Concepts and nomenclature of the Farro wheats, with special reference to Emmer, Triticum turgidum...

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

Michiel van Slageren1 & Thomas Payne2

Summary. This paper discusses the different taxonomic concepts of the wheat group as exemplified by threespecies, 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 inthe recognition of taxa at species level and below. The latter concept accepts far fewer taxa; it is advocated hereand its effect illustrated for the Farro wheats. Considerations regarding the nomenclature of the accepted taxa arepresented, applying both the International Code for the Nomenclature of Cultivated Plants (ICNCP) and theInternational Code of Nomenclature for algae, fungi and plants (ICN or the ‘Melbourne Code’). Within the genusTriticum L. we propose Farro wheats to be classified at subspecies rank, and to be defined as the total of theircultivars united under a botanical name. Einkorn is T. monococcum L. (the cultivars in subsp. monococcum) and SpeltT. aestivum L. subsp. spelta (L.) Thell.; if one chooses the species level the names are T. monococcum L. and T. speltaL., respectively. We show that, based on consideration of the original literature, the ICN and relevant type speci-mens, 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 themore widely known T. dicoccum Schrank ex Schübl. from 1818, and provide arguments for rejecting T. dicocconSchrank 1789, T. album Gaertn. 1790, T. spelta sensu Host 1809 (non Linnaeus) and (possibly) T. atratum Host 1809.

Key Words. taxonomy, wheat.

IntroductionFarro is an Italian ethnobotanical concept (Szabó &Hammer 1996: 3) and applies to three ‘hulled’, or ‘notfree-threshing’ wheat taxa, that is to say wheats inwhich the spike rachis breaks up under pressure andthe spikelets fall with their glumes still attached. Free-threshing or not is an important agronomic characteras it means the difference between an easy, almostspontaneous, separation of the grains from the re-mains of the flower and this being not so. In the lattercase threshing can still create the desired result butachieving it is much more elaborate and thus costly:an example from South Africa cited the cost for hulledspelt wheat at roughly 5.5 times that of bread wheat(D. van Papendorp, pers. comm.). The Farro wheatsare Einkorn (Triticum monococcum L. subsp.monococcum), Emmer (T. turgidum L. subsp. dicoccum(Schrank ex Schübl.) Thell.), and Spelt (T. aestivumL. subsp. spelta (L.) Thell.); in terms of our generalconcept of wheat taxa they are all considered atsubspecies rank. These three wheats are part of theprimary gene pool (or GP-1) of cultivated wheat. Thisdesignation of gene pools is based on ease of

crossability with a particular cultivated taxon (Harlan& de Wet 1971). In Triticum the GP-1 consists of fourgroups, based around Einkorn, turgidum wheat(T. turgidum L.), Timofeev wheat (T. timopheevii(Zhuk.) Zhuk.), and bread wheat (T. aestivum L.)(Harlan&deWet 1971:fig. 2). Szabó&Hammer (loc. cit.)point out that ‘hulled wheats’ are not the same as ‘notfree-threshing’, as many species from the wider wheatgene pool (GP-2 and -3 in the scheme of Harlan & deWet) possess this character too, for example species ofgenera such as Aegilops L., Amblyopyrum (Jaub. & Spach)Eig (separation fromAegilops is sensu van Slageren 1994),and Secale L. However, within GP-1 Triticum the twoare effectively the same: hulled wheats = Farro (butsee below).

Conversely, Szabó & Hammer consider the free-threshing Triticum sinskajae Filat. & Kurkiev part ofFarro to illustrate it does not automatically mean‘hulled’. However, we question whether one can speakof a ‘species’ in this case (see the nomenclaturesummary below). To our knowledge it has never beencommercially released as a cultivar to replace the

Accepted for publication 9 May 2013.1 Royal Botanic Gardens Kew, Seed Conservation Department, Wakehurst Place, Ardingly, West Sussex, RH17 6TN, UK. e-mail: [email protected] The International Wheat and Maize Improvement Center (CIMMYT, Int.), Genetic Resources Program, Apdo. Postal 6-641, 06600 Mexico, DF, Mexico.

e-mail: [email protected].

KEW BULLETIN VOL. 68: 1 Y 18 (2013)DOI 10.1007/S12225-013-9459-8

ISSN: 0075-5974 (print)ISSN: 1874-933X (electronic)

© The Board of Trustees of the Royal Botanic Gardens, Kew, 2013

more cumbersome hulled Einkorn, and rather thandescribing and conferring a name and type we feel itshould have remained ‘research material of plantbreeding interest’. As T. sinskajae is a published speciesand is discussed here, its nomenclature is includedbelow, with typography indicating our opinion on thestatus of this taxon.

While we equate Farro in this publication with thethree well-known, hulled wheat species, other Triticumspecies are classified as being ‘spelt’ type in contrast to‘naked’ or ‘free-threshing’ type. They may thereforeexhibit hulled characteristics as the Farro species do.Hexaploid T. aestivum L. subsp. macha (Dekapr. &Menabde) Mac Key from Georgia is considered spelt-type by Mac Key (1954: 579), while Sears (1959: 164),Briggle & Reitz (1963: 35) and Gontcharov (2011: 6)add T. aestivum L. subsp. vavilovii (Tumanian) Á. Löve,a branched mutant type of bread wheat originatingfrom Turkey. However, these are not Farro wheats ascommonly understood, and are not further consid-ered here.

Grouping hulled (as in: Farro) wheats separatefrom the free-threshing ones is an old concept, datingback to pre-Linnaean times (Morrison 1998: 708). Asearly as 1623, Bauhin (1623: 21 – 22) listed them in theseparate (from Triticum that is) genus Zea. As theirphrase-names include such words as ‘Monococos’,‘dicoccos’ and ‘Triticospeltum’ (the latter cited byTrinius (1822: 377) as ‘Tritico - speltum’ and identifiedas T. spelta L.), it is clear which species are meant.Several overviews and monographs of the wheat groupuse hulled vs free-threshing, for example Bayle-Barelle(1809), Seringe (1818, in the ‘Tableau méthodique’),Metzger (1824), Alefeld (1866), Harz (1885), Körnicke(1885), Flaksberger (1915), Thellung (1918a, b) and,more recently, Sears (1959) and Briggle & Reitz (1963:35). The juxtaposition can also be found in floraswhen all hulled species are present, for example inthose from Schübler & von Martens (1834: 46), Döll(1857 (1855): 124, as ‘II. Zea, Spelzweizen’), orAscherson & Graebner (1898: 124).

Different approaches in the classificationof Farro speciesThe precise naming and taxonomic position matters,as the Farro species are somewhat of a resurgent crop.Accordingly, the breeding value of all three ispresently the subject of renewed interest (Stallknechtet al. 1996; Zaharieva et al. 2010). It is comparativelyeasy to cross them with selections of the two mainwheat species, durum wheat (Triticum turgidumL. subsp. durum (Desf.) Husn.) and bread wheat(T. aestivum subsp. aestivum). Emmer enjoys increasedinterest for a potential role in high-quality food;accessions are known with comparatively high (thoughvariable) protein and mineral content, can be a source

for agronomically useful characters such as tilleringand grain weight, while resistance to rusts and buntare also recorded (Zaharieva et al. 2010). Transferringthese traits is not without problems but should becomparatively easy as, for example, complete genomiccompatibility between the Emmer and durum wheatgenomes is reported (Zaharieva et al. 2010: 953,quoting a study by Yanchenko).

We realise that much wheat research is carried outby plant breeders, genetic resources scientists, pathol-ogists, entomologists and the like, and that thereforethe correct botanical name of the Farro species isperhaps not a priority. However, taxonomic opinionaffects the naming of the genetic resources used bythese scientists, and reality is often different from theideal that correctly identified material is available. It ishere that the effect of two schools of thought, bothstill very much in evidence, is most apparent.

The ‘morphology’ schoolFirstly, there has been the historical development (andacceptance) of a detailed infraspecific classificationthat aims to describe and appreciate the richness ofmorphological variation found in the species, be thiswild or cultivated ones (Hammer et al. 2011). Whilethere is quite some history here, our exposure mainlycentres on a relatively recent paper by Szabó &Hammer (1996). Although accepting many micro-taxa,these authors nevertheless advocated (1996: 4 – 5) theneed for a practical scheme to be widely adopted inwhich ‘…wild taxa are consequently treated on thesubspecies and varietal levels, and cultivated taxa on theconvariety and (pro)variety levels…’ However, theadditional ‘(supra)convar.’ as a second equivalent of‘subsp.’ for cultivated material is then introduced, as arethe smallest categories of ‘f. (forma)’ for wild taxa and‘cv. (cultivar)’ for cultivated taxa. The distinction withinthe ranks for cultivated groups is for ‘larger’ vs ‘smaller’ones: several cvs. into a provar.; then, for reasons ofgeographical distribution (tom. cit. 16), one or severalprovars. are joined into a convar. or sometimes a(supra)convar. The problems with this proposed practi-cal scheme become immediately apparent when theauthors proceed to classify the Farro species. It shouldalso be noted that the ranks of provar., convar.,supraconvar., grex, race, and proles as used byFlaksberger, Hammer, Szabó and others for units of(predominantly) cultivated plants are not cited but canbe intercalated between formally recognised ranksthrough the International Code of Nomenclature(ICN, McNeill et al. 2012). These additional ranks areunder the ICN’s Art. 4.3 as the International Code ofNomenclature for Cultivated Plants (ICNCP) both at thetime (Trehane et al. 1995) as well as today (Brickell et al.2009) only deals with the naming of the unit of cultivar.(The one exception, grex, was (1995: Art. 4.6) and still is(2009: Art. 4.1) recognised by the ICNCP— but only for

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orchids.) The ICN allows additional ranks to be interca-lated, but demands they do not introduce confusion. Wedemonstrate here that Szabó & Hammer’s ranks do, asdid Flaksberger’s ranks before them (Table 1).

With Einkorn, Szabó & Hammer (1996) present twoseparate classifications. (1) Four geographical provenancegroups, called ‘races’ are distinguished — we assumethem equivalent to the geographically-definedsupraconvar. of their dicoccum classification. Races wereearlier used by Flaksberger (1935: 359) who, however,called them ‘proles’ and classifies them under a givensubspecies: for example, ‘race’ Helotinum Flaksb. withSzabó & Hammer equals ‘prol.’ heothinum Flaskb. (2)They then present an agrobotanical classification withprovarieties in the ‘convar. (subsp.)’ monococcum; anumber of these are sometimes synonymised with the‘vars.’ of other authors. The Einkorn provars. are listed assimply ‘var.’ (tom. cit. 14), and thus the proposedseparation of terms is abandoned: ‘varieties’ in theirbotanical and cultivated classifications are both equaland indicate the same rank. Szabó &Hammer’s Einkornclassification may appear exhaustive, but when com-pared with Flaksberger’s (1935: 355 – 368) earlier one,many of the latter’s taxa are missing. The impression isalso raised that a Szabó & Hammer provar. equalsFlaksberger’s ‘grex’ unit of cultivated taxa, but this isnot always so: provar. nigricultum Flaksb. is ‘grex’nigricultum, but provar. pseudohornemannii Dekapr. &Menabde is a ‘var.’ of the same name withFlaksberger.

When classifying Emmer (tom. cit. 15) the authors donot make ‘convar.’ but ‘supraconvar.’ equal to subspeciesand use convar. as a not further defined subgrouptherein. Further confusion is then introduced in that ‘…Convariety is a category in use for cultivated plantsindicating groups of botanical (our italics) varieties…’.The cultivated varieties of Emmer are listed with one as‘(pro)var.’, and all others as ‘var.’, but it is unclear if this isintentional or shorthand. This contrasts remarkably withthe Einkorn scheme. With spelta Szabó & Hammer(1996: 19) use both ‘subsp. (supraconvar.)’ and ‘subsp.’for the same subgroup: kuckuckianumGökgöl exDorof. etal. carries both denominations. Lastly, a new combina-tion of sinskajae within monococcum is presented as‘convar. et provar.’ — two different ranks at the sametime. As with the ICN, earlier botanical Codes

demanded a ‘clear indication of rank’ for a newcombination, which patently is not the case here.

These examples are cited to demonstrate theconfusion and easy contradiction that a hierarchical,detailed system can produce while attempting toclassify each and every slightly different cultivar. Itemanates from the so-called ‘German-Russian School’that has been proposing wheat classifications ofincreasing complexity (examples are Alefeld 1866,Körnicke 1885, Harz 1885, Vavilov 1923, Flaksberger1915 and 1935, and Mansfeld 1958). Before it isbrushed aside one should realise that it is stillconsidered relevant by authors like Hammer et al.(2011: 4) who present it as the only basis for formaldescription of new ‘races’ (probably meaning cultivarsand unlikely to be the categories of the Einkornclassification, discussed above).

We wonder what plant breeders would make of allthis as there is definitely another way for the classifi-cation of new cultivars.

The ‘genetic’ schoolIn contrast to the above, Thellung (1918a, b) hadalready built a classification of the wheats using,amongst others, considerations on phylogeny andpractical use of cereals. Notable in his scheme are:(1) a limitation in taxonomic ranks to species,subspecies and variety only (although referring tothe schemes of Alefeld and Körnicke & Werner), (2) adistinction as a single group of the three ‘spelt-wheats’[‘Spelzweizen’] (the Farro wheats of this paper) asopposed to the ‘naked’ wheats [‘Nacktweizen’], (3) anotion of phylogeny that has the naked wheats derivedfrom wild ancestors via the spelting ones, (4) arecognition that the evolution of the ‘naked’aestivum-compactum group has not derived from anyoriginal wild forms but from the spelting Triticumspelta, and (5) a distinction of three parallel,‘natürliche spezifische Gruppen oder Stämme’ [natu-ral specific groups or strains] (1918a: 470) exactlyalong what is now a sectional division of the wheats byploidy level! Thellung accepted 10 species, which here-grouped and classified formally at a lower rankunder only three accepted ones: T. monococcum,turgidum and aestivum. Thellung came to this conclu-sion in view of the ‘systematics and nomenclature’ (loc.

Table 1. Comparison of subgroups within cultivated species.

Flaksberger 1915/1935 Szabó & Hammer 1996 this publication

species/conspecies a species speciessubspecies supra(convar.)/subspecies/race subspeciesproles/subproles convar.grex/var. provar./var.var./forma cultivar (cv.)

aBoth species and conspecies were simultaneously used by Flaksberger in his 1915 Determination of Wheats paper.

CONCEPTS AND NOMENCLATURE OF THE FARRO WHEATS


cit.) of the group. The ground for a simple, flexibleand phylogenetically informative system was laid.

It is remarkable, therefore, that the extensive bibliog-raphy in Flaksberger’s later wheat monograph (1935:369 – 404) does not include Thellung’s 1918a, b paper.That omission, however, becomes more understandablewith the notion that Thellung’s points 2 – 5, as we quotethem above, are already present in a very similar (thoughmore elaborate) scheme in Flaksberger’s earlier‘Determination of wheats’ paper (1915: 17), which usedthree ‘conspecies’ where Thellung has ‘groups orstrains’. While the fundamental difference betweenFlaksberger and Thellung is the detailed subdivision ofTriticum species by the former, the ideas on phylogenyreflected in both schemes are essentially similar. (Theonly significant difference is the treatment of theexceptional polonicum forms: a ‘Monstrosität’ [monstros-ity] emanating from T. durum Desf. by Thellung, and anunexplained but entirely separate lineage byFlaksberger.) It would be easy to conclude thatThellung, although quoting Flaksberger’s 1915 paperfor a small nomenclatural matter, borrowed his ideas,changed them slightly (as in: simplifying with simulta-neous omission of some of Flaksberger’s more question-able lineages) but then did not acknowledge him as hissource. Obviously this cannot be proven.

More recently, Mac Key (1981) reflected on hisstudies since 1954 of wheat and other crops, and setout very similar principles to Thellung’s; van Slageren(1994) has agreed to this concept in the past and wedo so here again. Mac Key (1954, 1981) and others,such as Sears (1959) and Briggle & Reitz (1963), seethe wheat species, their names and possible subdivi-sions from a purely practical, often plant breeders’point of view. Genetics and crossability in the aestivumgroup lead Mac Key (1954: 582) to observations thatpolyploidy implied increased variation, but that theexpression of this is actually restricted by the bufferingeffect of gene duplication inherent in such polyploids.Moreover, easy crossability among the potentialgroups within aestivum sensu lato points at lack ofgenetic isolation (only one gene keeps the compactum,spelta and sphaerococcum groups apart from the coreaestivum, cf. Briggle & Reitz 1963: 35). These consid-erations pose a direct challenge to any purely (or atleast mainly) morphology-based system.

ConclusionsPlant breeders can exploit the Farro wheat species byidentifying desirable traits in individual accessions andthen crossing these with elite lines of bread and durumwheat, nowadays the only two wheat species of commer-cial importance. Therefore, where would that leave awheat ‘species’ beyond the levels of species and/orsubspecies as defined by their genome type, as Mac Key(1981: 205 ‘…more appropriate to discard the variety

concept and go directly to the cultivars…’) questions?The species in his system are de facto identified by theirgenome type, but using a subspecific status allows foruseful distinction within such a lumped group: Triticumaestivum can still be split into subsp. aestivum (sensustricto), compactum (Host) Domin, macha (Dekapr. &Menabde) Mac Key, spelta (L.) Thell., and sphaerococcum(Percival)Mac Key, even though these are closely relatedand all possess the hexaploid BAD genome. Wetherefore disagree with Szabó & Hammer’s (1996: 15)assertion that lumping in this way cannot distinguishbetween hulled and naked-seeding wheats: within tetra-ploid T. turgidum the two subsp. dicoccum (Emmer) anddicoccoides (Körn. ex Asch. & Graebn.) Thell.) are hulled,but all other subspecies, for example turgidum, carthlicum(Nevski) Á. Löve & D. Löve, polonicum (L.) Thell., anddurum, are not. Defined in this way species andsubspecies are meaningful units, provided that reason-ably observable differences among them exist. Incontrast to Hammer et al. (2011) we think that the valueof Farro and other wheats lies at the cultivar level and intheir possible possession of desirable traits for a partic-ular breeding programme.

In terms of communication the name of any cultivarcan simply be stated as prescribed by Art. 8 of the 2009edition of the ICNCP: ‘…a combination of the name ofthe genus or lower taxon to which it is assigned with acultivar … epithet.’ (for example, Triticum aestivumsubsp. aestivum ‘Florence Aurore’, or T. turgidum subsp.durum ‘Nab el Jamal’ [camel’s tooth], the latter an olddurum cultivar from Lebanon and Syria). This approachhas another, societal effect. We define a cultivated wheatspecies as a collection of cultivars, ‘held together’ by abotanical name with its reference point, the type (not anew concept, actually, but one already applied bybreeders like de Vilmorin in 1905; see below). With thecultivar in the spotlight, attention turns to the effortsneeded in creating them, whether by (participatoryplant-) breeding methods, leading to ‘old’ or ‘modern’cultivars, or through selection by the farmers themselvesover lengthy periods of time— the so-called ‘landraces’.Protection of the ‘intellectual capital’ of cultivarsthrough registration with the UPOV, the Unioninternationale pour la Protection des ObtentionsVégétales/International Union for the Protection ofNew Varieties of Plants, can be achieved as long as thematerial is novel and fulfils criteria on Distinctness,Uniformity and Stability (‘DUS’) (Z. Bishaw, pers. comm.).For farmer-developed landraces recognition of ‘Farmer’sRights’ cannot be obtained through UPOV as theirmaterial is unlikely to be considered novel, but this ispossible since 2004 under the International Treaty onPlant Genetic Resources for Food and Agriculture(ITPGRFA, see: http://www.planttreaty.org/content/farmers-rights). Our short digression illustrates what isat stake, for instance when considering Szabó &Hammer’s (1996: 14) relatively recent description of



http://www.planttreaty.org/content/farmers-rights

http://www.planttreaty.org/content/farmers-rights

the Einkorn provar. clusii A. T. Szabó & K. Hammer,which is defined as a ‘group of landraces’. (The citedtype collection, not seen by us, is thus either aheterogeneous gathering or obscures all but one ofthese landraces.) At the time they did not indicatewhether any measure of protection or recognition wasbeing considered; however, under the ITPGRFA this isnow (since 2004) possible. We wonder if formal recog-nition in a provar was actually relevant. Why not sampleand keep each landrace separate (assuming this can bedone, of course), describe each, name them (if stillneeded) according to the ICNCP, and then submit forprotection under the ITPGRFA?

The correct names of Einkorn and Spelt wheatWith the cultivar the primary tool for both the geneticresources scientist and the farmer, we believe its name,as constructed above, will be sufficient and thus amorphology-driven superstructure unnecessary. Thisconcept is adopted here and we see Einkorn, Emmerand Spelt as species in the sense of de Vilmorin andMac Key. As a result all published names of taxa belowthe level of (sub-)species will not be considered, if notactively discouraged from being used. Hence ourdiscarding of 18, 75 and 61 taxa within monococcum,dicoccum, and spelta respectively, as used in theclassification of Szabó & Hammer (and the manymore names proposed by, for example, Flaksberger(1935) and Dorofeev & Korovina (1979)).

The resulting names of Einkorn and Spelt are notdisputed and are presented in the nomenclaturesummary below.

Considerations on the correct name of EmmerwheatIn contrast to Einkorn and Spelt, for the correct, Latinname of Emmer three alternatives have been pro-posed. All of them have been supported in view ofambiguity of the protologue and the application — orlack of it — of the rules of the botanical Code:

1.Triticum dicoccon Schrank, Bayer. Fl. 1: 389 (1789)— dated to Schrank and at species level. The finalepithet uses the acceptable Greek neuter form,ending with –on, which does not need correctingto a Latinised form. This name was accepted by,for example, Mansfeld (1958: 238), Bor (1968:204; 1970: 207), Humphries (1978: 368), Tan(1985: 251), Szabó & Hammer (1996: 15) andHammer et al. (2011: 5)

2.Triticum spelta L. var. (‘*’) dicoccon Schrank, Bayer. Fl.1: 389 (1789) — dated to Schrank and interpreted asbeing at varietal rank (see below for a comment onthe asterisk). This name, which is cited as a synonymby, for example, Tzvelev (1973: 41, 1976: 165/1983:

237), Kerguélen (1975: 279), Cai et al. (1991: 221),and van Slageren (1994: 91), is the basionym of thethird alternative;

3.Triticum dicoccum ‘(Schrank) Schübl., Diss. char.descr. cereal. 29 (Schübler 1818)’ or with the authorsas ‘Schrank ex Schübl. (1818)’ or just ‘Schübl.(1818)’. This alternative uses the Latinised versionof the original Greek epithet. The authorship‘(Schrank) Schübl.’ is used by, for instance,Flaksberger (1935: 285), Tzvelev (1973: 41),Kerguélen (1975: 279), and Gontcharov (2011: 9),and is correct only when Schrank’s name isconsidered to have been published validly and at arank other than species. ‘Schrank ex Schübl.’ (aswith Morrison 2007: 272) or just ‘Schübl.’ should beused when Schrank’s name is considered invalid.

Schrank’s 1789 Bayerische Flora and Schübler’s 1818Dissertatio are the critical publications in addressingthe dicoccon/-um issue.

Schrank (1789: 388) described Triticum spelta as no.263 on p. 388 in his flora, adding ‘*dicoccon’ on p. 389(Fig. 1), followed by a diagnosis in German — not inLatin as Mansfeld (1958: 237) suggests — andindicating that dicoccon is not a particular form ofEinkorn wheat (‘…Es ißt das Emmerkorn gewiß keineSpielart des Einkorns…’), but, if anything, somethingcloser to spelta, as he writes at the end of that species:‘…Hierher, denke ich, gehört das Emmerkorn…’ [‘…here, I believe, belongs the Emmer wheat…’]. Hisdicoccon thus appears to be part of spelta and, inaddition, does not receive its own number but anasterisk (‘*’) instead. The use of the asterisk before theepithet indicated that it had at the time not yet beenfound in the region of his flora, i.e. Bavaria (Mansfeld1958: 237) — indeed, Schrank refers to its cultivation inthe nearby region of Württemberg. In contrast to this,however, Schrank does not use for his dicoccon a Greekletter (α, β, etc.) as he does for his infraspecific taxa(for example with the subdivision into α aestivum andβ hybernum of his T. cereale). Tzvelev, Kerguélen, Cai et al.,and van Slageren are therefore erroneous in theirinterpretation as an intended variety. That is not to saythat Schrank sees Emmer as a species, however. Henot only indicates its closeness with spelta, but thenwrites: ‘…wenn es nicht eine eigene Art ißt …’ [whenit is not a separate species] (Fig. 1). In other words,did he actually consider it less distinct than speltaitself, therefore not deserving species status, and thusimplicitly as belonging under it? Then status as avariety under spelta is logical, and this was the earlierinterpretation as ‘spelta var. dicoccon’ by the quotedauthors. Doubt about separate species status is alsoexpressed by Humphries (1978: 368); however, hethen immediately states that ‘…although Schrank mayhave had some doubt about its status he still wentahead and published it as a species.’



We present these three alternatives for the same nameand their respective arguments in order to show thatconfusion has dogged the Schrank name almost to thepresent day. However, after his remarks on the status ofEmmerkorn Schrank writes ‘…; ich nenne es einsweilen…’

[‘…; I call it for the time being…’]. The ‘einsweilen’makes Triticum dicoccon Schrank provisional and thusinvalid for Art. 36.1(b) (Greuter in Morrison 1998: 708).We agree, and this causes the name T. dicoccum (Schrankex) Schübl. from 1818 to come into view.

Schübler’s 1818 publication describes Triticumdicoccum (thus with a Latin ending of the epithet) whilereferring indirectly to the earlier Schrank name. Herefers in a footnote (loc. cit. 30 – 31) to Roemer &Schultes (1817: 766), commenting that the latter ‘…citeT. dicoccon Schrank as a synonym of T. zea Host while

there are many differences between the two, such as theglumes [of T. zea] only partly covering the spikelets withmature seeds and [the spike] rachis flexible…’ (bothgood characters of T. spelta where Host’s T. zea actuallybelongs). Thus a link with Schrank exists, albeit indirect.Shortly thereafter Schübler (1820: 451) re-described andillustrated dicoccum in great detail, listing dicocconSchrank — but not T. zea — as a synonym, thusconsidering the earlier dicoccon the same species butdid not accept Schrank’s name as the ‘valid’ one.According to the ICN Art. 46.5 the name should beascribed to the validating author and his/her spelling,but reference to the earlier, invalid publication at thesame time can be made: Triticum dicoccum ‘Schrank exSchübl.’ or just ‘Schübl.’. To recognise Schrank’s role inall this we recommend the first form of author citation; it

Fig. 1. Triticum dicoccon as described by Schrank in his Bayerische Flora, Vol. 1.



can be based on the 1818 Schübler publication, but ifone considers this not a strong enough reference, thencertainly on his 1820 one.

Hammer et al. (2011: 5) are prepared to put the ICBNrules aside for the sake of stability in order tomaintain thatTriticum dicoccon Schrank 1789 was validly published. Theyrefer to Art. 34.1(b) of the ICBN (McNeill et al. 2006) —now Art. 36.1(b) of the ICN — ruling provisional namesinvalid, but declare this to lead to ‘nomenclatural un-certainties’ that should be avoided. In this they are wrong.The question is whether T. dicoccon Schrank is, or is not,invalid, and not whether the Code is inconvenient or not.Hammer et al., and earlier Mansfeld (1958: 238), conclud-ed that in case Schrank’s name is not acceptable, T. farrumBayle-Bar. from 1809 should be adopted as the speciesname. No explanation is provided, however, as to why thiswould be so. Even if they were right there are optionsbetween 1789 and 1809 that should have been consideredas well. Most of these have appeared in synonymy, but notin treatments by genetic resources scientists such asMansfeld, Hammer and others, but by botanists such asKerguélen (1975: 279 – 280) and Humphries (1978: 368).

Names published before Schübler 1818 relatingto EmmerSpecies names of taxonomic synonyms of Triticumdicoccon Schrank, published between 1789 and Nov.1818, the date of Schübler’s valid publication as T.dicoccum Schrank ex Schübl.:

1.Triticum album Gaertn., Fruct. Sem. Pl. 2(1): 8, Tab.81, fig. 1 album ‘a–i’ (Gaertner 1790) (Fig. 2).(Our notation relating to the illustration summa-rises the letters connected with the analyticaldrawings ofT. album.) This species is notmentionedby Humphries in 1978, while cited by Kerguélen(1975: 278) under T. aestivum as ‘T. album Gaertnerex Steudel, Nomencl. Bot. ed. 1: 853 (1821), prosyn.’. Kerguélen apparently relied on Steudel’sNomenclator, both for the name and its authorship,but examination of all original elements presents anambiguous identity. This problem could be solved ifa type specimen were known but the herbariumTUB that houses Gaertner’s types does not hold anysuch collection. Gaertner’s album illustration depictsa dicoccum as here understood, including the prom-inent glume keel running from apex to base, whichis a turgidum-wheat character (Fig. 2). The phrasedescriptions on p. 8 present amixed identity: Bauhin(1623: 21), Morison (1699: 175), and Linnaeus(1774: 108) all refer to either awned aestivum orawnless hybernum forms of bread wheat. Malpighi(1687: 89, Tab. 54, fig. 325 A – G) only shows someviews of a wheat grain and is not conclusive. DeTournefort (1719, 1: 512, and its reference to Vol. 3)Tab. 293, Fig. F is different. Tabula 293 shows twospikes, of which Fig. Q is T. dicoccum and Fig. T is

T. turgidum (Fig. 3); the referred Fig. F depicts awheat grain in Tab. 292. Thus, Gaertner’s ownillustration and (part of) de Tournefort’s Tab.293 present links with Emmer. Kerguélen did notlectotypify album when he placed it under breadwheat, and were it not for the two illustrations itmay well belong there. Lectotypification may bedesirable for stability of use but can never dojustice to all elements associated with the name.This makes it unsuitable as the oldest correctname for dicoccum at species level.

2. Triticum spelta sensu Host, Icon. Descr. Gram. Austriac.3: 21, Tab. 30 (1805), non Linnaeus (1753). Whilethere is direct reference to Linnaeus’s Species Plantarum(1762: 127) the illustration of Tab. 30 clearly depicts adicoccum as here understood; in addition no collectionin the Host herbarium at W exists in connection withthis name.Mansfeld (1958) considers theHost ‘species’a synonym of his T. dicoccon Schrank. While older thanthe following two it is effectively a misapplication of theLinnaean name and not a contender.

3. Triticum atratum Host, Icon. Descr. Gram. Austriac. 4:5, Tab. 8 (1809). We were informed that as withT. spelta sensu Host no specimen for this name isknown in the Host herbarium in W, and haveassigned a type in the form of Tab. 8 (Fig. 4). Wecould not establish the date of publication withmore precision than the year 1809 (as in Stafleu& Cowan 1979: 340); this is the same year as thenext entry, T. farrum.

4. Triticum farrum Bayle-Bar., Monogr. Agron. Cereal. 1:50, Tab. 4, figs 1 – 2 (Bayle-Barelle 1809) (Fig. 5). Asfor atratum we have selected an illustration as the type(see below) as there is no herbarium known to existthe can be linked to Giuseppe Bayle-Barelle(Stafleu & Mennega 1992: 398; confirmed incorrespondence with PAV, the most likely placefor any such collection).

5. Triticum bauhini Lag., T. cienfuegos Lag. andT. gaertnerianum Lag. from Lagasca’s ElenchusPlantarum 6 (1816) will not be further consideredas T. farrum is older, validly published and taxo-nomically Emmer wheat. These (and many other)species are now heterotypic synonyms under theaccepted T. turgidum subsp. diccocum.

6. Triticum amyleum Ser. Mél. Bot. 1(2): 124 (Seringe,Sept. 1818) has been used as the species ‘representing’dicoccum, for example by Seringe (1818) himself,Metzger (1824) and Henri de Vilmorin (1880). It hasalso beenpresented in addition to dicoccum, for exampleby Schübler (1820, next to dicoccon Schrank) andPhilippe de Vilmorin (1905). A type specimen in Ghas been selected (see below).

We agree with Kerguélen (1975: 280) that a choicecannot, in fact, be made between Triticum atratum andT. farrum, and there is no reason why farrum shouldhave preference until an earlier publication date in1809 can be proven for it. It is only in following the



established ‘tradition’ of Mansfeld and Hammer,discussed above, that we recommend T. farrum as theaccepted name at species level for Emmer wheat.

Conclusion: considerations for the name of Emmerwheat at subspecific levelAs to what Emmer wheat represents we agree withPhilippe de Vilmorin’s (1905: 328) Hortus Vilmorinianus:the wheat groups are what he calls Variétés agricoles[agricultural varieties], united under a botanical name,whether this is at species (as ‘vulgare Vill.’ where T.aestivum would be correct), subspecies (for example‘vulgare Vill. subspec. T. durum Desf.’ — just ‘subspec.durum’ would be correct) or varietal (‘var. dicoccumSchrank’) rank. Interestingly, de Vilmorin does not addthe notion ‘Variétés agric.’ after his var. dicoccum, butwe consider this unintentional. His location of dicoccumas a variety under cultivated monococcum is not arguedbut illustrates their supposedly close link. With spelta asubspecies under T. vulgare the practical grouping ofthe hulled Farro species vs the free-threshing ones isnot adopted.

Building on this concept and taking note ofThellung’s and Mac Key’s considerations for an overallwheat classification, Emmer (and the other Farrowheats) can be located with precision within thegenus, now defined by the Gene Pool concept andsubdivided along ploidy levels and genome types. Theoldest available name for Emmer at subspecific level isthen Triticum turgidum L. subsp. dicoccum (Schrank exSchübl.) Thell. (see below).

Annotated summary of nomenclature (acceptednames at both species and subspecies rankin bold; only homotypic synonyms and selectedheterotypic synonyms included)

EINKORN (2x = 2n = 14)Triticum monococcum L. (Linnaeus 1753: 86); Metzger(1824: 35 – 36); (Körnicke 1885: 104); H. de Vilmorin(1880: 13, 21); Hackel (1887: 80); Fiori (1896: 108);Husnot (1899: 81); Ph. de Vilmorin (1905: 328);Thellung (1912: 140, Thellung 1918a: 470, 1918b: 146);

Fig. 2. Gaertner’s (1790) De Fructibus et Seminibus Plantarum Volumen Alterum, Tab. 81, fig. 1 showing Triticum album withdetails ‘a–i’. (Adapted from the original Gaertner plate.).



Fig. 3. De Tournefort’s (1719) Institutiones Rei Herbariæ, Tomus Tertius, with Tab. 293, fig. Q, showing a spike of Triticumdicoccum.



Fig. 4. The illustration with analysis of Tab. 8 in Host’s (1809) Icones et Descriptiones Graminum Austriacorum, Vol. 4, selected asthe lectotype of Triticum atratum.



Percival (1921: 154, 170); Flaksberger (1915: 21, 1935:355); Kerguélen (1975: 279); Hammer et al. (2011: 4);

Gontcharov (2011: 9). Type: [not located] (lectotypeLINN 104.4!, selected by Bowden (1959: 664)).

Fig. 5. The illustrations with analysis of Tav. IV [Tab. 4], figs. 1 – 2, in Bayle-Barelle’s (1809) Monografia Agronomica dei Cereali,selected as the lectotype of Triticum farrum. IMAGE © BRITISH LIBRARY BOARD, C.21.C44 PG 118. REPRODUCED WITH PERMISSION.



Nivieria monococca (L.) Ser. (Seringe 1842: 114,‘monococcum’). Type: as for T. monococcum. Note:although the separate genus Nivieria was an-nounced earlier in 1841 by Seringe, the actualname was not validly published until the yearafter.

Triticum vulgare Vill. convar. [‘Var.-Gr.’] monococcum(L.) Alef. (Alefeld 1866: 333). Type: as for T.monococcum.

Triticum monococcum L. var. vulgare Körn. (Körnicke1885: 111, 112); Percival (1921: 175). Type: as forT. monococcum. Note: epithet and thus name is notinvalid for Art. 24.3, although indicating the type ofthe next higher taxon.

Triticum sativum Lam. subsp. monococcum (L.) Voss(1895: 1218). Type: as for T. monococcum.

Triticum monococcum L. subsp. (‘B.’) cereale Asch. &Graebn. (Ascherson & Graebner 1901: 702);Thellung (1918a: 470, 1918b: 146), nom. illegit.(Art. 52.1). Type: as for T. monococcum. Note:rankless name and superfluous final epithet(monococcum should have been used). The rankappears to be considered a variety by Thellung(1918a: 470, ‘subsp. II. cereale (Ascherson etGraebner pro var)’) who then raises it to subspe-cies. However, this seems to be a contradiction tohis interpretation elsewhere of A., B., I., II., etc.from Ascherson & Graebner’s Synopsis as subspe-cies. For example, Thellung (1912: 143) earlierconsidered T. ovatum (L.) Rasp. ‘B. I.’ triaristatum(Willd.) Asch. & Graebn. a subspecies; vanSlageren (1994: 325) in a wider discussion on therankings in the Synopsis agreed. Mansfeld (1958:237) considers T. sativum Lam. ‘B. II.’ dicoccum(Schrank ex Schübl.) Asch. & Graebn. to be asubspecies also.

Triticum aestivum L. var. monococcum (L.) L. H. Bailey(1923: 133). Type: as for T. monococcum.

Crithodium monococcum (L.) Á. Löve (1984: 490). Type:as for T. monococcum.

Triticum monococcum L. subsp. monococcum.Selected literature using this subspecies to indicatecultivated material as opposed to the ‘wild’ (or‘spontaneous’, as with Szabó & Hammer 1996: 6)subspecies: Mac Key (1966: 267, 2005: 39); de Wet(1981: 192); van Slageren (1994: 89).

(diploid species with 2x = 2n = 14)Triticum sinskajae A. Filat. & Kurk. (Filatenko &Kurkiev 1975: 239); van Slageren (1994: 85);Morrison (2007: 272); Hammer et al. (2011: 4). Type:(Russian Federation, Daghestan autonomous region)[a free-threshing mutant,] isolated at the Derbentbranch of the N. I. Vavilov Institute from a collectionof cultivated T. monococcum made by P. M. Zhukovskyin 1926 in Dadaj, Kastamonu region, Turkey, [andpreserved as a herbarium voucher on] 29 June 1970

[by] U. Kurkiev & A. Filatenko (holotype WIR K-48993;isotypes WIR-Derbent no. 20970, [former WIR branchat] Tashkent [Uzbekistan]).Triticum monococcum L. ‘convar. et provar.’ sinskajae (A.

Filat. & Kurk.) A. T. Szabó & K. Hammer (1996:14), nom. invalid. (Art. 37.1: no clear indication ofrank). Note: Szabó & Hammer (1996: 6) expresslyconsider provar. and convar. of separate rank intheir classification. Hence their new combination ispublished at two ranks at the same time andtherefore invalid.

Triticum monococcum L. var. sinskajae (A. Filat. & Kurk.)Mac Key (2005: 39). Type: as for T. sinskajae.

Triticum monococcum L. subsp. sinskajae (A. Filat. &Kurk.) Valdés & H. Scholz (2006: 661). Type: as forT. sinskajae. Note: this name was cited earlier bySzabó & Hammer (1996: 14) but as ‘auct. dif.’without indicating who these might be; indeed,they were only formally established 10 years later.

EMMER (2x = 4n = 28)Triticum turgidum L. subsp. dicoccum (Schrank exSchübl.) Thell. (Thellung 1918a: 470, 1918b: 146); MacKey (1966: 268, 2005: 41); Kerguélen (1975: 279); vanSlageren (1994: 91). Type: the newly selected type ischosen for the basionym, T. dicoccum Schrank exSchübl. (q.v.)Triticum dicoccon Schrank (1789: 389); Schübler & von

Martens (1834: 46, ‘dicoccum’); Harz (1885: 1217,‘dicoccum’); Flaksberger (1915: 22, ‘dicoccumSchrnk.’); Mansfeld (1958: 238), Bor (1968: 204;1970: 207); Humphries (1978: 368), Tan (1985:251); Szabó & Hammer (1996: 15); Hammer et al.(2011: 5), nom. invalid. (Art. 36.1(b)).

Triticum dicoccum Schrank ex Schübl. (Schübler 1818:29; 1820: 450, Plate 1, fig. 2a–c – with ‘Trit. dicocconSchrank’ as a synonym on p. 451); Percival (1921:155, 186); Flaksberger (1935: 285); Tzvelev (1973:41); Kerguélen (1975: 279); Morrison (2007: 272,‘Schrank ex Schübl.’); Gontcharov (2011: 9). Type:a specimen in the von Schreber herbarium in M,bearing the label annotations ‘Triticum aestivumspicis incanis 1768’ and ‘hoc nominum missum estTriticum Dicoccum Sch.’ (lectotype M-0187352!,selected here (Fig. 6)). Notes: (1) Of the labelannotations, the line ‘Triticum aestivum spicisincanis’ and the year ‘1768’ were written bySchrank, while ‘hoc nomine missum est Triticumdicoccum Sch.’ was not written at the same time, isin a different ink, and most probably not bySchrank. This makes sense as Schrank would nothave written ‘missum est’ [was sent]. These wordscan be compared with those on the separateSchreber label on the sheet — it could well bethe same hand (Schreber’s?, which would makesense). It seems quite possible that Schrank sentthe specimen to Schreber in 1768, under the



name ‘Triticum aestivum spicis incanis’, and thatSchreber later added ‘This is Triticum dicoccumSch[rank].’. It thus appears that the epithet dicoccumwas coined already some time before 1789 but withthe Latin –um ending, not the Greek –on that was usedwith the formal publication. (2) Earlier than thislectotypification, Bor (1970: 207) and Kerguélen(1975: 279) noted at ‘type’: ‘Die von Stuttgartgesandten Samen gägen nicht nur im kaltenBoden, sondern sogar im Blumentöpfevierblütige zweisaamige Aehrchen’, Schrank.’.This is a major part of the comment bySchrank at his *dicoccon publication in his Flora(Fig. 1), only omitting ‘H. R. Kerner’, the personin Stuttgart who sent him the seeds. Of course itis not a typification as such and its use ofGerman is also incorrect in parts. (3) Citedliterature attributes the species to Schübler 1818;however, Flaksberger, Tzvelev, Kerguélen, andGontcharov cite the author as ‘(Schrank)Schübl.’, while Morrison is one of the fewinstances of a correct author citation for thisspecies. Note also that Flaksberger changesposition compared to his 1915 paper (see atdicoccon Schrank). [The identification of the typespecimen, the interpretation of its annotations,and comments on the Schrank statement werekindly provided by Dr Hajo Esser (M).]

Triticum vulgare Vill. convar. [‘Var.-Gr.’] dicoccum(Schrank ex Schübl.) Alef. (Alefeld 1866: 331).Type: as for T. dicoccum.

Triticum vulgare Vill. subsp. dicoccum (Schrank exSchübl.) Körn. (Körnicke 1885: 41, 81, as (subsp.)‘Triticum dicoccum Schrk.’, to be corrected for Art.24.4); Mansfeld (1958: 237, cited as a synonym of T.dicoccon Schrank). Type: as for T. dicoccum. Note:Körnicke accepted only three species inTriticum (tom.cit. 40) and dicoccum and spelta (and others) areconsidered ‘Gruppen (Unterarten)’ [groups (subspe-cies)] of T. vulgare. Still the typography suggests thathe sees them at species level and the purported rankis not further mentioned. As all of these ‘groups’ havefew or many ‘Varietäten’ [varieties], both interpreta-tions of his classification would not be in conflict withthe accepted taxonomic hierarchy.

Sitopyrum dicoccum Döll, nom. invalid. (Art. 36.1(c)). Asquoted by Harz (1885: 1217) with the author of thename only as ‘Döll’ rather than ‘(Schrank exSchübl.) Döll’. Note: a name cited only in synonymyof T. dicoccum Schrank that appears to refer toDöll’s (1857 (1855): 123) unranked subgroup ‘I.Pyros’ of Triticum. The generic name is not foundelsewhere.

Triticum sativum Lam. subsp. dicoccum (Schrank exSchübl.) Hackel (1887: 81, 84); Voss (1895: 1218);Ascherson & Graebner (1901: 679, rankless as ‘B. II.’but this interpreted as subsp. — see at T. monococcum

Fig. 6. The voucher M-0187352, showing the lectotype ofTriticum dicoccum in herbarium M. IMAGE © BOTANISCHE

STAATSSAMMLUNG MÜNCHEN. REPRODUCED WITH PERMISSION.



subsp. cereale); Mansfeld (1958: 237, ascribes thename to Ascherson & Graebner 1901). Type: asfor T. dicoccum.

Triticum monococcum L. var. dicoccum (Schrank exSchübl.) P. Vilm. (Ph. de Vilmorin 1905: 328).Type: as for T. dicoccum.

Triticum aestivum L. var. (‘θ’) dicoccum (Schrank exSchübl.) Fiori (1896: 108). Type: as for T. dicoccum.

Triticum spelta L. subsp. [‘sous-esp. T. dicoccum’ — tobe corrected for Art. 24.4] dicoccum (Schrank exSchübl.) Husn. (Husnot (1899: 81). Type: as for T.dicoccum.

Triticum aestivum L. subsp. dicoccum (Schrank exSchübl.) Thell. (Thellung: 1912: 141). Type: as forT. dicoccum.

Gigachilon polonicum (L.) Seidl ex Á. Löve subsp.dicoccum (Schrank ex Schübl.) Á. Löve (1984: 497).Type: as for T. dicoccum.

Triticum album Gaertn. (Gaertner 1790: 8, Tab. 81, fig.1 album ‘a, g-i’) pro parte.

Triticum spelta sensu Host (1805: 21, Tab. 30);Mansfeld (1958: 237), non Linnaeus (1753).

Triticum atratum Host (1809: 5, Tab. 8); Metzger (1824:34, under T. amyleum). Type: the illustration withanalysis of Tab. 8 in Host’s 1809 Icones et DescriptionesGraminum Austriacorum Vol. 4, selected here(Fig. 4).

Triticum dicoccum Schrank ex Schübl. var. (‘β’) atratum(Host) Schübl. & G. Martens (Schübler & vonMartens 1834: 46, as ‘β T. atratum’ to be correctedfor Art. 24.4); Bluff et al. (1836: 203); Schrader invon Schlechtendahl (1838: 465); Körnicke 1885: 84,89); Percival (1921: 201); Flaksberger (1935: 317).Type: as for T. atratum.

Triticum vulgare Vill. (convar. [‘Var.-Gr.’] monococcum(L.) Alef.) var. atratum (Host) Alef. (Alefeld 1866:333). Type: as T. atratum. Note: Alefeld has atratumunder monococcum after investigating material fromMetzger in which he only found one seed perspikelet.

Triticum farrum Bayle-Bar. (Bayle-Barelle 1809: 50,Tav. 4, figs 1 – 2); Harz (1885: 1219, ‘Farrum’ and asan invalid ‘species’ under T. dicoccum). Type: theillustrations in Bayle Barelle’s 1809 Monografiaagronomica dei Cereali, Tav. 4, figs 1 – 2, selectedhere (Fig. 5). Note: the figures show the awnless(fig. 1) and awned (fig. 2) forms of T. farrumspikes, together with individual spikelets andglumes. Both figures are expressly included toshow the variety of forms that make up thisspecies as was intended by Bayle-Barelle. Theawnless form of Tav. 4, fig. 1 is named ‘T. farrummuticum’ but this is not formally described onpp. 50 – 52 of the book as a separate entity. On thecontrary, ‘Triticum farrum (mihi)’ is headed on p. 50 by‘Specie IX. Tav. 4. Fig. 1. e 2 [our italics].’, expresslyreferring to both figures.

Triticum vulgare Vill. (convar. dicoccum (Schrank exSchübl.) Alef.) var. farrum (Bayle-Bar.) Alef.(Alefeld 1866: 331). Type: as for T. farrum.

Triticum dicoccum Schrank ex Schübl. var. farrum(Bayle-Bar.) Körn. (Körnicke 1885: 84, 87);Flaksberger (1915: 64, 184); Percival (1921: 197).Type: as for T. farrum.

Triticum dicoccum Schrank ex Schübl. (subsp.europaeum Vavilov proles tardo-europaeum Flaksb.)grex farrum (Bayle-Bar.) Flaksb. (Flaksberger 1935:306). Type: as for T. farrum. Note: the grex containstwo varieties (‘var.’) and two formae (‘f.’), but noneis named farrum!

Triticum monococcum L. (var.) majus Dum. Cours(Dumont de Courset 1811: 110); Seringe (1818: 125;as a synonym of T. amyleum, and author as ‘Dum. deCours.’); Schübler (1820: 451, author as ‘Dumeril deCours’ [sic]); Harz (1885: 1217, as ‘T. monococcummajus Dumeril’ [sic]). Type: not indicated, andherbarium unknown (Stafleu & Cowan 1976). Note:Dumont de Courset writes at his entry formonococcum:‘Grande épeautre, T. monococcum majus’. Althoughthe French name is generally linked with T. spelta,when seen as a series of increasing size the next ‘stepup’ would logically refer to dicoccum where the majusname has indeed been located.

Triticum amyleum Ser. (Seringe 1818: 124); Metzger(1824: 30 – 35); H. de Vilmorin (1880: 13, 21).Type: a specimen from the herbarium of Charles-Isaac Fauconnet, acquired by Delessert for theherbarium in Geneva in 1879, bearing the printedlabel: ‘6. A. Triticum amyleum Ser. Spica aristata,alba, glabra; glumae mucrone incurvo. Ser.[Seringe] Mél. 1. p. 125.’ (lectotype G-00359729!,selected here). Note: although clear references toadequate illustrations by the Bauhin brothers arepresented by Seringe, plant material has preference(sensu Art. 9.12 of the Code, the lectotype being an‘uncited specimen’ as referred to in the Article),and the lectotype from G serves as such.

Spelta amylea (Ser.) Ser. (Seringe 1842: 114). Type: asfor T. amyleum. Note: although the genus name isillegitimate (for Art. 53.1) after Spelta Wolf thespecies name is not (Art. 55.1).

Triticum vulgare Vill. subsp. amyleum (Ser.) P. Vilm.(Ph. de Vilmorin 1905: 328, as ‘subspec. T.amyleum Ser.’ to be corrected for Art. 24.4). Type:as for T. amyleum.

Triticum tricoccum Schübl. (Schübler 1820: 458); Harz(1885: 1218). Type: the illustration with analysis ofPlate 1, fig. 3a – c in Flora 3(2) of 1820, referred toin Schübler’s paper on p. 458, selected here. Note:Dr Hajo Esser (M) informed us that no material wasfound in M that could be connected with Schüblerwith enough certainty to take precedence. Theillustration of a complete spike with spikelet andgrain details is perfectly adequate to show the



purported difference of three grains per spikeletcompared with the 2-grained ‘normal’ dicoccum spike-let and the 1-grained monococcum one.

Triticum dicoccon (Schrank) Schübl. var. (‘γ’) tricoccum(Schübl.) Schübl. & G. Martens (Schübler &Martens 1834: 46, as ‘γ T. tricoccum’ to be correctedfor Art. 24.4); Schrader in von Schlechtendahl(1838: 465); Döll (1857 (1855): 125). Type: as forT. tricoccum.

Triticum vulgare Vill. (convar. [‘Var.-Gr.’] dicoccon(Schrank ex Schübl.) Alef.) var. tricoccum (Schübl.)Alef. (Alefeld 1866: 332, ‘subtricoccum’ but withdirect reference to ‘Trit. tricoccum, Schübler’).Type: as for T. tricoccum.

SPELT (2x = 6n = 42)Triticum aestivum L. subsp. spelta (L.) Thell.(Thellung 1918a: 471, 1918b: 147); Mac Key (1966:268, 2005: 43); van Slageren (1994: 94); Soreng (2003:677). Type: as for T. spelta.Triticum spelta L. (Linnaeus 1753: 86, ‘Spelta’);

Metzger (1824: 26 – 30, ‘Spelta’); Schübler & vonMartens (1834: 46, ‘Spelta’ and with new varieties forawned (α aristatum) and unawned (β muticum)forms); Harz (1885: 1210, ‘Spelta’ and includingnumerous invalidly (for Art. 37.6) published‘species’); Husnot (1899: 80); Percival (1921: 158,329); Flaksberger (1935: 126); Kerguélen (1975:279); Cai et al. (1991: 222, with subsp. spelta);Hammer et al. (2011: 6); Gontcharov (2011: 9).Type: [not located] (lectotype LINN 104.1!,selected by Morrison (1998: 709)).

Spelta vulgaris Ser. (Seringe 1842: 114). Type: as forT. spelta.Note: although the genus name is illegitimate (forArt. 53.1) after Spelta Wolf 1776 the species name isnot (Art. 55.1). Seringe called this species ‘Epautrecommun’, clearly suggesting it is a renaming ofTriticum spelta and not of another species such asT. amyleum (q.v.).

Triticum vulgare Vill. convar. [‘Var.-Gr.’] spelta (L.) Alef.(Alefeld 1866: 334). Type: as for T. spelta.

Triticum vulgare Vill. subsp. spelta (L.) Körn. (Körnicke1885: 41, 75, as (subsp.) ‘Triticum Spelta L.’, to becorrected for Art. 24.4). Type: as for T. spelta.

Triticum sativum Lam. subsp. [‘Rasse’] spelta (L.) Hack.(Hackel 1887: 81, ‘Spelta’); Voss (1895: 1217);Ascherson & Graebner (1901: 676). Type: as forT. spelta.

Triticum vulgare Vill. subsp. spelta (L.) P. Vilm. (Ph. deVilmorin 1905: 328, as ‘subspec. T. spelta L.’ to becorrected for Art. 24.4). Type: as for T. spelta.

Triticum aestivum L. var. (‘η’) spelta (L.) Fiori (1896:108, ‘Spelta’); Bailey (1923: 133); Soreng (2003: 677,author as ‘(L.) L. H. Bailey’). Type: as for T. spelta.

Zeia spelta (L.) Lunell (1915: 226). Type: as for T. spelta.Triticum x aestivum L. emend. Bowden ‘cultivar group’

spelta (L.) Bowden (1959: 674). Type: as for T. spelta.

Triticum aestivum L. ‘varietal group’ spelta (L.) Morris &Sears (1967: 21) pro parte (considered to be basedon ‘T. spelta L. + T. macha Dek. & Men.’). Type: asfor T. spelta.

Triticum zea Host, Icon. Descr. Gram. Austriac. 3: 20, Tab.29 (1809); Roemer & Schultes (1817: 766); Harz(1885: 1214, under T. spelta). Type: the illustrationwith analysis of Tab. 29 from Host’s Icones etDescriptiones Graminum Austriacorum Vol. 3, selectedhere. Note: the Tab. 29 shows a clear but awnedspelta, which is less common than awnless cultivars.

AcknowledgementsAt K we thank Melanie Thomas for advice and transla-tion of Schübler’s (1818) Latin text, Paul Little for hisproduction of high-resolution images of the relevantillustrations, the staff at the Library, and Dr WolfgangStuppy (Seed Conservation Department) for his ‘Stuppy-fication’ of most of the illustrations, making them asuperior version of the originals. Furthermore we thankthe curators of G, M, PAV, TUB and W for their help inidentifying (possible) type collections, and the staff at theLibraries of the Natural History Museum (NHM) andthe British Library for their help in making the relevantliterature available. We also acknowledge the granting ofpermission by the British Library Board and theBotanische Staatssammlung München (M) to publishimages under their copyright. Dr Zewdie Bishaw of theInternational Center for Agricultural Research in theDry Areas (ICARDA) advised on UPOV and theITPGRFA. We thank the reviewers for useful suggestionsand improvements. This research is financiallysupported by the Stichting Elise Mathilde Fonds,CIMMYT, Int., and the COFRA foundation.

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