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

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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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Concepts and nomenclature of the Farro wheats, with special
reference to Emmer, Triticum turgidum subsp. dicoccum (Poaceae)
Michiel van Slageren
1
& Thomas Payne
2
Summary. This paper discusses the different taxonomic concepts of the wheat group as exemplied 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 classied at subspecies rank, and to be dened 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 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 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.
Key Words. taxonomy, wheat.
Introduction
Farro is an Italian ethnobotanical concept (Szabó &
Hammer 1996: 3) and applies to three hulled,ornot
free-threshingwheat taxa, that is to say wheats in
which the spike rachis breaks up under pressure and
the spikelets fall with their glumes still attached. Free-
threshing or not is an important agronomic character
as it means the difference between an easy, almost
spontaneous, separation of the grains from the re-
mains of the ower and this being not so. In the latter
case threshing can still create the desired result but
achieving it is much more elaborate and thus costly:
an example from South Africa cited the cost for hulled
spelt wheat at roughly 5.5 times that of bread wheat
(D. van Papendorp, pers. comm.). The Farro wheats
are Einkorn (Triticum monococcum L. subsp.
monococcum), Emmer (T. turgidum L. subsp. dicoccum
(Schrank ex Schübl.) Thell.), and Spelt (T. aestivum
L. subsp. spelta (L.) Thell.); in terms of our general
concept of wheat taxa they are all considered at
subspecies rank. These three wheats are part of the
primary gene pool (or GP-1) of cultivated wheat. This
designation 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 four
groups, based around Einkorn, turgidum wheat
(T. turgidum L.), Timofeev wheat (T. timopheevii
(Zhuk.) Zhuk.), and bread wheat (T. aestivum L.)
(Harlan & de Wet 1971:g. 2). Szabó & Hammer (loc. cit.)
point out that hulled wheatsarenotthesameasnot
free-threshing, as many species from the wider wheat
gene pool (GP-2 and -3 in the scheme of Harlan & de
Wet) possess this character too, for example species of
genera such as Aegilops L., Amblyopyrum (Jaub. & Spach)
Eig (separation from Aegilops is sensu van Slageren 1994),
and Secale L. However, within GP-1 Triticum the two
are effectively the same: hulled wheats = Farro (but
see below).
Conversely, Szabó & Hammer consider the free-
threshing Triticum sinskajae Filat. & Kurkiev part of
Farro to illustrate it does not automatically mean
hulled. However, we question whether one can speak
of a speciesin this case (see the nomenclature
summary below). To our knowledge it has never been
commercially released as a cultivar to replace the
Accepted for publication 9 May 2013. Published online 22 June 2013
1
Royal Botanic Gardens Kew, Seed Conservation Department, Wakehurst Place, Ardingly, West Sussex, RH17 6TN, UK. e-mail: m.slageren@kew.org
2
The International Wheat and Maize Improvement Center (CIMMYT, Int.), Genetic Resources Program, Apdo. Postal 6-641, 06600 Mexico, DF, Mexico.
e-mail: T.Payne@cgiar.org.
KEW BULLETIN VOL. 68: 477 Y494 (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 than
describing and conferring a name and type we feel it
should have remained research material of plant
breeding interest.AsT. sinskajae is a published species
and is discussed here, its nomenclature is included
below, with typography indicating our opinion on the
status of this taxon.
While we equate Farro in this publication with the
three well-known, hulled wheat species, other Triticum
species are classied as being spelttype in contrast to
nakedor free-threshingtype. They may therefore
exhibit 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 originating
from Turkey. However, these are not Farro wheats as
commonly understood, and are not further consid-
ered here.
Grouping hulled (as in: Farro) wheats separate
from the free-threshing ones is an old concept, dating
back to pre-Linnaean times (Morrison 1998: 708). As
early as 1623, Bauhin (1623:2122) listed them in the
separate (from Triticum that is) genus Zea. As their
phrase-names include such words as Monococos,
dicoccosand Triticospeltum(the latter cited by
Trinius (1822: 377) as Tritico - speltumand identied
as T. spelta L.), it is clear which species are meant.
Several overviews and monographs of the wheat group
use 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 oras
when all hulled species are present, for example in
those from Schübler & von Martens (1834: 46), Döll
(1857 (1855): 124, as II. Zea,Spelzweizen), or
Ascherson & Graebner (1898: 124).
Different approaches in the classication
of Farro species
The precise naming and taxonomic position matters,
as the Farro species are somewhat of a resurgent crop.
Accordingly, the breeding value of all three is
presently the subject of renewed interest (Stallknecht
et al. 1996; Zaharieva et al. 2010). It is comparatively
easy to cross them with selections of the two main
wheat species, durum wheat (Triticum turgidum
L. subsp. durum (Desf.) Husn.) and bread wheat
(T. aestivum subsp. aestivum). Emmer enjoys increased
interest for a potential role in high-quality food;
accessions are known with comparatively high (though
variable) protein and mineral content, can be a source
for agronomically useful characters such as tillering
and grain weight, while resistance to rusts and bunt
are also recorded (Zaharieva et al. 2010). Transferring
these traits is not without problems but should be
comparatively easy as, for example, complete genomic
compatibility between the Emmer and durum wheat
genomes is reported (Zaharieva et al. 2010:953,
quoting a study by Yanchenko).
We realise that much wheat research is carried out
by plant breeders, genetic resources scientists, pathol-
ogists, entomologists and the like, and that therefore
the correct botanical name of the Farro species is
perhaps not a priority. However, taxonomic opinion
affects the naming of the genetic resources used by
these scientists, and reality is often different from the
ideal that correctly identied material is available. It is
here that the effect of two schools of thought, both
still very much in evidence, is most apparent.
The morphologyschool
Firstly, there has been the historical development (and
acceptance) of a detailed infraspecic classication
that aims to describe and appreciate the richness of
morphological variation found in the species, be this
wild or cultivated ones (Hammer et al. 2011). While
there is quite some history here, our exposure mainly
centres on a relatively recent paper by Szabó &
Hammer (1996). Although accepting many micro-taxa,
these authors nevertheless advocated (1996:45) the
need for a practical scheme to be widely adopted in
which ‘…wild taxa are consequently treated on the
subspecies and varietal levels, and cultivated taxa on the
convariety and (pro)variety levels…’ However, the
additional (supra)convar.as a second equivalent of
subsp.for cultivated material is then introduced, as are
the smallest categories of f. (forma)for wild taxa and
cv. (cultivar)for cultivated taxa. The distinction within
the ranks for cultivated groups is for largervs smaller
ones: several cvs. into a provar.; then, for reasons of
geographical distribution (tom. cit. 16), one or several
provars. are joined into a convar. or sometimes a
(supra)convar. The problems with this proposed practi-
cal scheme become immediately apparent when the
authors proceed to classify the Farro species. It should
also be noted that the ranks of provar., convar.,
supraconvar., grex, race, and proles as used by
Flaksberger, Hammer, Szabó and others for units of
(predominantly) cultivated plants are not cited but can
be intercalated between formally recognised ranks
through the International Code of Nomenclature
(ICN, McNeill et al. 2012). These additional ranks are
under the ICNs Art. 4.3 as the International Code of
Nomenclature for Cultivated Plants (ICNCP) both at the
time (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)andstillis
(2009: Art. 4.1) recognised by the ICNCP but only for
478 KEW BULLETIN VOL. 68(3)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2013
orchids.) The ICN allows additional ranks to be interca-
lated, but demands they do not introduce confusion. We
demonstrate here that Szabó & Hammers ranks do, as
did Flaksbergers ranks before them (Table 1).
With Einkorn, Szabó & Hammer (1996) present two
separate classications. (1) Four geographical provenance
groups, called racesare distinguished we assume
them equivalent to the geographically-dened
supraconvar. of their dicoccum classication. Races were
earlier used by Flaksberger (1935: 359) who, however,
called them prolesand classies them under a given
subspecies: for example, raceHelotinum Flaksb. with
Szabó & Hammer equals prol.heothinum Flaskb. (2)
They then present an agrobotanical classication with
provarieties in the convar. (subsp.)monococcum;a
number of these are sometimes synonymised with the
vars.of other authors. The Einkorn provars. are listed as
simply var.(tom. cit. 14), and thus the proposed
separation of terms is abandoned: varietiesin their
botanical and cultivated classications are both equal
and indicate the same rank. Szabó & HammersEinkorn
classication may appear exhaustive, but when com-
pared with Flaksbergers(1935:355368) earlier one,
many of the latters taxa are missing. The impression is
also raised that a Szabó & Hammer provar. equals
Flaksbergersgrexunit of cultivated taxa, but this is
not always so: provar. nigricultum Flaksb. is grex
nigricultum, but provar. pseudohornemannii Dekapr. &
Menabde is a var.ofthesamenamewith
Flaksberger.
When classifying Emmer (tom. cit. 15) the authors do
not make convar.but supraconvar.equal to subspecies
and use convar. as a not further dened subgroup
therein. Further confusion is then introduced in that ‘…
Convariety is a category in use for cultivated plants
indicating 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 is
intentional or shorthand. This contrasts remarkably with
the Einkorn scheme. With spelta Szabó & Hammer
(1996: 19) use both subsp. (supraconvar.)and subsp.
for the same subgroup: kuckuckianum kgöl ex Dorof. et
al. carries both denominations. Lastly, a new combina-
tion of sinskajae within monococcum is presented as
convar. et provar.’—two different ranks at the same
time. As with the ICN, earlier botanical Codes
demanded a clear indication of rankfor a new
combination, which patently is not the case here.
These examples are cited to demonstrate the
confusion and easy contradiction that a hierarchical,
detailed system can produce while attempting to
classify each and every slightly different cultivar. It
emanates from the so-called German-Russian School
that has been proposing wheat classications of
increasing complexity (examples are Alefeld 1866,
Körnicke 1885, Harz 1885, Vavilov 1923, Flaksberger
1915 and 1935, and Mansfeld 1958). Before it is
brushed aside one should realise that it is still
considered relevant by authors like Hammer et al.
(2011: 4) who present it as the only basis for formal
description of new races(probably meaning cultivars
and unlikely to be the categories of the Einkorn
classication, discussed above).
We wonder what plant breeders would make of all
this as there is denitely another way for the classi-
cation of new cultivars.
The geneticschool
In contrast to the above, Thellung (1918a,b) had
already built a classication of the wheats using,
amongst others, considerations on phylogeny and
practical use of cereals. Notable in his scheme are:
(1) a limitation in taxonomic ranks to species,
subspecies and variety only (although referring to
the schemes of Alefeld and Körnicke & Werner), (2) a
distinction as a single group of the three spelt-wheats
[Spelzweizen] (the Farro wheats of this paper) as
opposed to the nakedwheats [Nacktweizen], (3) a
notion of phylogeny that has the naked wheats derived
from wild ancestors via the spelting ones, (4) a
recognition that the evolution of the naked
aestivum-compactum group has not derived from any
original wild forms but from the spelting Triticum
spelta, and (5) a distinction of three parallel,
natürliche spezische Gruppen oder Stämme[natu-
ral specic groups or strains] (1918a: 470) exactly
along what is now a sectional division of the wheats by
ploidy level! Thellung accepted 10 species, which he
re-grouped and classied formally at a lower rank
under 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 species
subspecies supra(convar.)/subspecies/race subspecies
proles/subproles convar.
grex/var. provar./var.
var./forma cultivar (cv.)
a
Both species and conspecies were simultaneously used by Flaksberger in his 1915 Determination of Wheats paper.
479CONCEPTS AND NOMENCLATURE OF THE FARRO WHEATS
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2013
cit.) of the group. The ground for a simple, exible
and phylogenetically informative system was laid.
It is remarkable, therefore, that the extensive bibliog-
raphy in Flaksbergers later wheat monograph (1935:
369 404) does not include Thellungs1918a,bpaper.
That omission, however, becomes more understandable
with the notion that Thellungspoints25, as we quote
them above, are already present in a very similar (though
more elaborate) scheme in Flaksbergers earlier
Determination of wheatspaper (1915: 17), which used
three conspecieswhere Thellung has groups or
strains. While the fundamental difference between
Flaksberger and Thellung is the detailed subdivision of
Triticum species by the former, the ideas on phylogeny
reected in both schemes are essentially similar. (The
only signicant difference is the treatment of the
exceptional polonicum forms: a Monstrosität[monstros-
ity] emanating from T. durum Desf. by Thellung, and an
unexplained but entirely separate lineage by
Flaksberger.) It would be easy to conclude that
Thellung, although quoting Flaksbergers1915 paper
for a small nomenclatural matter, borrowed his ideas,
changed them slightly (as in: simplifying with simulta-
neous omission of some of Flaksbergersmorequestion-
able lineages) but then did not acknowledge him as his
source. Obviously this cannot be proven.
More recently, Mac Key (1981)reected on his
studies since 1954 of wheat and other crops, and set
out very similar principles to Thellungs; van Slageren
(1994) has agreed to this concept in the past and we
do so here again. Mac Key (1954,1981) and others,
such as Sears (1959) and Briggle & Reitz (1963), see
the wheat species, their names and possible subdivi-
sions from a purely practical, often plant breeders
point of view. Genetics and crossability in the aestivum
group lead Mac Key (1954: 582) to observations that
polyploidy implied increased variation, but that the
expression of this is actually restricted by the buffering
effect of gene duplication inherent in such polyploids.
Moreover, easy crossability among the potential
groups within aestivum sensu lato points at lack of
genetic isolation (only one gene keeps the compactum,
spelta and sphaerococcum groups apart from the core
aestivum, cf. Briggle & Reitz 1963: 35). These consid-
erations pose a direct challenge to any purely (or at
least mainly) morphology-based system.
Conclusions
Plant breeders can exploit the Farro wheat species by
identifying desirable traits in individual accessions and
then crossing these with elite lines of bread and durum
wheat, nowadays the only two wheat species of commer-
cial importance. Therefore, where would that leave a
wheat speciesbeyond the levels of species and/or
subspecies as dened 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 identied by their
genome type, but using a subspecicstatusallowsfor
useful distinction within such a lumped group: Triticum
aestivum can still be split into subsp. aestivum (sensu
stricto),compactum (Host) Domin, macha (Dekapr. &
Menabde) Mac Key, spelta (L.) Thell., and sphaerococcum
(Percival) Mac Key, even though these are closely related
and all possess the hexaploid BAD genome. We
therefore disagree with Szabó & Hammers(1996:15)
assertion that lumping in this way cannot distinguish
between hulled and naked-seeding wheats: within tetra-
ploid T. turgidum the two subsp. dicoccum (Emmer) and
dicoccoides (Körn. ex Asch. & Graebn.) Thell.) are hulled,
but all other subspecies, for example turgidum,carthlicum
(Nevski) Á. Löve & D. Löve, polonicum (L.) Thell., and
durum,arenot.Dened in this way species and
subspecies are meaningful units, provided that reason-
ably observable differences among them exist. In
contrast to Hammer et al. (2011) we think that the value
of Farro and other wheats lies at the cultivar level and in
their possible possession of desirable traits for a partic-
ular breeding programme.
In terms of communication the name of any cultivar
can simply be stated as prescribed by Art. 8 of the 2009
edition of the ICNCP: ‘…a combination of the name of
the genus or lower taxon to which it is assigned with a
cultivar epithet.(for example, Triticum aestivum
subsp. aestivum Florence Aurore,orT. turgidum subsp.
durum Nab el Jamal[camels tooth], the latter an old
durum cultivar from Lebanon and Syria). This approach
has another, societal effect. We dene a cultivated wheat
species as a collection of cultivars, held togetherby a
botanical name with its reference point, the type (not a
new concept, actually, but one already applied by
breeders like de Vilmorin in 1905; see below). With the
cultivar in the spotlight, attention turns to the efforts
needed in creating them, whether by (participatory
plant-) breeding methods, leading to oldor modern
cultivars, or through selection by the farmers themselves
over lengthy periods of time the so-called landraces.
Protection of the intellectual capitalof cultivars
through registration with the UPOV, the Union
internationale pour la Protection des Obtentions
Végétales/International Union for the Protection of
New Varieties of Plants, can be achieved as long as the
material is novel and fulls criteria on Distinctness,
Uniformity and Stability (DUS) (Z. Bishaw, pers. comm.).
For farmer-developed landraces recognition of Farmers
Rightscannot be obtained through UPOV as their
material is unlikely to be considered novel, but this is
possible since 2004 under the International Treaty on
Plant Genetic Resources for Food and Agriculture
(ITPGRFA, see: http://www.planttreaty.org/content/
farmers-rights). Our short digression illustrates what is
at stake, for instance when considering Szabó &
Hammers(1996: 14) relatively recent description of
480 KEW BULLETIN VOL. 68(3)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2013
the Einkorn provar. clusii A. T. Szabó & K. Hammer,
which is dened as a group of landraces.(Thecited
type collection, not seen by us, is thus either a
heterogeneous gathering or obscures all but one of
these landraces.) At the time they did not indicate
whether any measure of protection or recognition was
being considered; however, under the ITPGRFA this is
now (since 2004) possible. We wonder if formal recog-
nition in a provar was actually relevant. Why not sample
and keep each landrace separate (assuming this can be
done, of course), describe each, name them (if still
needed) according to the ICNCP, and then submit for
protection under the ITPGRFA?
The correct names of Einkorn and Spelt wheat
With the cultivar the primary tool for both the genetic
resources scientist and the farmer, we believe its name,
as constructed above, will be sufcient and thus a
morphology-driven superstructure unnecessary. This
concept is adopted here and we see Einkorn, Emmer
and Spelt as species in the sense of de Vilmorin and
Mac Key. As a result all published names of taxa below
the level of (sub-)species will not be considered, if not
actively discouraged from being used. Hence our
discarding of 18, 75 and 61 taxa within monococcum,
dicoccum,andspelta respectively, as used in the
classication of Szabó & Hammer (and the many
more names proposed by, for example, Flaksberger
(1935) and Dorofeev & Korovina (1979)).
The resulting names of Einkorn and Spelt are not
disputed and are presented in the nomenclature
summary below.
Considerations on the correct name of Emmer
wheat
In contrast to Einkorn and Spelt, for the correct, Latin
name of Emmer three alternatives have been pro-
posed. All of them have been supported in view of
ambiguity of the protologue and the application or
lack 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 nal
epithet uses the acceptable Greek neuter form,
ending with on, which does not need correcting
to 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) and
Hammer et al. (2011:5)
2.Triticum spelta L. var. (*)dicoccon Schrank, Bayer. Fl.
1: 389 (1789)dated to Schrank and interpreted as
being at varietal rank (see below for a comment on
the asterisk). This name, which is cited as a synonym
by, 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 the
third alternative;
3.Triticum dicoccum (Schrank) Schübl., Diss. char.
descr. cereal. 29 (Schübler 1818)or with the authors
as Schrank ex Schübl. (1818)or just Schübl.
(1818). This alternative uses the Latinised version
of 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 Schranksnameis
considered to have been published validly and at a
rank other than species. Schrank ex Schübl.(as
with Morrison 2007: 272) or just Schübl.should be
used when Schranks name is considered invalid.
Schranks1789 Bayerische Flora and Schüblers1818
Dissertatio are the critical publications in addressing
the dicoccon/-um issue.
Schrank (1789: 388) described Triticum spelta as no.
263 on p. 388 in his ora, adding *dicocconon p. 389
(Fig. 1), followed by a diagnosis in German not in
Latin as Mansfeld (1958: 237) suggests and
indicating that dicoccon is not a particular form of
Einkorn wheat (‘…Es ißt das Emmerkorn gewiß keine
Spielart des Einkorns…’), but, if anything, something
closer to spelta, as he writes at the end of that species:
‘…Hierher, denke ich, gehört das Emmerkorn…’ [‘…
here, I believe, belongs the Emmer wheat…’]. His
dicoccon thus appears to be part of spelta and, in
addition, does not receive its own number but an
asterisk (*) instead. The use of the asterisk before the
epithet indicated that it had at the time not yet been
found in the region of his ora, i.e. Bavaria (Mansfeld
1958:237)indeed, Schrank refers to its cultivation in
the nearby region of Württemberg. In contrast to this,
however, Schrank does not use for his dicoccon aGreek
letter (α,β, etc.) as he does for his infraspecic taxa
(for example with the subdivision into αaestivum and
βhybernum of his T. cereale). Tzvelev, Kerguélen, Cai et al.,
and van Slageren are therefore erroneous in their
interpretation as an intended variety. That is not to say
that Schrank sees Emmer as a species, however. He
not only indicates its closeness with spelta,butthen
writes: ‘…wenn es nicht eine eigene Art ißt …’ [when
it is not a separate species] (Fig. 1). In other words,
did he actually consider it less distinct than spelta
itself, therefore not deserving species status, and thus
implicitly as belonging under it? Then status as a
variety under spelta is logical, and this was the earlier
interpretation as spelta var. dicocconby the quoted
authors. Doubt about separate species status is also
expressed by Humphries (1978: 368); however, he
then immediately states that ‘…although Schrank may
have had some doubt about its status he still went
ahead and published it as a species.
481CONCEPTS AND NOMENCLATURE OF THE FARRO WHEATS
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2013
We present these three alternatives for the same name
and their respective arguments in order to show that
confusion has dogged the Schrank name almost to the
present day. However, after his remarks on the status of
Emmerkorn Schrank writes ‘…; ich nenne es einsweilen …’
[‘…; I call it for the time being…’]. The einsweilen
makes Triticum dicoccon Schrank provisional and thus
invalid for Art. 36.1(b)(GreuterinMorrison1998: 708).
We agree, and this causes the name T. dicoccum (Schrank
ex) Schübl. from 1818 to come into view.
Schüblers1818 publication describes Triticum
dicoccum (thus with a Latin ending of the epithet) while
referring indirectly to the earlier Schrank name. He
refers in a footnote (loc. cit. 30 31) to Roemer &
Schultes (1817: 766), commenting that the latter ‘…cite
T. dicoccon Schrank as a synonym of T. zea Host while
there are many differences between the two, such as the
glumes [of T. zea] only partly covering the spikelets with
mature seeds and [the spike] rachis exible…’ (both
good characters of T. spelta where HostsT. zea actually
belongs). Thus a link with Schrank exists, albeit indirect.
Shortly thereafter Schübler (1820: 451) re-described and
illustrated dicoccum in great detail, listing dicoccon
Schrank but not T. zea as a synonym, thus
considering the earlier dicoccon the same species but
did not accept Schranks name as the validone.
According to the ICN Art. 46.5 the name should be
ascribed to the validating author and his/her spelling,
but reference to the earlier, invalid publication at the
same time can be made: Triticum dicoccum Schrank ex
Schübl.or just Schübl.. To recognise Schranks role in
all this we recommend the rst form of author citation; it
Fig. 1. Triticum dicoccon as described by Schrank in his Bayerische Flora, Vol. 1.
482 KEW BULLETIN VOL. 68(3)
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can be based on the 1818 Schübler publication, but if
one considers this not a strong enough reference, then
certainly on his 1820 one.
Hammer et al. (2011: 5) are prepared to put the ICBN
rules aside for the sake of stability in order to maintain that
Triticum dicoccon Schrank 1789 was validly published. They
refer to Art. 34.1(b) of the ICBN (McNeill et al. 2006)
now Art. 36.1(b)oftheICNruling provisional names
invalid, but declare this to lead to nomenclatural un-
certaintiesthat 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 Schranks name is not acceptable, T. farrum
Bayle-Bar. from 1809 should be adopted as the species
name. No explanation is provided, however, as to why this
would be so. Even if they were right there are options
between 1789 and 1809 that should have been considered
as well. Most of these have appeared in synonymy, but not
in treatments by genetic resources scientists such as
Mansfeld, Hammer and others, but by botanists such as
Kerguélen (1975:279280) and Humphries (1978:368).
Names published before Schübler 1818 relating
to Emmer
Species names of taxonomic synonyms of Triticum
dicoccon Schrank, published between 1789 and Nov.
1818, the date of Schüblers valid publication as T.
dicoccum Schrank ex Schübl.:
1.Triticum album Gaertn., Fruct. Sem. Pl. 2(1): 8, Tab.
81, g. 1 album ai(Gaertner 1790) (Fig. 2).
(Our notation relating to the illustration summa-
rises the letters connected with the analytical
drawings of T. album.) This species is not mentioned
by Humphries in 1978, while cited by Kerguélen
(1975: 278) under T. aestivum as T. album Gaertner
ex Steudel, Nomencl. Bot. ed. 1: 853 (1821), pro
syn.. Kerguélen apparently relied on Steudels
Nomenclator, both for the name and its authorship,
but examination of all original elements presents an
ambiguous identity. This problem could be solved if
a type specimen were known but the herbarium
TUB that houses Gaertners types does not hold any
such collection. Gaertnersalbum illustration depicts
adicoccum as here understood, including the prom-
inent glume keel running from apex to base, which
is a turgidum-wheat character (Fig. 2). The phrase
descriptions on p. 8 present a mixed identity: Bauhin
(1623: 21), Morison (1699: 175), and Linnaeus
(1774: 108) all refer to either awned aestivum or
awnless hybernum forms of bread wheat. Malpighi
(1687: 89, Tab. 54, g. 325 A G) only shows some
viewsofawheatgrainandisnotconclusive.De
Tournefort (1719, 1: 512, and its reference to Vol. 3)
Tab. 293, Fig. F is different. Tabula 293 shows two
spikes, of which Fig. Q is T. dicoccum and Fig. T is
T. turgidum (Fig. 3); the referred Fig. F depicts a
wheat grain in Tab. 292. Thus, Gaertnersown
illustration and (part of) de TournefortsTab.
293 present links with Emmer. Kerguélen did not
lectotypify album when he placed it under bread
wheat, and were it not for the two illustrations it
may well belong there. Lectotypication may be
desirable for stability of use but can never do
justice to all elements associated with the name.
Thismakesitunsuitableastheoldestcorrect
name for dicoccum at species level.
2. Triticum spelta sensu Host, Icon. Descr. Gram. Austriac.
3: 21, Tab. 30 (1805), non Linnaeus (1753). While
there is direct reference to LinnaeussSpecies Plantarum
(1762:127)theillustrationofTab.30clearlydepictsa
dicoccum as here understood; in addition no collection
in the Host herbarium at W exists in connection with
this name. Mansfeld (1958) considers the Host species
a synonym of his T. dicoccon Schrank. While older than
the following two it is effectively a misapplication of the
Linnaean name and not a contender.
3. Triticum atratum Host, Icon. Descr. Gram. Austriac. 4:
5, Tab. 8 (1809). We were informed that as with
T. spelta sensu Host no specimen for this name is
known in the Host herbarium in W, and have
assigned a type in the form of Tab. 8 (Fig. 4). We
could not establish the date of publication with
more precision than the year 1809 (as in Staeu
&Cowan1979:340);thisisthesameyearasthe
next entry, T. farrum.
4. Triticum farrum Bayle-Bar., Monogr. Agron. Cereal. 1:
50, Tab. 4, gs 1 2 (Bayle-Barelle 1809)(Fig.5). As
for atratum we have selected an illustration as the type
(see below) as there is no herbarium known to exist
the can be linked to Giuseppe Bayle-Barelle
(Staeu & Mennega 1992:398;conrmed in
correspondence with PAV, the most likely place
for any such collection).
5. Triticum bauhini Lag., T. cienfuegos Lag. and
T. gaertnerianum Lag. from LagascasElenchus
Plantarum 6(1816) will not be further considered
as T. farrum is older, validly published and taxo-
nomically Emmer wheat. These (and many other)
species are now heterotypic synonyms under the
accepted 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 has
also been presented in addition to dicoccum,forexample
by Schübler (1820,nexttodicoccon Schrank) and
Philippe de Vilmorin (1905).AtypespecimeninG
has been selected (see below).
We agree with Kerguélen (1975: 280) that a choice
cannot, in fact, be made between Triticum atratum and
T. farrum, and there is no reason why farrum should
have preference until an earlier publication date in
1809 can be proven for it. It is only in following the
483CONCEPTS AND NOMENCLATURE OF THE FARRO WHEATS
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2013
established traditionof Mansfeld and Hammer,
discussed above, that we recommend T. farrum as the
accepted name at species level for Emmer wheat.
Conclusion: considerations for the name of Emmer
wheat at subspecic level
As to what Emmer wheat represents we agree with
Philippe de Vilmorins(1905:328)Hortus Vilmorinianus:
the wheat groups are what he calls Variétés agricoles
[agricultural varieties], united under a botanical name,
whetherthisisatspecies(asvulgare Vill.where T.
aestivum would be correct), subspecies (for example
vulgare Vill. subspec. T. durum Desf.’—just subspec.
durumwould be correct) or varietal (var. dicoccum
Schrank) rank. Interestingly, de Vilmorin does not add
the notion Variétés agric.after his var. dicoccum,but
we consider this unintentional. His location of dicoccum
as a variety under cultivated monococcum is not argued
but illustrates their supposedly close link. With spelta a
subspecies under T. vulgare the practical grouping of
the hulled Farro species vs the free-threshing ones is
not adopted.
Building on this concept and taking note of
Thellungs and Mac Keys considerations for an overall
wheat classication, Emmer (and the other Farro
wheats) can be located with precision within the
genus, now dened by the Gene Pool concept and
subdivided along ploidy levels and genome types. The
oldest available name for Emmer at subspecic level is
then Triticum turgidum L. subsp. dicoccum (Schrank ex
Schübl.) Thell. (see below).
Annotated summary of nomenclature (accepted
names at both species and subspecies rank
in bold; only homotypic synonyms and selected
heterotypic synonyms included)
EINKORN (2x = 2n = 14)
Triticum monococcum L. (Linnaeus 1753: 86); Metzger
(1824:3536); (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. Gaertners(1790)De Fructibus et Seminibus Plantarum Volumen Alterum, Tab. 81, g. 1 showing Triticum album with
details ai. (Adapted from the original Gaertner plate.).
484 KEW BULLETIN VOL. 68(3)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2013
Fig. 3. De Tourneforts(1719)Institutiones Rei Herbariæ, Tomus Tertius, with Tab. 293, g. Q, showing a spike of Triticum
dicoccum.
485CONCEPTS AND NOMENCLATURE OF THE FARRO WHEATS
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2013
Fig. 4. The illustration with analysis of Tab. 8 in Hosts(1809)Icones et Descriptiones Graminum Austriacorum, Vol. 4, selected as
the lectotype of Triticum atratum.
486 KEW BULLETIN VOL. 68(3)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2013
Percival (1921: 154, 170); Flaksberger (1915:21,1935:
355); Kerguélen (1975: 279); Hammer et al. (2011:4);
Gontcharov (2011: 9). Type: [not located] (lectotype
LINN 104.4!, selected by Bowden (1959: 664)).
Fig. 5. The illustrations with analysis of Tav. IV [Tab. 4], gs. 1 2, in Bayle-Barelles(1809)Monograa Agronomica dei Cereali,
selected as the lectotype of Triticum farrum.IMAGE © BRITISH LIBRARY BOARD, C.21.C44 PG 118. REPRODUCED WITH PERMISSION.
487CONCEPTS AND NOMENCLATURE OF THE FARRO WHEATS
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2013
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 actual
name was not validly published until the year
after.
Triticum vulgare Vill. convar. [Var.-Gr.]monococcum
(L.) Alef. (Alefeld 1866: 333). Type: as for T.
monococcum.
Triticum monococcum L. var. vulgare Körn. (Körnicke
1885: 111, 112); Percival (1921: 175). Type: as for
T. monococcum. Note: epithet and thus name is not
invalid for Art. 24.3, although indicating the type of
the 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 superuous nal epithet
(monococcum should have been used). The rank
appears to be considered a variety by Thellung
(1918a:470,subsp. II. cereale (Ascherson et
Graebner pro var)) who then raises it to subspe-
cies. However, this seems to be a contradiction to
his interpretation elsewhere of A., B., I., II., etc.
from Ascherson & GraebnersSynopsis as subspe-
cies. For example, Thellung (1912: 143) earlier
considered T. ovatum (L.) Rasp. B. I.triaristatum
(Willd.)Asch.&Graebn.asubspecies;van
Slageren (1994: 325) in a wider discussion on the
rankings in the Synopsis agreed. Mansfeld (1958:
237) considers T. sativum Lam. B. II.dicoccum
(Schrank ex Schübl.) Asch. & Graebn. to be a
subspecies 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 indicate
cultivated 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);vanSlageren(1994:85);
Morrison (2007: 272); Hammer et al. (2011: 4). Type:
(Russian Federation, Daghestan autonomous region)
[a free-threshing mutant,] isolated at the Derbent
branch of the N. I. Vavilov Institute from a collection
of cultivated T. monococcum made by P. M. Zhukovsky
in 1926 in Dadaj, Kastamonu region, Turkey, [and
preserved as a herbarium voucher on] 29 June 1970
[by] U. Kurkiev & A. Filatenko (holotype WIR K-48993;
isotypes WIR-Derbent no. 20970, [former WIR branch
at] 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 of
rank). Note: Szabó & Hammer (1996: 6) expressly
consider provar. and convar. of separate rank in
their classication. Hence their new combination is
published at two ranks at the same time and
therefore 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 for
T. sinskajae. Note: this name was cited earlier by
Szabó & 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 ex
Schübl.) Thell. (Thellung 1918a: 470, 1918b: 146); Mac
Key (1966: 268, 2005: 41); Kerguélen (1975: 279); van
Slageren (1994: 91). Type: the newly selected type is
chosen for the basionym, T. dicoccum Schrank ex
Schübl. (q.v.)
Triticum dicoccon Schrank (1789: 389); Schübler & von
Martens (1834: 46, dicoccum); Harz (1885: 1217,
dicoccum); Flaksberger (1915:22,dicoccum
Schrnk.); 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, g. 2acwith Trit. dicoccon
Schrankas 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 aestivum
spicis incanis 1768and hoc nominum missum est
Triticum Dicoccum Sch.(lectotype M-0187352!,
selected here (Fig. 6)). Notes: (1) Of the label
annotations, the line Triticum aestivum spicis
incanisand the year 1768were written by
Schrank, while hoc nomine missum est Triticum
dicoccum Sch.was not written at the same time, is
in a different ink, and most probably not by
Schrank. This makes sense as Schrank would not
have written missum est[was sent]. These words
can be compared with those on the separate
Schreber label on the sheet it could well be
the same hand (Schrebers?, which would make
sense). It seems quite possible that Schrank sent
the specimen to Schreber in 1768, under the
488 KEW BULLETIN VOL. 68(3)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2013
name Triticum aestivum spicis incanis, and that
Schreber later added This is Triticum dicoccum
Sch[rank]..Itthusappearsthattheepithetdicoccum
was coined already some time before 1789 but with
the Latin um ending, not the Greek on that was used
with the formal publication. (2) Earlier than this
lectotypication, Bor (1970: 207) and Kerguélen
(1975:279)notedattype:Die von Stuttgart
gesandten Samen gägen nicht nur im kalten
Boden, sondern sogar im Blumentöpfe
vierblütige zweisaamige Aehrchen,Schrank..
This is a major part of the comment by
Schrank at his *dicoccon publication in his Flora
(Fig. 1), only omitting H. R. Kerner,theperson
in Stuttgart who sent him the seeds. Of course it
is not a typication as such and its use of
German is also incorrect in parts. (3) Cited
literature attributes the species to Schübler 1818;
however, Flaksberger, Tzvelev, Kerguélen, and
Gontcharov cite the author as (Schrank)
Schübl., while Morrison is one of the few
instances of a correct author citation for this
species. Note also that Flaksberger changes
position compared to his 1915 paper (see at
dicoccon Schrank). [The identication of the type
specimen, the interpretation of its annotations,
and comments on the Schrank statement were
kindly 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 ex
Schübl.) Körn. (Körnicke 1885: 41, 81, as (subsp.)
Triticum dicoccum Schrk.,tobecorrectedforArt.
24.4); Mansfeld (1958: 237, cited as a synonym of T.
dicoccon Schrank). Type: as for T. dicoccum.Note:
Körnicke accepted only three species in Triticum (tom.
cit. 40) and dicoccum and spelta (and others) are
considered Gruppen (Unterarten)[groups (subspe-
cies)] of T. vulgare. Still the typography suggests that
he sees them at species level and the purported rank
is not further mentioned. As all of these groupshave
few or many Varietäten[varieties], both interpreta-
tions of his classication would not be in conict with
the accepted taxonomic hierarchy.
Sitopyrum dicoccum Döll, nom. invalid. (Art. 36.1(c)). As
quoted by Harz (1885: 1217) with the author of the
name only as Döllrather than (Schrank ex
Schübl.) Döll. Note: a name cited only in synonymy
of T. dicoccum Schrank that appears to refer to
Dölls(1857 (1855): 123) unranked subgroup I.
Pyrosof Triticum. The generic name is not found
elsewhere.
Triticum sativum Lam. subsp. dicoccum (Schrank ex
Schübl.) 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 of
Triticum dicoccum in herbarium M. IMAGE © BOTANISCHE
STAATSSAMMLUNG MÜNCHEN. REPRODUCED WITH PERMISSION.
489CONCEPTS AND NOMENCLATURE OF THE FARRO WHEATS
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2013
subsp. cereale); Mansfeld (1958:237,ascribesthe
name to Ascherson & Graebner 1901). Type: as
for T. dicoccum.
Triticum monococcum L. var. dicoccum (Schrank ex
Schübl.) P. Vilm. (Ph. de Vilmorin 1905: 328).
Type: as for T. dicoccum.
Triticum aestivum L. var. (θ)dicoccum (Schrank ex
Schübl.) Fiori (1896: 108). Type: as for T. dicoccum.
Triticum spelta L. subsp. [sous-esp. T. dicoccum’—to
be corrected for Art. 24.4] dicoccum (Schrank ex
Schübl.) Husn. (Husnot (1899:81).Type:asforT.
dicoccum.
Triticum aestivum L. subsp. dicoccum (Schrank ex
Schübl.) Thell. (Thellung: 1912: 141). Type: as for
T. 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, g.
1album 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 with
analysis of Tab. 8 in Hosts1809 Icones et Descriptiones
Graminum Austriacorum Vol. 4, selected here
(Fig. 4).
Triticum dicoccum Schrank ex Schübl. var. (β)atratum
(Host) Schübl. & G. Martens (Schübler & von
Martens 1834: 46, as βT. atratumto be corrected
for Art. 24.4); Bluff et al. (1836: 203); Schrader in
von 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 atratum
under monococcum after investigating material from
Metzger in which he only found one seed per
spikelet.
Triticum farrum Bayle-Bar. (Bayle-Barelle 1809: 50,
Tav. 4, gs 1 2); Harz (1885: 1219, Farrumand as
an invalid speciesunder T. dicoccum). Type: the
illustrations in Bayle Barelles1809 Monograa
agronomica dei Cereali, Tav. 4, gs 1 2, selected
here (Fig. 5). Note: the gures show the awnless
(g. 1) and awned (g. 2) forms of T. farrum
spikes, together with individual spikelets and
glumes. Both gures are expressly included to
show the variety of forms that make up this
species as was intended by Bayle-Barelle. The
awnless form of Tav. 4, g. 1 is named T. farrum
muticumbut this is not formally described on
pp. 50 52 of the book as aseparate entity. On the
contrary, Triticum farrum (mihi)is headed on p. 50 by
Specie IX. Tav. 4. Fig. 1. e 2 [our italics]., expressly
referring to both gures.
Triticum vulgare Vill. (convar. dicoccum (Schrank ex
Schübl.) Alef.) var. farrum (Bayle-Bar.) Alef.
(Alefeld 1866: 331). Type: as for T. farrum.
Triticum dicoccum SchrankexSchübl.var.farrum
(Bayle-Bar.) Körn. (Körnicke 1885: 84, 87);
Flaksberger (1915: 64, 184); Percival (1921: 197).
Type: as for T. farrum.
Triticum dicoccum SchrankexSchübl.(subsp.
europaeum Vavilov proles tardo-europaeum Flaksb.)
grex farrum (Bayle-Bar.) Flaksb. (Flaksberger 1935:
306). Type: as for T. farrum. Note: the grex contains
two varieties (var.) and two formae (f.), but none
is 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. de
Cours.); Schübler (1820: 451, author as Dumeril de
Cours[sic]); Harz (1885: 1217, as T. monococcum
majus Dumeril[sic]). Type: not indicated, and
herbarium unknown (Staeu & Cowan 1976). Note:
Dumont de Courset writes at his entry for monococcum:
Grande épeautre, T. monococcum majus.Although
the French name is generally linked with T. spelta,
when seen as a series of increasing size the next step
upwould logically refer to dicoccum where the majus
name has indeed been located.
Triticum amyleum Ser. (Seringe 1818: 124); Metzger
(1824:3035); H. de Vilmorin (1880: 13, 21).
Type: a specimen from the herbarium of Charles-
Isaac Fauconnet, acquired by Delessert for the
herbarium in Geneva in 1879, bearing the printed
label: 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 to
adequate illustrations by the Bauhin brothers are
presented by Seringe, plant material has preference
(sensu Art. 9.12 of the Code, the lectotype being an
uncited specimenas referred to in the Article),
and the lectotype from G serves as such.
Spelta amylea (Ser.) Ser. (Seringe 1842: 114). Type: as
for T. amyleum. Note: although the genus name is
illegitimate (for Art. 53.1) after Spelta Wolf the
species 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 of
Plate 1, g. 3a cinFlora 3(2) of 1820, referred to
in Schüblers paper on p. 458, selected here. Note:
Dr Hajo Esser (M) informed us that no material was
found in M that could be connected with Schübler
with enough certainty to take precedence. The
illustration of a complete spike with spikelet and
grain details is perfectly adequate to show the
490 KEW BULLETIN VOL. 68(3)
© The Board of Trustees of the Royal Botanic Gardens, Kew, 2013
purported difference of three grains per spikelet
compared with the 2-grained normaldicoccum spike-
let and the 1-grained monococcum one.
Triticum dicoccon (Schrank) Schübl. var. (γ)tricoccum
(Schübl.) Schübl. & G. Martens (Schübler &
Martens 1834: 46, as γT. tricoccumto be corrected
for Art. 24.4); Schrader in von Schlechtendahl
(1838: 465); Döll (1857 (1855): 125). Type: as for
T. tricoccum.
Triticum vulgare Vill. (convar. [Var.-Gr.]dicoccon
(Schrank ex Schübl.) Alef.) var. tricoccum (Schübl.)
Alef. (Alefeld 1866:332,subtricoccumbut with
direct 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 (1824:2630, Spelta); Schübler & von
Martens (1834: 46, Speltaand with new varieties for
awned (αaristatum)andunawned(βmuticum)
forms); Harz (1885: 1210, Speltaand including
numerous 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:asforT. spelta.
Note: although the genus name is illegitimate (for
Art. 53.1) after Spelta Wolf 1776 the species name is
not (Art. 55.1). Seringe called this species Epautre
commun, clearly suggesting it is a renaming of
Triticum spelta and not of another species such as
T. 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. (Körnicke
1885: 41, 75, as (subsp.) Triticum Spelta L.,tobe
corrected 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 for
T. spelta.
Triticum vulgare Vill. subsp. spelta (L.) P. Vilm. (Ph. de
Vilmorin 1905: 328, as subspec. T. spelta L.to be
corrected 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:asforT. spelta.
Triticum x aestivum L. emend. Bowden cultivar group
spelta (L.) Bowden (1959: 674). Type: as for T. spelta.
Triticum aestivum L. varietal groupspelta (L.) Morris &
Sears (1967: 21) pro parte (considered to be based
on T. spelta L. + T. macha Dek. & Men.). Type: as
for 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 illustration
with analysis of Tab. 29 from HostsIcones et
Descriptiones Graminum Austriacorum Vol. 3, selected
here. Note: the Tab. 29 shows a clear but awned
spelta, which is less common than awnless cultivars.
Acknowledgements
At K we thank Melanie Thomas for advice and transla-
tion of Schüblers(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-
cationof 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 nancially
supported by the Stichting Elise Mathilde Fonds,
CIMMYT, Int., and the COFRA foundation.
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... Leaves alternating, simple, whole, linear, 15-40 Â 1-2 cm. Spike distichous spike 4-12 cm long, spikelets 10-15 mm long, laterally compressed, 4-7-flowered (Maniee et al. 2009;Pecetti and Damania 1996;Van Slageren and Payne 2013;Zeven 1990). Archaeological have yielded 8000BC-dated remains of primitive kinds of farmed Triticum turgidum L. (Abinasa et al. 2011;Chaachouay et al. 2020Chaachouay et al. , 2021Elhani et al. 2007;Levy and Feldman 1988;Ozbek et al. 2007a;b;Pecetti and Damania 1996). ...
... For this reason, Bowden (1959) suggested to unite the two genera into one genus, Triticum. Yet, despite their close genetic relationship, most taxonomists (e.g., Zhukovsky 1928;Eig 1929;Mac Key 1966, 1981Hammer 1980;van Slageren 1994) prefer, based on morphological differences, to regard them as two separate genera. Classification of the genera Aegilops and Triticum is complicated due to: (1) allopolyploid species, which have bi-phyletic or tri-phyletic origin, resulting from intergeneric hybridization between species from two different genera; ...
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This chapter reviews dilemmas concerning the classification of the three genera of the wheat group. It discusses the differences between the taxonomic species concept and the biological species concept, and refers to the debated issue of designation and classification of synthetic allopolyploids.
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The chemical, physical and physicochemical characteristics of Einkorn wheat (Triticum monococcum), a representative of diploid wheat and Emmer wheat (Triticum dicoccum), a representative of tetraploid wheat have been studied. Based on the obtained results, technological characteristics of the milling and baking quality of the studied cereals have been made, with respect to their future application in the production of wholemeal flours for food purposes. The specific features of the studied grains and their technological potential for obtaining wholemeal flours with different product composition and properties are taken into account.
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Intraspecif ic classif ication of cultivated plants is necessary for the conservation of biological diversity, study of their origin and their phylogeny. The modern cultivated wheat species originated from three wild diploid ancestors as a result of several rounds of genome doubling and are represented by di-, tetra- and hexaploid species. The identif ication of wheat ploidy level is one of the main stages of their taxonomy. Such classif ication is possible based on visual analysis of the wheat spike traits. The aim of this study is to investigate the morphological characteristics of spikes for hexa- and tetraploid wheat species based on the method of high-performance phenotyping. Phenotyping of the quantitative characteristics of the spike of 17 wheat species (595 plants, 3348 images), including eight tetraploids (Triticum aethiopicum, T. dicoccoides, T. dicoccum, T. durum, T. militinae, T. polonicum, T. timopheevii, and T. turgidum) and nine hexaploids (T. compactum, T. aestivum, i:ANK-23 (near-isogenic line of T. aestivum cv. Novosibirskaya 67), T. antiquorum, T. spelta (including cv. Rother Sommer Kolben), T. petropavlovskyi, T. yunnanense, T. macha, T. sphaerococcum, and T. vavilovii), was performed. Wheat spike morphology was described on the basis of nine quantitative traits including shape, size and awns area of the spike. The traits were obtained as a result of image analysis using the WERecognizer program. A cluster analysis of plants according to the characteristics of the spike shape and comparison of their distributions in tetraploid and hexaploid species showed a higher variability of traits in hexaploid species compared to tetraploid ones. At the same time, the species themselves form two clusters in the visual characteristics of the spike. One type is predominantly hexaploid species (with the exception of one tetraploid, T. dicoccoides). The other group includes tetraploid ones (with the exception of three hexaploid ones, T. compactum, T. antiquorum, T. sphaerococcum, and i:ANK-23). Thus, it has been shown that the morphological characteristics of spikes for hexaploid and tetraploid wheat species, obtained on the basis of computer analysis of images, include differences, which are further used to develop methods for plant classif ications by ploidy level and their species in an automatic mode.
Chapter
Wheat, which originated around 12,000 years ago in the Fertile Crescent, is one of the most important crops worldwide providing around 20% of the total calories and protein of the human diet. During its long history across the whole world, several species and thousands of landraces have evolved making wheat the crop with more and diverse genetic resources available currently. The conservation, evaluation and characterization of these genetic resources are basic in wheat sustainable cultivation because this genetic diversity could be key to face the challenges of agriculture nowadays. In this chapter, we describe an important part of these wheat genetic resources, the so-called ancient wheat. These are materials that were used by ancient civilizations but that know have a marginal use in agriculture and that in most of the cases are conserved in Germplasm Banks. Ancient wheat host huge variability that could be used in breeding programs to enhance the resistance to biotic and abiotic stresses, and the grain quality properties of the wheat grain, including its nutritional properties.
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Мета. Проаналізувати практику найменувань основних видів і підвидів пшениці. Результати. Сучасні системи роду Triticum базуються на різних морфологічних та генетичних критеріях, і нараховують від 7(10) до 27 видів. В Україні культивують види (підвиди): пшениця літня (T. aestivum), пшениця кулястозерна (T. aestivum subsp. sphaerococcum; T. sphaerococcum), пшениця щільна (T. aestivum subsp. compactum; T. compactum), пшениця спельта (T. aestivum subsp. spelta; T. spelta), пшениця однозерна (T. monococcum L.), пшениця тверда (T. durum; T. turgidum subsp. durum), пшениця картлійська (T. carthlicum; T. turgidum subsp. carthlicum, T. persicum), пшениця польська (T. polonicum, T. turgidum subsp. polonicum) та пшениця двозернянка (T. turgidum subsp. dicoccon) або пшениця фаро (T. farrum, T. dicoccon, T. dicoccum). Поряд із системою ботанічних назв існує агробіологічна, що базується на назвах культур, але є менш упорядкованою. Через це один і той же таксон може мати понад десяток назв, що застосовують різні автори в агрономії. Водночас, правильний переклад видової назви T. aestivum – пшениця літня трапляється лише в одному джерелі з чотирнадцяти проаналізованих. Переклад латинських триноміналів підвидового рівня українськими біноміналами згладжує розбіжності між морфологічними і генетичними системами роду Triticum. У науковому стилі української мови домінує прямий порядок слів за якого прикметник передує іменнику, за виключенням наукових видових назв рослин. Це дозволяє легко розрізняти назви, що належать до різних терміносистем. «Тверда пшениця» є назвою культури в агробіологічній номенклатурі, а «пшениця тверда» це видова назва в ботанічній номенклатурі. Висновки. Унаслідок опрацювання етимології упорядковано українські наукові назви основних таксонів пшениці, які відповідають оригінальним латинським назвам. У неорганізованій агробіологічній системі назв сільськогосподарських рослин використовують суміш ботанічних назв таксонів та назв культур, які інколи тією чи іншою мірою збігаються. Назви культур та назви таксонів різняться між собою, відповідно, прямим та зворотнім порядком слів. Помилкове чи неусвідомлене заміщення одних назв іншими та нехтування правилами української мови призводить до небажаної плутанини. Її можна подолати дотриманням норм літературної української мови та Правил номенклатури, таксономії та культономії рослин.