Veget Hist Archaeobot (2015) 24:9–18
DOI 10.1007/s00334-014-0498-3
ORIGINAL ARTICLE
Invading a new niche: obligatory weeds at Neolithic Atlit-Yam,
Israel
Anat Hartmann-Shenkman • Mordechai E. Kislev
Ehud Galili • Yoel Melamed • Ehud Weiss
•
Received: 9 February 2014 / Accepted: 22 October 2014 / Published online: 8 November 2014
Springer-Verlag Berlin Heidelberg 2014
Abstract A characteristic group of obligatory weeds was
found in the well of the submerged Pre-Pottery Neolithic C
site of Atlit-Yam, Israel. Identifying these finds to species
level was crucial for defining them as obligatory weeds.
We deal here with the earliest and largest assemblage of
obligatory and facultative weeds in the southwest Asian
Neolithic. Atlit-Yam may reflect a stage in the establishment of weeds in cultivated fields. Weeds are an important
resource for reconstructing the agricultural situation in
archaeological sites, as weed-crop interactions reflect an
agricultural lifestyle. Some of the weeds of Atlit-Yam grow
in fields as well as in Mediterranean herbaceous habitats.
This may indicate that the local herbaceous ecosystem was
the original habitat of the weeds and the place where the
first fields were planted. Presence in a single context of the
earliest identified obligatory grain pest beetle (Sitophilus
granarius) along with obligatory weeds reflects a novel
change made to the ecosystem by the farmers, in which
stored crops were invaded by pests.
Communicated by F. Bittmann.
A. Hartmann-Shenkman E. Weiss (&)
The Martin (Szusz) Department of Land of Israel Studies and
Archaeology, The Institute of Archaeology, Bar-Ilan University,
5290002 Ramat-Gan, Israel
e-mail: eweiss@biu.ac.il
M. E. Kislev Y. Melamed
The Mina & Everard Goodman Faculty of Life Sciences, BarIlan University, 5290002 Ramat-Gan, Israel
E. Galili
Israel Antiquities Authority and Zinman Institute of
Archaeology, The University of Haifa, POB 180, 3030000 Atlit,
Israel
Keywords Pre-Pottery Neolithic C (PPNC) Submerged
settlement Agriculture Obligatory weeds Sitophilus
granarius
Introduction
Weeds invade most agricultural land and are therefore
considered to be characteristic of cultivated fields. The
prehistoric establishment by humans of tilled fields (as well
as other disturbed habitats) enabled numerous unwanted
plants to invade these newly made habitats and evolve as
weeds. Although our herbicide-addicted agriculture is
troubling, weed control in ancient farming was an even
greater problem. Since weed evolution went hand in hand
with crop cultivation, our ancestors had to invest greater
efforts to control these field invaders as agriculture
advanced. The evolutionary adaptation of these weeds to
the agricultural niche moved along multiple paths, which
included increasing seed numbers, mimicking cultivars,
adapting to the cultivars’ growth cycles, and developing
dormancy. We can therefore use weeds as ‘‘index fossils’’
of agricultural ecosystems. In addition, some common
weeds are native to southwest Asia, and their origin may be
linked to the beginning of cultivation (Harlan 1995; Willcox 2012; Zohary 1950; Zohary et al. 2012).
Botanists and ecologists have long dealt with the issue
of weeds and their evolution. Generally, weeds were
described negatively as being useless, injurious, obnoxious
or harmful (Webster 1966), and even homeless (Zohary
1950). For some, like Pieters (1935), weeds took on an
anthropocentric cast with epithets such as ‘‘evil’’ or creatures that ‘‘should be destroyed’’. However, weeds are not
always the farmer’s nemesis, as they sometimes turn out to
produce valuable seeds. The farmer could then plant weed
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seeds, which could then lead to species domestication. For
example, several Old World crops are regarded as ‘‘secondary crops,’’ which entered domestication through the
back door of weed evolution. Such cases are Avena sativa
(oat), Secale cereale ssp. cereale (rye) and Camelina sativa
(gold of pleasure), which actually turned from weeds into
domesticated plants (Vavilov 1949–1950, 1987; Zohary
et al. 2012). In addition, several facultative weed taxa
found at the site are edible, for example Chenopodium
album, Chrozophora tinctoria, Emex spinosa, Hordeum
spontaneum, Scorpiurus muricatus and Silybum marianum
(Zohary 1966, 1972; Feinbrun-Dothan 1978, 1986; Dafni
1984; Feinbrun-Dothan and Danin 1998). Therefore we
need to consider the possibility that they may also have
been gathered for consumption. Importantly, this does not
change their role as weeds if they were gathered in the
fields.
We present here the weed assemblage from Atlit-Yam.
The main importance of this paper is that it presents the
earliest and largest assemblage of obligatory and facultative weeds in the southwest Asian Neolithic. Moreover,
Atlit-Yam is a Pre-Pottery Neolithic C (PPNC) site in the
heartland of the Levantine Neolithic, and therefore belongs
to a stage following the establishment of the full-fledged
agricultural economy of the PPNB. This economy is
manifested in the site’s complex of crop plants, weeds and
a particular insect pest.
The site of Atlit-Yam
The village of Atlit-Yam is currently a submerged PPNC
settlement (Fig. 1), radiocarbon dated to 9,200–8,500 cal
Veget Hist Archaeobot (2015) 24:9–18
BP. It is located 200–400 m west of the present Mediterranean coastline of Israel, 8–12 m below current sea level.
The site was excavated by E. Galili on behalf of the Israel
Antiquities Authority and the University of Haifa. It was a
coastal agro-pastoral village that subsisted on agriculture
and marine resources.
The plant material under discussion, mostly waterlogged, was taken from a single well, Structure no. 11. The
fill of the well contained plant remains, large quantities of
animal bones and various flint, bone and stone artefacts
(Galili and Nir 1993; Galili et al. 1993; Galili and Rosen
2011). It included thousands of seeds and fruit remains
belonging to about 90 species. The finds include:
•
•
•
•
•
Grains of domesticated cereals: Triticum dicoccum
(emmer wheat), T. parvicoccum (small-grained wheat),
T. durum (durum wheat) and H. vulgare (two/six-rowed
barley);
Pulses: Lens culinaris (lentil) and Cicer arietinum
(chickpea);
Fruits: Ficus carica (fig), Pistacia atlantica (atlantic
pistachio), P. palaestina (terebinth tree), P. lentiscus
(lentisk), as well as Vitis sylvestris (wild grape vine);
Wild vegetables: Malva parviflora (small-flowered
mallow) and Rumex pulcher (fiddle dock); and
Herbs and weeds.
This rich assemblage of wild and domesticated
crop plants indicates the centrality of agriculture to the
inhabitants’ life style. Other material remains, such as
dwellings and domesticated animal bones, further demonstrate the site’s domesticated culture.
Since the crop remains are an admixture with a large
assemblage of weeds, we have a unique opportunity to deal
with the evolution of weeds and their early appearance
during the Neolithic period.
The definition of ‘‘weed’’
Fig. 1 The location of the Atlit-Yam site near the coastal plain of
Israel
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The term ‘‘weed’’ has many definitions. In 1935, Pieters
(1935) quoted a restricted definition of weeds from the
Farmers’ Bulletin: ‘‘A plant that does more harm than good
and has the habit of intruding where not wanted’’.
Harlan’s (1965) use of the term was broad:
‘‘For the purpose of this discussion, the word weed is
taken to mean a species or race which is adapted to conditions of human disturbance (Harlan and DeWet 1965).
By this definition, weeds are not confined to plants. Animals such as the English sparrow, the starling, the ‘statuary’ pigeon, the house mouse, Drosophila melanogaster,
and others are especially fitted to environments provided
by human disturbance’’.
These definitions imply that the appearance and abundance of weeds in archaeological material can influence
Veget Hist Archaeobot (2015) 24:9–18
our understanding of ancient cultures. For example, several
generations of archaeobotanists have shown methodologically that finding weeds with crop plants in an archaeological assemblage can indicate that grain remains were
grown in planted fields. In other words, the presence of
weed species in crop remains is associated with past agricultural activities (Zohary 1950; Harlan 1965; Harlan and
DeWet 1965; Wasylikova 1981; Greig 1990; Hillman
1991; Fahmy 1997; Colledge 1998; Jones 2002; Kreuz
et al. 2005; Fuller 2007; Eliáš et al. 2010; Jones et al. 2010;
Kreuz and Schäfer 2011; Willcox 2012; Riehl et al. 2013).
Since obligatory weeds (see below) are clearly associated
with contemporary farming, they are acceptable indicators
for past agricultural activities. Because obligatory weeds
do not grow outside fields, their presence in an archaeobotanical context strongly indicates that these plants had
already entered the favourable niche of farmed lands.
Unfortunately, once there, they have remained an agricultural curse for thousands of years, through to our own
times.
Classification of weeds
In weed research, weedy taxa are clustered into several
categories. The main groups are obligatory and facultative
weeds, which were defined by Michael Zohary (1950):
Obligatory weeds are ‘‘not found as yet outside segetal
[field] habitats. Many of them are rather widespread….
These are, undoubtedly, ancient plants which have lost
their primary habitats [i.e. natural vegetation] during
the long history of cultivation. Many of them are
known from prehistoric times. They have been termed
archaeophytes…’’
Facultative weeds are ‘‘plants occurring in both segetal
and primary habitats.’’
We relate here to both of these groups. However, following the terminology of Zohary and other previously
mentioned researchers, obligatory weeds are the best
markers for agricultural activity in archaeobotanical
assemblages. This is because, in most cases, their only
habitats are cultivated fields.
Materials and methods
Sampling
The first stage of sampling was on site. The samples were
excavated underwater with dredger suction, mostly in
20 cm layers of the well depth. The divers took the samples
to shore in bags, which were placed separately into plastic
boxes with fresh water. On the shore, large stones and
artefacts were removed manually from the boxes and the
11
fine material underwent several stages of sieving: floating
and hovering waterlogged organic material was collected
in a 0.5 mm strainer, while the heavy material sank to the
bottom of the box. Some of the material was collected by
picking. The waterlogged organic material was kept in
sealed jars for wet storage and analysis.
Laboratory work
The analysis work was conducted in the laboratory. The
carbonized material was taken out of the water and dried.
The waterlogged material was left in the water. To prevent
mould development, we replaced the original water with a
4 % solution of the fungicide Nipagin (methyl paraben/
para-hydroxybenzoic acid methyl ester; 4 g of Nipagin per
1 l of water). Sorting the samples was carried out using a
stereoscopic microscope at a magnification of up to 509;
mainly seeds and fruits were sorted out (other types of
remains were mostly disregarded). The plants were identified mainly on the basis of their morphology, following
sorting into types, sizes, shapes and surface textures. The
identification was aided by our reference collection. We
followed Fragman-Sapir et al. (1999) for English plant
names.
Finally, some of the seeds were photographed by stereoscopic light microscope using an Olympus SZX10, or
by SEM using an XL 30 FE (field emission), FEI. The SEM
samples were coated with gold and examined at 200–500x
magnifications.
Results
Some 8,500 seeds and fruit remains representing some 90
species were identified from the well. The implication of
the large number of weed species there is discussed below.
The weed assemblage
The obligatory and facultative weeds from Atlit-Yam are
an important resource for reconstructing the agricultural
activities of its inhabitants. Following Zohary’s grouping
(1950), we found remains of weeds representing some 35
species representing two groups, five obligatory and 30
facultative weed species (Table 1; Figs. 2, 3, 4, 5 and 6).
The weeds and other finds were distributed throughout the
various depths of the well assemblage (Table 2).
As far as we know, this is the earliest and largest
assemblage of weeds from the southwest Asian Neolithic.
Furthermore, identifying these five obligatory weeds to
species level is significant, as all of them are non-monotypic, so they have other species in their genera.
Undoubtedly, another species of the genus could have a
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Veget Hist Archaeobot (2015) 24:9–18
Table 1 The weeds of Atlit-Yam; all these species still function as
weeds in cultivated fields, those with an asterisk also grow in
Mediterranean semi-steppe herbaceous habitats (see text; see Figs. 2,
3, 4, 5 and 6)
Fig. 2 Bupleurum subovatum. Mericarp, dorsal view. The fruit is
long, ovoid, prominently ribbed and densely tuberculate in the wide,
shallow furrows (SEM)
totally different ecological signature and could not serve as
a proxy for agricultural activity. Luckily, the excellent
preservation of these waterlogged samples allowed for
species level identification.
The identification to species also provides us with an
improved understanding of weed emergence. Some of
these weeds (see Table 1, the asterisked species) still grow
in local fields, as well as in Mediterranean semi-steppe
herbaceous habitats (Zohary 1962). These examples may
indicate that the herbaceous ecosystem was their original,
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Fig. 3 Lolium temulentum. Grain, ventral view. The narrow ventral
furrow becomes wider and shallower on the upper part of the grain
(SEM)
pre-weed habitat, and that the first fields originated in this
ecosystem.
The obligatory weeds
The following are the descriptions and the identification
criteria of the obligatory weeds from Atlit-Yam.
Veget Hist Archaeobot (2015) 24:9–18
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Fig. 6 Tordylium aegyptiacum. a ventral wall of flattened, small
mericarp; b recent mericarp with distinct, somewhat moniliform
margin for comparison, dorsal view. Unfortunately, the delicate
mericarp was broken during photo processing, revealing the dorsal
ridge. The inner part of the margin is viewed at the perimeter, while
its moniliform outer part is missing (stereo microscope photo)
Fig. 4 Phalaris paradoxa. Grain, side view. The fertile grain is
enveloped by the lemma and the palea. The minute two lemmas (at
the bottom) are actually reduced sterile florets (SEM)
characters that distinguish this species, which is exclusively segetal in Israel. The second character is the fusion
of two mericarps into a single dispersal unit (Zohary 1972;
Danin 2004), which are similar in length and width to those
of T. parvicoccum grain. The fused mericarps, which are
unique to the genus and also unusual in the Apiaceae,
suggest that the fruit of the segetal B. subovatum adapted to
the T. parvicoccum grain in shape and size. Moreover, the
wide and generally perfoliate shaped leaves of the two
species are unique to their genus. This resemblance might
suggest that B. subovatum evolved from the closely related
B. lancifolium.
Lolium temulentum L. (Poaceae), Darnel
Fig. 5 Raphanus raphanistrum. Sterile portion of the siliqua upper
joint, side view. The neighbouring joints are fertile (SEM)
Bupleurum subovatum Spreng. (=Bupleurum lancifolium
var. lancifolium; B. intermedium Poir.) (Apiaceae), False
thorow-wax
Two mericarps were found (Fig. 2), one at a well depth of
310–350 cm, and the second is from an unknown depth.
They are characteristic in shape and pattern (Zohary 1972).
Beupleurum subovatum was formerly the typical variety
of B. lancifolium, of which the other varieties are nonweedy, wild plants, and which still belong to the last species. A 4 mm long mericarp is one of the two main
A single carbonized grain without hulls was identified
(Fig. 3), found at an unknown depth in the well. Lolium
grains have a small and elliptic embryo, and a hilum in a
longitudinal furrow which equals the length of the grain
(Bor 1968). The grain was identified to this species because
of its relative size, width and elliptic shape.
Phalaris paradoxa L. (Poaceae), Bristle-spiked Canary
grass
Nine carbonized hulled grains of Phalaris paradoxa were
found (Fig. 4), all of them from a depth of 250–270 cm. It
was easy to identify its genus, thanks to the characteristic
shape of the grain, where its thickness is much greater than
its width. In addition, the grain is tightly enclosed by the
palea and glossy lemma, which are clearly visible in the
archaeological find. One of the grains was subtended at its
base by 2 min, unequal lemmas of sterile florets (Feinbrun-
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Veget Hist Archaeobot (2015) 24:9–18
Table 2 Crop and weed species (?pest beetle) with depth (cm) in the Atlit-Yam well
Dothan 1986). Based on these sterile lemmas, other Phalaris species were rejected.
Raphanus raphanistrum L. (Brassicaceae), Wild radish
Two carbonized and two waterlogged degraded portions of
the siliqua upper joint were found (Fig. 5), all of them from
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a depth of 250–270 cm. They are characteristic of this
species in shape, pattern, texture and dimensions. The
upper joint has 2–8 seeded portions (and sometimes a few
sterile ones) with constrictions between them, readily
breaking into longitudinally ridged 1-seeded (or sterile)
portions. In Israel, this species is common in the coastal
plain (Zohary 1966).
Veget Hist Archaeobot (2015) 24:9–18
Tordylium aegyptiacum (L.) Lam. (Apiaceae), Egyptian
hartwort
Six waterlogged umbel peripheral mericarps were found
(Fig. 6), all of them from a depth of 250–270 cm. They are
strongly flattened, orbicular to suborbicular, somewhat
notched at base and apex.
The middle dorsal ridge can be observed. We compared our
remains with all local species of the tribe Peucedaneae, especially
the other species of Tordylium, as well as the closely related
Peucedanum junceum, but they were not similar to our finds.
Discussion
This large group of weeds from Atlit-Yam is a case-study
for weed research, because it represents their very early
establishment in cultivated fields. In addition, the weeds
provide persuasive evidence for the antiquity of the interrelationship between weed flora and agricultural fields.
There are several broad implications of this study.
(a) Atlit-Yam is the earliest known assemblage of obligatory weeds: In order to understand the obligatory weed
community from Atlit-Yam in relation to contemporary
agricultural communities, we checked archaeobotanical
assemblages from other Pre-Pottery Neolithic southwest
Asian sites. Although many PPN sites revealed domesticated
crops, their weed finds are remarkably limited. In most
reports, obligatory weeds are completely absent. Some PPN
sites include finds which can potentially be regarded as
obligatory weeds. However, these taxa were only identified
to genus or sp./cf. level, not to species level (Table 3). As we
mentioned earlier, weeds are useful ecological markers only
if they are identified to species level.
There are several earlier reported cases where obligatory
weeds were identified to species level, all in southwestern
Asian Pre-Pottery Neolithic B agrarian sites. Galium tricornutum was found in Can Hasan III, Turkey (Hillman
1972, 1978), and in Yiftah’el, Israel (Garfinkel et al. 1988).
Lolium temulentum was found at Ramad; Vaccaria hispanica was found at Aswad and Ramad, Syria (van Zeist and
Bakker-Heeres 1982 (1985)), and also in PPNA M’lefaat
(Savard et al. 2003). Two possible additional sites are Tepe
Ali Kosh, Ali Kosh phase, where Phalaris cf. paradoxa
was found (Helbæk 1969) and Nevali Çori where Lolium
cf. temulentum was found (Damania et al. 1998).
The comprehensive ensemble of weeds from Atlit-Yam
includes five obligatory and 30 facultative weeds. It represents, therefore, the establishment of weeds as a characteristic
group within the agricultural cycle. This is, of course, not the
‘‘beginning of weeds,’’ but rather the earliest representation of
a complex weed population and not the sporadic appearance
of a single weed.
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(b) Weed-crop relationships reflect local agriculture:
Contemporary observations have shown that particular
weeds have a preference, or adaptation, for specific crops,
or group of crops (Harlan 1965). At Atlit-Yam, we can
distinguish a group of obligatory weeds, typical of cereal
fields with P. paradoxa, L. temulentum, B. subovatum, R.
raphanistrum and T. aegyptiacum (Zohary 1941).
Therefore, the combined plant assemblage of crops and
weeds from Atlit-Yam can give additional information
regarding the establishment of agriculture in the PPNC.
Even in this prehistoric period we can see the state of coevolution between certain obligatory weeds and their crops.
(c) A grain pest beetle and obligatory weeds invade the
same human-made ecological niche: Interestingly, in
addition to the noxious weeds from Atlit-Yam, we also
identified the remains of some 27 specimens of S. granarius (granary weevil). Their numbers are based on finds of
the prothorax, the fore segment of the thorax, which is the
largest and hardest part of the insect and bears the front pair
of legs (Fig. 7). In addition, the samples also contained six
elytra, the protective wing cases that generally protect the
functional wings, the latter being only vestigial in this
species. Sitophilus granarius is a locally extinct pest beetle
of stored grains, reviewed by Kislev et al. (2004). Together
with contemporary finds from Hacılar, southwest Anatolia,
Turkey (Helbæk 1970), these remains are the earliest
known evidence of a pest associated with domesticated
grains. Today, this beetle is widespread, mainly in the
north temperate zone, and it is naturally absent from our
warmer region. The best conditions for its development are
cool environments, down to 11 C, with high relative
humidity, and the beetle is therefore more successful in
temperate regions (Howe 1965; Dobie et al. 1991). It seems
that the cooler and more humid climate prevailing in the
prehistoric southwest Asian Neolithic (Bar-Matthews et al.
1997; Kislev et al. 2004), along with human grain storage,
enabled the spread of S. granarius to our region. Later,
when the area became warmer and drier, S. granarius
suffered extinction.
The remains of obligatory weeds together with grain
weevils suggest that Atlit-Yam represents an advanced agricultural economy of crop growing and storage. In addition,
agriculture not only served to provide food, but also functioned as a significant ecological factor that influenced a vast
array of organisms taking up residence in new human-made
ecological niches.
Conclusions
Atlit-Yam was occupied some 2,000 years after the
beginning of sowing and some 1,000 years after the first
morphological signs of domesticated cereals.
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Veget Hist Archaeobot (2015) 24:9–18
Table 3 Potential obligatory weeds in Pre-Pottery Neolithic southwest Asian sites
Studying the site has enabled us to demonstrate aspects
of the complex interactions operating in a more advanced
agriculture. Aside from the domesticated cereals and pulses
found there, we have identified the earliest and largest
weed assemblage from the southwest Asian Neolithic.
There are several plant associations where herbs could
have turned into weeds. These include semi-steppe and
savanna-like vegetation, such as the Ziziphus spina-christi
vegetation association in the southwest Golan and eastern
lower Galilee, and open woods, such as the Quercus
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ithaburensis-Styrax officinalis and Ceratonia siliqua-Pistacia lentiscus associations, both of which can be found
near Atlit-Yam (Zohary 1962). Some of these weeds may
indicate that the original habitats of various contemporary
weeds were in the Mediterranean herbaceous ecosystem,
which was also the ecosystem of the first fields.
D. Zohary described the adaptation of annual crops to
their new field habitats’’ as follows: ‘‘Unconscious or
automatic selection brought about by the fact that the
plants concerned were taken from their original wild
Veget Hist Archaeobot (2015) 24:9–18
Fig. 7 Sitophilus granarius. Prothorax of an adult beetle, dorsal
view. The punctures are distinctly oblong. In each of them there is a
short hair, most of them are preserved (SEM)
habitat and placed in new (and usually very different)
human-made or human-managed environments. The shift
in the ecology led automatically to drastic changes in
selection pressures. Numerous adaptations vital for survival in the wild environments lost their fitness under the
[new] sets of conditions’’ (Zohary 2004).
We suggest that the weeds-to-be were also adapted to
the field habitat by similar unconscious selection. Only
those herbs that succeeded in adapting to their new ecosystems could pass over various ecological (and possibly
evolutionary) barriers and become weeds.
Acknowledgments We would like to thank E. Donahaye of the
Department of Plant Protection, Beit-Dagan, Israel, for confirming the
identification of S. granarius; J. Langsam for the SEM micrographs;
Y. Mahler-Slasky for the stereo-microscope pictures and we thank the
anonymous reviewers for their constructive comments. We also thank
the COST program, action TD0902, SPLASHCOS of the European
Commission, the National Geographic Research Foundation, CARE
Archaeological Foundation, MAFCAF Foundation, and Sandy and
Joseph Lepelstat for their financial support of research on the submerged Carmel coast sites. The Israel Antiquities Authority, the Israel
Prehistoric Society and Haifa University are acknowledged for their
financial and administrative support of the underwater excavations.
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