The lost ancestor of the broad bean (Vicia faba L.) and the origin of plant cultivation in the Near East

Abstract

The broad bean (Vicia faba L.) was among the founder crops of the Near East; nevertheless, its wild close relatives remain unknown. Presumably, its missing wild progenitor had a small range within the Levant and was associated with restricted habitats, so that it was domesticated entirely as a species. Its habitats are supposed to have been situated along floodplain/slope borders (“transeluvial-accumulative barriers”) providing favorable edaphic conditions. These restricted natural habitats of the broad bean could be foci of early cultivation activities, thus becoming nascent fields. It is hypothesized that the broad bean, a conspicuous plant with large seeds and restricted habitats, could be the Near Eastern “primer crop”, which provoked the first emergence of the idea and practice of plant cultivation and “invention” of the field.

Full text

831
ВаВилоВский журнал генетики и селекции, 2014, том 18, № 4/1
THE LOST ANCESTOR OF THE BROAD BEAN (Vicia faba L.)
AND THE ORIGIN OF PLANT CULTIVATION IN THE NEAR EAST
© 2014 г. O.E. Kosterin
Institute of Cytology and Genetics SB RAS, Novosibirsk, Russia,
е-mail: kosterin@bionet.nsc.ru;
Novosibirsk State University, Novosibirsk, Russia
Поступила в редакцию 29 августа 2014 г. Принята к публикации 1 сентября 2014 г.
The broad bean (Vicia faba L.) was among the founder crops of the Near East; nevertheless, its wild close
relatives remain unknown. Presumably, its missing wild progenitor had a small range within the Levant
and was associated with restricted habitats, so that it was domesticated entirely as a species. Its habitats are
supposed to have been situated along floodplain/slope borders (“transeluvial-accumulative barriers”) provid-
ing favorable edaphic conditions. These restricted natural habitats of the broad bean could be foci of early
cultivation activities, thus becoming nascent fields. It is hypothesized that the broad bean, a conspicuous
plant with large seeds and restricted habitats, could be the Near Eastern “primer crop”, which provoked the
first emergence of the idea and practice of plant cultivation and “invention” of the field.
Key words: Vicia faba L., Near East, origin of plant cultivation, plant domestication, founder crops, primer
crop, slope/floodplain joint.
УДК 631.9+581.6
INTRODUCTION
The broad bean, Vicia faba L., is known as a
cultivated plant from the very onset of agriculture
(Hanelt, 1972; Zohary, Hopf, 2000) and is an im-
portant crop until the present. According to the most
recent revision, V. faba represents a monospecic
section in its genus, nearly warranting upgrade to
generic rank (Maxted, 1993). Until the present,
neither wild representatives of this species nor any
closely related species have been found (Ladizinsky,
1975; Maxted et al., 1991; Maxted, Kell, 2009). The
absence of a link to the extant wild ora made the
broad bean scarcely considered in reconstructions of
the origin of plant cultivation and domestication in
the Near East. However, the absence of a wild rela-
tive of some specic crop may be as such meaningful
and indicative, as discussed below.
BROAD BEAN AMONG FOUNDER CROPS
The broad bean has been found in quite a
number of the earliest archaeological sites in the
Levant, namely in ten sites referring to the Pre-
Pottery Neolithic B (PPNB, ca 8,700-6,000 BC),
the time of early plant cultivation: three in the
Jordan basin: Jericho II (Hopf, 1983), Yiftahel
(Kislev, 1985), “Ain Ghazal (Rollefson et al.,
1985); one at the Orontes: Tell el-Kerkh (Tanno,
Willcox, 2006a); four in the upper Euphrates basin: upper Euphrates basin:
Tell Abu Hureyra II (Hillman, 1975; de Moulins,
2000), Tell Halula, Dja’de (Willcox, 1996), Nevali
Çori (Pasternak, 1998), Cafer Höyük IX-XIII (de
Moulins, 1997) and one at the upper Tigris: Çayönü
(van Zeist, de Roller, 1992). The broad bean was
also reported for two sites in the Jordan valley refer-
ring to the earlier Pre Pottery Neolithic A (PPNA,
ca 9,800–8,700 BC), when nascent plant cultiva-
tion was probably arising, namely Jericho I (dating
range 9,150–8,350 BC) and, less surely, Iraq ed-
Dub (9,700–8,800 BC), but these records remain
dubious with respect to exact archaeological layer
and species identication (Colledge, 2001). There
is some uncertainty with respect to identication of
V. faba seeds, which in some cases could not be dis-
tinguished from those of Vicia narbonensis L. and
were identied as Vicia sp., so the actual number
of Neolithic sites where V. faba was found may

832 O.E. Kosterin
be greater (Kislev, 1985; Tanno, Willcox, 2006a).
The broad been seeds found in archaeological sites
provide no information if the plant was collected
in the wild or cultivated. It is noteworthy that in
Yiftahel (Kislev, 1985) and Tell el-Kerkh (Tanno,
Willcox, 2006a), broad been seeds were found in
very large quantities. V. faba was not included in
the set of the so-called “founder crops” of the Near
East (Zohary, Hopf, 2000; Weiss, Zohary, 2011).
Fuller et al. (2012) referred to it as one of the “lost
crops”, although it was the ancestor which is lost,
not the crop. Tanno and Willcox (2006a) and Abbo
et al. (2013. P. 816) reasonably proposed to include
broad bean into the founder crop set: “We see no
reason why the broad bean cannot be added to the
classical eight “founder” crops list (Zohary, Hopf,
2000), as advocated earlier by Tanno and Willcox
(2006b). We presume that Zohary and Hopf (2000,
and in earlier editions) refrained from doing so
simply because the wild ancestor of broad bean is
still elusive (Zohary, Hopf, 2000)”.
The area of the broad bean domestication is sup-
posed either very broadly as “between Afghanistan
and East Mediterranean” (Hanelt, 1972; Maxted,
Kell, 2009. P. 133) or as two options: “the Near East,
which is the centre of diversity of section Faba [in
a broad sense by Kupicha], and Afghanistan, where
the most primitive forms of V. faba occur” (Maxted
et al., 1991). The second option (Afghanistan),
proposed by Ladizinsky (1975) is, however, un-
likely since the archeological records of cultivated
V. faba in the Near East far predate the onset of
agriculture in Afghanistan (Cubero, 1984). The
diagnostic characters of V. faba subsp. paucijuga
Murat., cultivated in Afghanistan, Pakistan and N
India, namely, the greater number of leaet pairs
per leaf (3–4) and owers per inorescence (4–11),
were supposed to be plesiomorphic (Muratova,
1937; Maxted et al., 1991; Maxted, Kell, 2009) but
the arguments are not convincing. Moreover, the
very necessity of subspecies division of V. faba, is
not well justied (Cubero, 1973, 1984). The sub-
stantial diversity found in recent broad beans has
most probably been accumulated already under
cultivation, by mutation, isolation during expansion
of the crop, and selection, since little intraspecies
divergence was found in in the entire species (“a
strong nucleus, which carries the maximum of po-
tentialities of the species and from which different
populations branch”) (Cubero, 1973. P. 59).
AN ANCESTOR MISSING
Since Alphonse de Candolle (1882), the standard
logic for locating the area of plant domestication,
at least in the Near East, was the search for areas
where all or most of wild relatives of the founder
crops still exist until the present (Lev-Yadun et al.,
2000; Abbo et al., 2011a, 2012). This approach has
to leave aside a founder crop which no longer has
its extant wild representatives or close relatives.
One may, however, suppose that such a species was
domesticated entirely as is, rather than was “taken
for cultivation from nature”, so that its natural range
and habitats were small and entirely converted to
nascent agrocenoses.
The wild progenitor of the broad bean is sup-
posed to be either extinct (Hanelt, 1972; Schäfer,
1973; Abbo et al. 2013) or not yet found (De Wouw
et al., 2001; Abbo et al., 2013). However, perhaps
the simplest explanation could be that it had a
restricted natural range and habitat and so was
domesticated entirely. Its cautious version is found
in literature: “... material in the centre of origin was
bred extensively so obliterating the remains of the
original forms” (Maxted et al., 1991. P. 136).
A PUTATIVE HABITAT
What conjectures can we make about the lost
broad bean wild ancestor based on the present
day crop? This is a large and conspicuous annual
herb with stiff erect stems and broad foliage. It is
hygrophilous and demands heavy clayey humid
soils rich in humus and carbonates (Muratova,
1937). These traits suggest for the broad bean wild
progenitor an open habitat with fertile soil and a
good water supply. This kind of habitat is far from
widespread in the Near East. In spite of a consid-
erable variability of the seed size among present
day cultigens, and although the earliest cultivated
forms were rather small-seeded (Maxted, 1993),
the broad bean is still a champion for this param-
eter among legume crops of Near Eastern origin.
Thus there is little doubt that its wild ancestor was
among wild Near Eastern herbaceous legumes with
the largest seeds.
The large seeds may be advantageous in growth
competition of seedlings among annual plant spe-
cies; or an adaptation to drought after germination
as allowing early growth of a deep root. At last, it

833
e lost ancestor of the broad bean (Vicia faba L.) and the origin of plant cultivation in the Near East
may be supposed to serve for successful germina-
tion in conditions of relatively deep burial (note the
broad bean has a hypogeal type of germination).
Domestication of a crop was usually followed (or
accompanied) by gradual seed enlargement, which
was supposed to be an adaptation to deeper seed
burial under cultivation than achieved with natural
seed dispersal (Harlan et al., 1973; Smith, 2006;
Fuller, 2007), with germinating ability at deep
burial experimentally shown to correlate with seed
mass for ve of eight grain legume species tested
(Kluyver et al., 2013). Analogous situations of
deeper burial can be found in nature, for instance,
beneath occasional alluvium deposited by sudden
oods in river oodplains. In pre-agricultural times,
the oodplains of major rivers received seasonal
delivery of thick alluvium. They were swampy and
dominated by large perennial plants propagating
via rhizomata or stolons, with participation of tiny
“oodland ephemeretum” (annual plants with very
short life span specic to oodlands). However,
we may suppose that natural growth of the broad
bean progenitor occurred at the most elevated
levels of oodplains along their margins, where
they meet the valley slopes, thus locally rimming
the slope bases. In terms by the seminal work by
B.B. Polynov (1937) considering soil chemistry
in a geomorphological context, this position was
the border between the transeluvial position of a
hill/mountain slope and the accumulative position
of oodplain. This transitional landscape position,
once termed “transeluvial-accumulative barrier”
(Glazovskaya, 1964; Stebaev et al., 1993), is an
ecotone known for its very high biological produc-
tivity because of soils enjoying permanently suf-
cient amount of ground water (provided by springs
at slope bases as well as by the river), oxygen and
nutrients (Stebaev et al., 1993). It is irregularly (not
every year) reached by the highest oods and the
soil is expected to be irregularly and moderately
disturbed by slope creeps and linear erosion as well
as by occasional deposit of thin alluvium.
It is noteworthy that many species of the plant
communities of the slopes are usually found still
growing along the slope bases, partly because of
accumulation of seeds washed down or fallen from
above, so the slope bases are the habitats richest in
species across a valley. In the Near East they could
include wild progenitors of other annual founder
crops, with seeds smaller than those of the broad
bean. Moreover, there they could undergo some
natural selection for greater seed mass which in-
creases the chances of a seed descending the slope
by gravitation and ensures its germination from
under the thin alluvial deposit. Various species from
the slope plant communities could facultatively
grow at the slope margins, however the characters
of the contemporary broad bean tempts us to sup-
pose its wild progenitor to be specialised to this
very habitat. Judging from the archaeological evi-
dence from the Levant cited above, the wild broad
bean progenitor’s native range could comprise the
valley(s) of some of the upper Orontes, Euphrates
and Tigris Rivers or their major tributaries, but
unlikely spread over all these valleys.
It is not possible to indicate at a plant associa-
tion in which the faba bean wild progenitor once
participated; moreover, the below hypothesis on
its role in the origin of plant cultivation implies
that this association, as a nascent agrocoenosis,
disappeared in its natural state together with the
progenitor itself.
INVENTION OF THE FIELD
Origin of agriculture associated with domes-
tication of plants, together with domestication
of animals comprising the so-called Neolithic
Revolution, was followed by a dramatic rise in
human population density and hence was one of
the main pre-requisites of civilisation. No doubt
it took place several times in different continents,
independently and non-synchronously (Vavilov,
1951; Harlan, 1971; Zohary, 1999; Smith, 2006;
Abbo et al., 2010a; Fuller et al., 2012). Naturally,
these crucial events used to inspire a great interest
as to why and how they happened, giving rise to
hypotheses about factors, both natural and social,
which lead to the origin of plant cultivation and
domestication. Such attempts, however, often look
too deterministic as implying that domestication
must have taken place as soon as necessary condi-
tions appeared, such as useful plant species and
soils, and would better be reformulated in terms
of factors which made appearing of agriculture
possible rather than inevitable.
The seminal N.I. Vavilov’s concept of plant do-
mestication centres (Vavilov, 1951) faced the main
theoretical problem in explaining why agriculture
origin and plant domestication was localised in

834 O.E. Kosterin
space and time (Harlan, 1971). And indeed, some
Vavilov’s disciplees found it possible to expand the
centres to “regions” (Sinskaya, 1969) or “mega-
gene-centres”, the latter eventually occupying
nearly the entire land except for the extreme North
(Zhukovskiy, 1970). A pre-existing fortunate set of
especially useful plant species is a bad candidate
for a specic factor restricting the area of agricul-
ture origin: in most speculations of that kind (e.g.
Diamond, 1998), advantages of plant species which
have been domesticated used to be reasoned ad hoc
while advantages or disadvantages of edible spe-
cies which have not been domesticated are not (and
could hardly be) considered comparatively.
Currently a hot debate goes on concerning the
origin of plant cultivation and domestication in
the Near East. One party suggests that their ori-
gin was singular, rather fast (hundreds of years),
and took place in the so-called “core area” about
250 × 150 km situated at the sources of the Euphra-
tes and Tigris in NE Turkey (Dijarbakyr and Mardin
Vilayets); this was followed by cultural evolution
of domesticated crops improving their quality (Lev-
Yadun et al., 2000; Gopher et al., 2001; Abbo et
al., 2010a, 2011a, 2012, 2013). Adherents of the
contrary so-called protracted model of plant domes-
tication argue that both cultivation and domestica-
tion had multiple origins over the Fertile Crescent,
went on slowly and in parallel, with pre-domestica-
tion cultivation for 1–1.5 thousand years preceded
domestication (as a genetical phenomenon) which
was gradual, lasted for not less than 3,000 years and
was crowned by fully domesticated crops (Willcox,
2005; Tanno, Wilcox, 2006b; Weiss et al., 2006;
Fuller, 2007; Allaby et al., 2008; Brown et al.,
2009; Glémin, Battailon, 2009; Fuller et al., 2011,
2012; Asouti, Fuller, 2012). Both parties agree that
plant cultivation emerged as a conscious practice
but disagree whether selection for the domestica-
tion traits was conscious and exploited variation
pre-existing in wild population (the “Core Area
party”) or non-conscious and utilised spontaneous
mutations occurring in already cultivated crops (the
protracted domestication party).
While arguments of both parties about domes-
tication, essentially genetic changes of crops, are
strongly supported and diverse, those concerning
the origin of cultivation as an idea and practice are
not. This is not surprising since it would hardly
leave archaeological remnants (Willcox, 2007).
Thus, arguments in favour of pre-domestication
cultivation are largely based on nding seeds of cul-
tivation weeds admixed with those of non-domes-
ticated crops, or remnants of “pre-domestication
granaries” (Colledge, 1998; Willcox et al., 2008).
It may be argued that such evidence depends on the
methods of yield processing and may not reect the
origin and spread of cultivation as an idea. So the
issue is far from being solved whether the inven-
tion of cultivation in the Near East happened once
or many times over.
The very nature of the invention of cultivation
can still be put into question: was it
– “a cultural conscious and deliberate choice” (Abbo
et al., 2005. P. 495), “a fully conscious socio-
cultural move based on a well-educated choice
of specic food sources” (Abbo et al., 2010a. P.
325) and based on “deep cultural traditions of
plant tending” among hunter-gatherers (Fuller et
al., 2012. P. 642), as both parties believe;
– or a “cultural mutation”, a “meme” in Richard
Dawkins’s (1982) sense supported by “natural
selection of ideas” in Karl Popper’s (1978)
sense?
These options are rather aspects of our attempts
to speculate on motivations of the most primitive
farmer than mutually exclusive alternatives. But
there is a high danger of involving too much of
actualism while reconstructing motivations of
pre-historic people. All primitive societies are
characterised, if not to say overloaded, by numer-
ous and diverse traditions and rituals; it may be
said that rituals were the main mode of existence
of early humans. Some of them seem senseless
from the modern human’s point of view, while
some are obviously harmful biologically, such as
body modication, human sacrice etc. From an
evolutionary biologist’s perspective, traditions and
rituals can be viewed as the heredity of a society,
the diversity of which looks as if it resulted from
random “mutations”. Only societies with more or
less harmless traditions would survive and those
with useful traditions would propagate. (Some bio-
logically harmful traditions, e.g. body modication,
of course bring about social advantage to those who
execute them, but only within the context of these
traditions themselves, and a society is still viable
only if the biological harm is tolerable.) There is a
well-known example of a kind of useful traditions
found in most of the known societies including our

835
e lost ancestor of the broad bean (Vicia faba L.) and the origin of plant cultivation in the Near East
own: those preventing inbreeding and promoting
exogamy. It is noteworthy, however, that in primi-
tive societies, the actual harm of inbreeding is not
realised and is too weak to be noticed in practice,
and that traditions providing outbreeding are usu-
ally too overcomplicated for this purpose (Lévi-
Strauss, 1966). Such traditions most probably ap-
peared by chance and independently in some local
societies but were positively selected and enhanced
throughout the world by the benet of avoiding
inbreeding. Analogously, plant cultivation can be
imagined to originate from some ritual appearing
by chance and positively selected because of the
gain of food.
Some hypotheses refer to such usual evolution-
ary scenario as change of function and derive the
nascent agricultural technologies from otherwise
aimed human activities. For instance, primitive
agricultural habits could result as a by-product of
some complex ritual of a religious or magic nature.
Such rituals were indeed described as focused on
maize, considered by primitive farmers not just
food but also a magic plant (Anderson, 1952). Ini-
tial practices leading to plant cultivation could be
rituals associated with e.g. sacricing food, burial
imitations etc., probably unconscious in respect
of gain in food supply and perhaps quite odd from
our point of view. In fact it is fairly difcult to
reconstruct if the initial motivation of the most
primitive farmer was mostly trophic or spiritual,
or combining these components in a manner which
modern people would hardly understand. Anyway,
such speculations can by no means be tested.
Another example of the change-of-function
approach is the so-called “dump-heap hypothesis”
suggesting that soil disturbance around human
dwellings, its fertilisation by domestic wastes and
dropping of edible seeds would result in abundant
growth of useful annual plants and hence were pre-
requisites of plant cultivation (Engelbrecht, 1916;
Sauer, 1952; Anderson, 1952). However, it was
heavily criticised by Abbo et al. (2005) with respect
to the Near East cultivation. Note also that this hy-
pothesis does not solve the problem why plant do-
mestication was localised in space and time rather
than took place at early stages of human evolution
coherently throughout vast areas, as this hypothesis
predicts (Hawkes, 1983). Indeed, most of the land
was inhabited by humans and everywhere they used
to utilise some plants for food. Abbo et al. (2005)
additionally argued against the dump-heap hypoth-
esis noting that some of wild representatives of the
Near Eastern founder crops are not ruderals and can
hardly tolerate substantial human disturbance of
their natural ecosystems. In fact, as soon as a plant
species had been domesticated, the still wild and
already domesticated forms became subjected to
disruptive selection favouring them to retain either
wild or domesticated characteristics (Zohary, 2004;
Glémin, Battailon, 2009).
If practices of plant cultivation appeared by
chance as a cultural mutations, it is their low prob-
ability which could explain why centres of plant
cultivation origin, although quite a few worldwide,
were mostly localised in space and time in spite of
hunter-gatherers immense and intimate knowledge
of the surrounding nature and availability of some
useful plants wherever people lived. Still one can
consider objective pre-requisites favouring ap-
pearance of such cultural mutations (subjective in
essence) and/or providing their selective advantage
over non-farming societies, that is a common ap-
proach in the Darwinian theory of biological evo-
lution.
The amount of seasonal labour invested into plant
cultivation should have been associated with dra-
matic changes in societies of hunter-gatherers which
transited from utilising diverse but limited food re-
sources to cultivation of crops as a staple (Fuller et
al., 2010; Asouti, Fuller, 2012, but see Tzarfati et al.,
2013). So cultivation most probably could not appear
readily as soon as it became possible and benecial
in principle. Some additional factor is necessary to
facilitate invention of cultivation.
Hunter-gatherer societies depended on diver-
sied activities and used diverse food resources
distributed over considerable areas (Kelly, 1995;
Fuller et al., 2010; Asouti, Fuller, 2012). Perhaps
a crucial event in the transition to farming was the
“invention of the eld”, that is investment of large
amount of labour focused to certain restricted land
plots. Little is known about the earliest elds since
they used to leave no archaeological traces, so that
the archaeological record of nascent agriculture is
based on remnants of crop processing sites, dwell-
ings and burials (Fuller et al., 2010).
It is logical to suppose that the eld invention
would be facilitated if some very useful food plant
was conned to restricted habitats in the wild, es-
pecially if this plant was conspicuous and formed

836 O.E. Kosterin
pure growth. Let us call it the primer crop. Such
a situation would bring about natural focusing
of human attention to a special plant and of their
foraging activity to a restricted area. Addition of
some practice of soil treatment facilitating plant
growth either intentionally aimed at gain in food
or of a ritual nature, or most probably combining
both options, would result in converting these
habitats into elds. Hence a single primer crop is
supposed to trigger plant cultivation rather than a
set of founder crops.
Note this would to some extent solve the rst
problem of plant domestication: the wild type seed
dormancy, a trait considered critical for domes-
tication (Abbo et al., 2011b, 2012), which lead
Ladizinsky (1987) to his challenging concept of
“domestication before cultivation” in case of lentil:
sowing is efcient only with non-dormant seeds.
The inefciency (because of dormancy) of sowing
wild-type seeds without scarication was proved
by Abbo et al. (2011b) by experimental cultivation
of wild peas. If the earliest elds evolved from
natural habitats of a primer crop, harvesting seeds
could hardly be total, and the failure of sowing
dormant seeds would be compensated to some
extent by the soil seed bank. And if natural habitats
of the primer crops were scarce and restricted in
some large area, the focused human activity would
convert them all into nascent elds, thus solving
the second problem of plant domestication: gene
ow from the wild ancestor. Such a primer crop
would then undergo rather a rapid domestication,
that is genetic changes towards acquisition of the
so-called “domestication syndrome” (Hammer,
1984), characteristic for seed crops, in contrast to
slow domestication of those nascent crops which
co-existed with their wild relatives and hence
demonstrated the protracted pattern of acquisition
of the domesticated syndrome, because of gene
ow from the wild relatives (Jones, Brown, 2007;
Allaby, 2010) or just recurrent mixture with those
still being gathered from the wild (Barker, 2006;
Abbo et al., 2012) or entering elds through eld
shifting and fallowing (Fuller et al., 2010).
Fuller et al. (2012) argued than many independ-
ent cases of plant cultivation and domestication
were based on solitary crops, e.g. rice, Chinese mil-
let, pearl millet etc. Those solitarily domesticated
crops mostly were conspicuous plants conned to
specic habitats. A scenario with natural habitats
converted to elds could be suspected for the above
mentioned crops, and also maize. Note that the bot-
tleneck during maize domestication was estimated
to last 500–2000 years with the population size of
500–4000 maize individuals that suggests a fairly
small population of early farmers existing for about
a thousand years in isolation (Eyre-Walker et al.,
1998; Doebley, 2004). As supposed below, the
Near East plant cultivation may not have been an
exclusion and started from cultivation of a single
plant species serving as a primer crop”.
Whatever be the origin of the idea or tradition of
plant cultivation, it could spread over societies via
sociocultural inuences (Braidwood, 1967; Abbo
et al., 2010a) even decoupled from crops them-
selves, allowing recruitment elsewhere of suitable
species as crops from local wild oras (Jet, 1973).
Abbo et al. (2010a) noted that there is no proof of
independence of domestication of common bean
(Phaseolus vulgaris L.) and maize (Zea mays L.)
in Mesoamerica, which had taken place in an area
just several hundred kilometres away (Matsuoka et
al., 2002; Kwak et al., 2009), so that a sociocultural
inuence cannot be excluded. We may at least con-
sider as an option, and a version of the Core Area
hypothesis by Lev-Yadun et al. (2000), that in the
Near East, cultivation as an idea and practice had a
singular origin even if it was followed by multiple
domestication events. On the other hand, this sup-
position agrees with the protracted model of plant
domestication in assuming that the founder crops
may not have entered cultivation simultaneously.
BROAD BEAN AS A PRIMER CROP?
The European/West Asian civilisation sprouted
from the so-called “Neolithic revolution” following
the onset of plant cultivation in the Near East. This
centre of cultivation origin is characterised by quite
a number of founder crops: eight “traditional” ones
from three families: einkorn wheat (Triticum mono-
coccum L.), emmer wheat (T. dicoccum (Schrank)
Schuebl), barley (Hordeum vulgare L.), lentil (Lens
culinaris Medic), pea (Pisum sativum L.), chickpea
(Cicer arietinum L.), bitter vetch (Vicia ervilia (L.)
Willd.) and ax (Linum usitatissimum L.) (Lev-Ya-
dun et al., 2000; Zohary, Hopf, 2000; Abbo et al.,
2010a, b; Weiss, Zohary, 2011), plus broad bean
(V. faba) (Abbo et al., 2013), plus maybe some lost
crops (Melamed et al., 2008; Fuller et al., 2011,

837
e lost ancestor of the broad bean (Vicia faba L.) and the origin of plant cultivation in the Near East
2012 but see Abbo et al., 2013). It is noteworthy
that these crops are quite dissimilar in ecological
and biological respects demanding different ways
of treatment under cultivation: the cereals are
characterised by highly competitive determinate
growth, the legumes by indeterminate growth and
low competitive ability, with the pea being a tall
climbing plant, the broad bean a tall erect plant and
lentil and chickpea (Abbo et al., 2009, 2011b). For
this reason, the set of founder crops was supposed
to be complementary and deliberately chosen “on
the basis of intimate knowledge of their nutritional
value and potential to contribute to the nutritive
welfare of consumers” (Abbo et al., 2008. P. 928)
and because of “a very good yield buffering ability”
(Abbo et al., 2010b, 2012. P. 21).
However, such a variety of potential crop pro-
genitors, most of which grew over broad areas,
would hardly motivate transition from diverse
gathering activity to cultivation which demanded
extraordinary concentration of attention and labour
(Fuller et al., 2010). It may be supposed that among
those Near Eastern founder crops there was one
which served as the primer crop, to provoke an
idea and/or practice of plant cultivation and of the
eld as its focus. This could be the broad bean. It
is suggested above that the putative natural habitat
along the rims of river valleys was quite narrow and
probably patchy, and easy to convert into primary
elds. The relief position it probably occupied was
very favourable for cultivation, allowing a naturally
sufcient amount of ground water, nutrients and
oxygen. The broad bean is a tall conspicuous plant.
Its large seeds were not only useful for food but also
allow a clear observation of a plant germinating
from a seed and hence comprehension or stressing
the causal link between sowing and growing. (Note
that these are beans which are used for this very
purpose at present, as a model object` in school
education. Ironically, our hypothesis connotates
with the sacred nature of beans proclaimed by the
earliest philosopher of Pythagoras.) Absence of
extant wild relatives of the broad bean suggests
that the wild progenitor was not a common and
widespread plant. If so, all its scarce natural popula-
tions would be soon converted into primary elds
and then, without gene ow from wild relatives,
it would undergo rapid domestication, leaving no
remnants of the wild ancestor. Wild progenitors of
other founder crops were scattered over much larger
areas, as they still are at present, that would hardly
favour the emergence of the idea of cultivation at a
restricted eld. At the same time, presence of other
potential crops in the habitat of the broad bean wild
progenitor, being converted into primary elds, as
discussed above, would lead to their involvement
into cultivation as well. Moreover, conversion of
the broad bean natural habitats into primary elds
would most likely result in mixed species cultivation.
Probably this was an initial type of plant cultivation
in the Near East where the founder crop set included
so many species altogether. In this respect, the set
of founder crops could be balanced rather ecologi-
cally than nutritionally (as supposed by Abbo et al.,
2008, 2010b). If so, their involvement into cultiva-
tion could be interpreted in terms of endogenous
ecological succession of a biogeocenose which
included humans and plants being domesticated
as its constituents: human culture and genomes
of several plant species coevolved bringing about
drastic changes in the biogeocoenosis structure and
appearance – an approach related to consideration
of the plant cultivated origin in terms of symbiosis
(Rindos, 1980).
The proposed scenario would shape the pat-
tern we are observing: the founder crop, with the
largest seeds among others, known only as a culti-
vated plantd recorded archeologically from quite
a restricted area of the Levant. The hypothesis is
in line with supposition by Kislev and Bar-Yosef
(1988), based on favourable nutritional properties
and patchy distribution of wild legumes, that do-
mestication of pulses could predate domestication
of cereals in the Near East. It may draw attention
of archaeologists to a certain relief position where
the traces of the earliest elds could be sought for
if there appear methods to detect them. Finding
of archaeological remnants of a pre-agricultural
society using broad bean as a staple would be
decisive but there is little hope for this since this
hypothetical society, if existed, should have oc-
cupied a very restricted area and for a short time,
to rapidly evolve to an early agricultural society
using the founder crop set.
ACKNOWLEDGEMENTS
I am grateful to Dr. Shahal Abbo and four
anonymous referees for valuable critical com-
ments, to Dr. N.P. Goncharov for fruitful discussion

838 O.E. Kosterin
and to Dr. D. Thomas for the same and linguistic
corrections. This work was supported by the Rus-
sian State Program VI.53.1.3. “Genetic control of
mechanisms of uncompatibility between plant taxa
and their adaptation to unfavourable environmental
conditions”.
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«» (Vicia faba L.)

1, 2
1 Федеральное государственное бюджетное учреждение науки Институт цитологии и генетики
Сибирского отделения Российской академии наук, Новосибирск, Россия,
е-mail: kosterin@bionet.nsc.ru;
2 Новосибирский национальный исследовательский государственный университет,
Новосибирск, Россия
Конские бобы (Vicia faba L.) принадлежат к набору культур, с которых начиналось культивирование
растений на Ближнем Востоке, однако их дикий предок или близкие сородичи до сих пор неизвестны.
Предполагается, что дикий предок бобов имел ограниченный ареал в Леванте и был тесно связан
с растительным сообществом, ограниченным по площади, вследствие чего оказался одомашнен
целиком как вид. Возможно, его местообитания были связаны с границей речной поймы и склона
(так называемый «трансэлювиально-аккумулятивный барьер»), отличающейся благоприятными
почвенными условиями. Эти ограниченные природные местообитания бобов могли стать фокусом
приложения нарождающегося возделывания растений, становясь тем самым прообразом будущих
полей. Предполагается, что конские бобы, будучи заметными высокими растениями с крупными се-
менами и ограниченным местообитаниям, могли служить «стартовой культурой» для возникновения
сельского хозяйства на Ближнем Востоке и способствовать самому возникновению идеи и практики
возделывания растений и «изобретению» поля.
 Vicia faba L., конские бобы, Ближний Восток, происхождение возделывания
растений, доместикация растений, культуры-основатели, стартовая культура, трансиэлювиально-
аккумулятивный барьер.
Cite this article as:
Kosterin O.E. The lost ancestor of the broad bean (Vicia faba L.) and the origin of plant cultivation in the Near East.
Vavilov Journal of Genetics and Breeding. 2014. Volume 18, No. 4/1, pp. 831–840.