vugtman, F. (1995). International Code of Nomenclature for Cultivated Plants – 1995 (ICNCP or Cultivated Plant Code). Quarterjack Publishing, Wimborne.

    Google Scholar 

  78. Trinius, C. B. (1822). Clavis Agrostographiae Antiquioris. Biedermann, Coburg.

    Google Scholar 

  79. Tzvelev, N. N. (1973). Conspectus specierum tribus Triticeae Dum. familiae Poaceae in flora URSS. Novosti Sist. Vyssh. Rast. 10: 19 – 59.

    Google Scholar 

  80. ____ (1976). Zlaki SSSR 1. Nauka, Leningrad / (1983). Grasses of the Soviet Union, Part 1. (English translation; prepared for the Smithsonian Institution Libraries, and the National Science Foundation, Washington DC by Amerind Publishing Co. Pvt. Ltd., New Delhi).

  81. Valdés, B. & Scholz, H. (2006). The Euro + med treatment of the Gramineae — a generic synopsis and some new names. Willdenowia 36: 657 – 669.

    Google Scholar 

  82. Van Slageren, M. W. (1994). Wild wheats: a monograph of Aegilops L. and Amblyopyrum (Jaub. & Spach) Eig (Poaceae). Wageningen Agric. Univ. Pap. 94(7): 1 – 512.

    Google Scholar 

  83. Vavilov, N. I. (1923). [A contribution to the classification of soft wheats — Triticum vulgare Vill.], Trudy Prikl. Bot. Selekts. 13: 149 – 257. (In Russian with English summary pp. 216 – 257).

  84. Voss, A. (1895). In: A. Siebert & A. Voss, Vilmorin’s Blumengärtnerei (3rd ed.), 1(2): 1217 – 1218. Paul Parey, Berlin. (Entire publication 1894 – 1896; Vol. 1(2) pp. 833 – 1264 from 1895).

  85. Zaharieva, M., Ayana, N. G., Hakimi, A. A., Misra, S. C. & Monneveux, P. (2010). Cultivated emmer wheat (Triticum dicoccon Schrank), an old crop with a promising future: a review. Genet. Resources Crop Evol. 57: 937 962.

    Article  Google Scholar 

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Acknowledgements

At K we thank Melanie Thomas for advice and translation of Schübler’s (1818) Latin text, Paul Little for his production of high-resolution images of the relevant illustrations, the staff at the Library, and Dr Wolfgang Stuppy (Seed Conservation Department) for his ‘Stuppy-fication’ of most of the illustrations, making them a superior version of the originals. Furthermore we thank the curators of G, M, PAV, TUB and W for their help in identifying (possible) type collections, and the staff at the Libraries of the Natural History Museum (NHM) and the British Library for their help in making the relevant literature available. We also acknowledge the granting of permission by the British Library Board and the Botanische Staatssammlung München (M) to publish images under their copyright. Dr Zewdie Bishaw of the International Center for Agricultural Research in the Dry Areas (ICARDA) advised on UPOV and the ITPGRFA. We thank the reviewers for useful suggestions and improvements. This research is financially supported by the Stichting Elise Mathilde Fonds, CIMMYT, Int., and the COFRA foundation.

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van Slageren, M., Payne, T. Concepts and nomenclature of the Farro wheats, with special reference to Emmer, Triticum turgidum subsp. dicoccum (Poaceae). Kew Bull 68, 477–494 (2013). https://doi.org/10.1007/s12225-013-9459-8

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Key Words

  • taxonomy
  • wheat