The phytolith archaeological record: strengths and weaknesses evaluated

based on a quantitative modern reference collection from Greece

Georgia Tsartsidou
a,
*
, Simcha Lev-Yadun
b
, Rosa-Maria Albert
c
, Arlene Miller-Rosen
d
,
Nikos Efstratiou
e
, Steve Weiner
f
a
Ephoreia of Palaeoanthropology-Speleology of Southern Greece, Ardittou 34b, Athens 116 36, Greece
b
Department of Biology, University of Haifa, Oranim, Tivon 36006, Israel
c
Catalan Institution for Research and Advanced Studies (ICREA), Research Group for Palaeoecological and Geoarchaeological Studies,
Department of Prehistory, Ancient History and Archaeology, University of Barcelona, c/ Baldiri Reixac, s/n, 08028 Barcelona, Spain
d
Institute of Archaeology, University College London, 31e34 Gordon Square, London WC 1H OPY, UK
e
Department of Archaeology, Faculty of Philosophy, University of Thessaloniki, 54006 Thessaloniki, Greece
f
Department of Structural Biology, Weizmann Institute of Science, Rehovot 76100, Israel
Received 7 May 2006; received in revised form 22 September 2006; accepted 20 October 2006
Abstract
The phytolith record from archaeological strata is a powerful tool for reconstructing aspects of past human behaviour and ecology. Consider-
able insights into the problems and potentials of this record can be obtained by studying phytoliths in modern plants. We prepared a phytolith
reference collection of modern plants mostly from northern Greece that includes quantitative information on phytolith concentrations (number of
phytoliths per gram dry organic material) and morphological assemblages. Here we analyse this reference collection with the aim of evaluating
the strengths and weaknesses of the archaeological phytolith record. The reference collection comprises 62 different plant taxa. Of them, 28 are
trees, 1 conifer, 10 shrubs and 1 herb, 3 domesticated annual legumes and 19 monocotyledons. We demonstrate quantitatively the extent to which
woody species, legumes and fruits of dicots contribute only small amounts of phytoliths to the sediments per unit tissue dry weight, while leaves
of trees and shrubs produce signicant amounts of phytoliths and grasses are prolic phytolith producers. We compare the data of this reference
collection with a similar reference collection from another Mediterranean ecosystem (Israel). The comparison indicates that some aspects of
phytolith production are probably genetically controlled, whereas others are environmentally controlled. We note that despite the fact that woody
species produce few phytoliths per gram of tissue, their record can be most informative when taking into account other properties of the
sediments. We also note that jigsaw puzzle-shaped phytoliths normally form in plants that grow in humid conditions, but may form in arid
environments when the plants are irrigated. This study clearly shows the extent to which some plants can be under-represented and others
over-represented in the phytolith record. Knowing the extent of this bias can greatly improve our interpretation of the phytolith record.
2006 Elsevier Ltd. All rights reserved.
Keywords: Phytoliths; Quantitative reference collection; Greece; Plant microfossils; Archaeobotany
1. Introduction
Plants are an important component of the archaeological
record. Only the charred remains of the plants themselves
are usually preserved, and this often represents a small fraction
of the plants brought to the site by humans or their animals, or
those that grew at the site. A potentially more complete
botanical record can be obtained from the siliceous phytoliths.
These mineralized elements are produced, often in large quan-
tities, by many plants. They are relatively stable in many
sedimentary environments and thus may accumulate in large
amounts in archaeological sites. In order to best interpret the
* Corresponding author. Tel.: 30 2108310372; fax: 30 2107561438.
E-mail address: georgiat@arch.auth.gr (G. Tsartsidou).
0305-4403/$ - see front matter 2006 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jas.2006.10.017
Journal of Archaeological Science 34 (2007) 1262e1275
http://www.elsevier.com/locate/jas
phytolith record of an archaeological site it is necessary to
document which local plants are relevant to the archaeological
record, produce phytoliths and to determine the absolute quan-
tities produced per unit weight of plant material. It is also
necessary to differentiate between phytolith shapes that are
unique to a specic plant or group of plants, and those that
are common to many plant types. This can be done using
a so-called quantitative reference collection (Albert and
Weiner, 2001) to determine not only which plants were intro-
duced into the site, but also the approximate quantities of plant
material brought into the site. Furthermore, a detailed analysis
of the phytolith distribution within a site based on such a refer-
ence collection may also reveal the use of a particular area of
the site for a specic purpose. One such reference collection
was produced for plants in the area of the prehistoric Kebara
Cave, Mt. Carmel, Israel (Albert et al., 2000). Albert et al.
(2003) used this reference collection to estimate the amounts
of various dry plant material types that were brought into
Kebara and Hayonim caves, northern Israel, in the Mousterian
(Middle Paleolithic) and Natuan periods. Here we report the
results of a second, more comprehensive phytolith reference
collection mostly from northern Greece. These results, to-
gether with a comparison of the two collections, provide
new insights into the strengths and weaknesses of the phytolith
archaeological record.
Many phytolith reference collections have been produced in
different regions (Bozarth, 1992; Jones and Bryant, 1992;
Ollendorf et al., 1988; Piperno, 1984; Scott, 1992). Most of
these involve identifying only the phytolith types in a given
plant, and paying particular attention to those that are unique
to a particular plant taxon. This information is then used to
identify the use of that plant at a given archaeological site.
This application is often limited as many plants do not produce
uniquely shaped phytoliths. Nevertheless many studies have
been conducted with respect to size and shape measurements
in order to solve this problem and they succeed in differentiat-
ing phytoliths from closely related taxa (Ball et al., 1996,
1999; Berlin et al., 2003; Pearsall et al., 1995; Zhao et al.,
1998). Another approach is to divide the phytolith types into
broader categories, such as being derived from wood and
bark, from leaves, from grasses and in particular from cereals
(Rosen, 1992). This is archaeologically very useful, as the ex-
ploitation of such plant types is often indicative of a particular
behaviour, such as fuel used for res and fodder collected for
animals. There are however several inherent problems with
this approach. One is that the same phytolith types are some-
times produced in different plants. It is thus necessary to use
the modern reference collection to allocate the approximate
proportions of these common phytoliths to different plant cat-
egories. See Albert et al. (2003) for more details. Another
problem is that many phytoliths have variable morphologies,
with irregular and not identiable shapes, as opposed to those
with consistent morphologies. Albert et al. (1999) noted that
the ratio of variable to consistent morphology phytoliths (v/c)
can be indicative of certain plant types.
The absolute production of phytoliths in plants per unit
weight of dry plant material is for the most part genetically
determined (Hodson et al., 2005; Piperno, 1988). Plants that
produce large amounts of phytoliths (such as grasses, sedges
and palms), do so irrespective of geographical region
(Bamford et al., 2006; Piperno, 1988). Thus estimates of
amounts of plant types used can be made by comparing gen-
eral plant categories such as grasses, dicots, cereals and le-
gumes. Phytolith production is however also inuenced to
some extent by the environment; concentrations of monosilicic
acid in soils, the temperature and water content of soils, the pH
and the climate (Jones and Handreck, 1965; Madella et al.,
2002; Piperno, 1988; Rosen and Weiner, 1994). The reference
collection reported here was prepared as part of a broader
ethno-archaeological study of a traditional village called Sar-
akini, located in the Rhodope Mountains of northern Greece
(Fig. 1). In so doing it includes the most common plants of
the area relevant to everyday life of the ethnographic environ-
ment and it is not an exhaustive analysis of the plants of north-
ern Greece. Some species such as Zea mays, which is
a modern staple of the village, were also analysed even though
they do not have any local archaeological signicance. The
reference collection will also constitute the basis for analysing
the phytoliths from the Neolithic site of Makri, located on the
coast in the area of Thrace.
Here we highlight the relative amounts of phytoliths, as
well as the phytolith types produced by 62 different plant
taxa collected from northern and central Greece, including
woody dicots and a conifer, shrubs, dicot herbs and monocot-
yledons. This quantitative information can be used to offset the
biases of the phytolith record. We also focus on aspects of the
data that provide insights into the strengths and weaknesses of
the phytolith component of the archaeological record. All the
data is available upon request from G.T.
2. Materials and methods
Sixty-two plant species were collected during the summer
of 2003 from three different areas of Greece (Table 1). These
comprise 28 trees (woody dicotyledons), 1 conifer (gymno-
sperm), 10 shrubs, 1 herb, 3 domesticated annual legumes
and 19 monocotyledons, 5 of which were cereals and 1
a rush. All the dicotyledonous trees and shrubs, the conifer
and most of the grasses are from the mountainous area of
Rhodope, northern Greece, and the plain at the foot of the
mountains. The legumes and some monocotyledons (Lagurus
ovatus, Arundo sp., Hordeum vulgare, Hynaldia villosa, Briza
maxima, Juncus sp.) were collected during the same summer
from central Greece (Boeotia, Attika, Argolid). We chose
plants that are commonly used by humans now or in antiquity.
These include trees that are useful for construction (e.g., Ju-
glans regia), trees that are commonly used as fuel (Quercus
sp.), edible plants such as fruits, legumes and cereals, shrubs
that may have been used for medical or other reasons (e.g.,
Salvia ofcinalis, Thymus capitatus), wild grasses that animals
eat in the open or people feed them and cereals that are also
used for construction activities (Secale cereale).
Most of the plants were separated into different parts in
order to acquire information on the potential contribution of
1263 G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275
a specic plant part to the phytolith record. This is sometimes
most helpful, since different parts of plants may reect various
human or animal habits and activities, including seasonal ac-
tivities. Most of the grasses and all the cereals were separated
into two parts: inorescence (including the grains) and the
leaves/stems together. Tree species were mechanically sepa-
rated into ve parts whenever possible: wood, bark, leaves,
owers and fruits. Herbaceous plants were analyzed whole
and thus include stem, leaves and often the owers. A total
of 148 samples were analysed.
We compared our results with those of Albert and Weiner
(2001) for plants from Mt. Carmel, Israel. This collection con-
sists of 29 plant species, including 11 woody dicotyledons.
More recently a few grass species from Mount Carmel area
have also been analysed (Albert et al., submitted for publica-
tion). Table 2 lists all the species analysed.
2.1. Phytolith extraction
The plants were cleaned extensively by washing in de-
ionized water in a sonicator to avoid contamination from
phytoliths of other plants adhering to their surfaces. The
dried plants were then weighed. They were burnt in a fur-
nace at 500

for 4 h. The ash was treated with 1 N HCl
and centrifuged (3500 rpm for 10 min), resuspended in de-
ionized water and centrifuged again. The supernatant was
discarded and the washing was repeated three times. Since
there was charcoal in almost all the samples, the samples
were re-ashed at 500

C for 90 min and then weighed.
The ash was again treated with 1 N HCl and washed in
a centrifuge as above. This constituted the acid insoluble
fraction (AIF), which reects also the yield of silica
(opal) in the plants. The AIF fractions were then dried un-
der a heat lamp and weighed. We noted that in all the bark
and some of the wood and leaf samples the AIF did not
contain only silica, as was the case in most of the leaves
and all the monocotyledons, but also included mainly quartz
and clay. In two cases the calcium sulphate mineral, anhy-
drite, was a major component of the ash (Juglans regia,
Tamarix sp.). In order to obtain more accurate estimates
of the phytolith per gram plant material values, ash samples
with soil derived minerals were further puried by centri-
fuging in 2.4 g/ml density heavy liquid (sodium polytung-
state: Na
6
(H
2
W
12
O
40
)$H
2
O). The AIF was transferred
into a centrifuge tube and 5 ml of heavy liquid were added.
The suspension was sonicated for 10 min until it was well
dispersed. It was then centrifuged at 3500 rpm for 20 min.
The supernatant was transferred to another centrifuge tube,
1 ml of water was added and again centrifuged as above.
This cycle was repeated once more and then the tube was
lled with water for the nal centrifugation. In so doing
the denser minerals were separated from the less dense
silica in four stages. The plants that were subject to the ad-
ditional treatment included all the bark samples, and from
the wood samples Salix sp., Fagus sylvatica, Juglans regia,
Alnus glutinosa, Pistacia terebinthus, Quercus pubescens,
Fig. 1. Map of Greece. The areas from which the plants of the reference collection have been collected are shown (Thrace, Argolid, Boeotia).
1264 G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275
Table 1
List of all the plants collected from Greece for phytolith analysis
Latin name Family English name Provenance
Woody plants
Acer monospessulanum Aceraceae Maple Thrace M
Alnus glutinosa Betulaceae Black alder Thrace M
Arbutus adrachne Ericaceae Strawberry tree Thrace M
Carpinus betulus Betulaceae Hornbean Thrace M
Celtis australis Ulmaceae Nettle tree Thrace C
Corylus avelana Corylaceae Hazelnut Thrace M
Crataegus sp. Rosaceae Hawthorn Thrace M
Fagus sylvatica Fagaceae Beech Thrace M
Ficus carica Moraceae Fig tree Thrace M
Fraxinus ornus Oleaceae Manna ash Thrace M
Juglans regia Juglandaceae Walnut Thrace M
Juniperus oxycedrus Cupressaceae Juniper Thrace M
Olea europaea Oleaceae Olive tree Thrace C
Phillyrea latifolia Oleaceae Basswood Thrace P
Pistacia terebinthus Anacardiaceae Terebinth Thrace P
Populus alba Salicaceae White poplar Thrace M
Prunus amygdalus Rosaceae Almond tree Thrace C
Prunus avium Rosaceae Wild cherry Thrace M
Pyrus amygdaliformis Rosaceae Wild pear Thrace M
Quercus coccifera Fagaceae Kermes oak Thrace P
Quercus frainetto Fagaceae Hungarian oak Thrace M
Quercus petraea Fagaceae Durmast oak Thrace M
Quercus pubescens Fagaceae Downy oak Thrace M
Salix sp. Salicaceae Willow Thrace M
Sambucus nigra Caprifoliaceae Common elder Thrace M
Sorbus torminalis Rosaceae Wild service tree Thrace M
Tamarix sp. Tamaricaceae Tamarisk Thrace P
Tilia sp. Tiliaceae Lime Thrace M
Vitex agnus castus Verbenaceae Chaste tree Thrace P
Herbaceous plants and shrubs
Cichorium sp. Cichoriaceae Chicory Thrace M
Ephedra altissima Ephedraceae Ephedra Thrace C
Euphorbia sp. Euphorbiaceae Spurge Thrace M
Nicotiana tabacum Solanaceae Tobacco Thrace M
Paliurus spina christi Rhamnaceae Christs thorn Thrace P
Pteridium aquilinum Hypolepidaceae Fern Thrace M
Rubus sanctus Rosaceae Blackberry Thrace M
Salvia ofcinalis Labiatae Sage Thrace M
Thymus capitatus. Labiatae Thyme Thrace M
Urtica sp. Urticaceae Stinging nettle Thrace M
Vitis sylvestris Vitaceae Grape-vine Thrace M
Domesticated annual legumes
Cicer arietinum Leguminoseae Chickpea Boeotia P
Lens culinaris Leguminoseae Lentil Boeotia P
Vicia faba Leguminoseae Faba bean Boeotia P
Monocotyledons
Aegilops sp. Gramineae Goat grass (wild wheat) Thrace M
Arundo sp. Gramineae Reed Boeotia P
Briza maxima Gramineae Quaking grass Argolid P
Bromus sp. Gramineae Brome Thrace M
Cynodon dactylon Gramineae Bermuda grass Thrace M
Dactylis glomerata Gramineae Orchard grass Thrace M
Hynaldia villosa Gramineae Argolid P
Lagurus ovatus Gramineae Hare tail Attika P
Lolium sp. Gramineae Ryegrass Thrace M
Melica sp. Gramineae Melick grass Thrace M
Phleum sp. Gramineae Cats tail Thrace M
Polypogon monspeliensis Gramineae Beard grass Thrace M
Stipa bromoides Gramineae Needle grass Thrace M
Juncus sp. Juncaceae Rush Argolid P
(continued on next page)
1265 G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275
Quercus frainetto, Quercus coccifera, Juniperus oxycedrus,
Tamarix sp., Acer mosnospessulanum, Carpinus orientalis,
Olea europaea, Corylus avelana, Populus alba and Celtis
australis. The leaves of the following plants were also fur-
ther puried: Prunus avium, Rubus sp., Salix sp., Sambucus
nigra, Pyrus amygdaliformis, Tamarix sp., Crataegus sp., Vi-
tis sylvestris, Vicia faba, Lens cullinaris, Salvia ofcinalis
and Thymus capitatus.
The two last fractions contained the phytoliths and after
drying under a heat lamp they were weighed. Slides of both
fractions were prepared and counted. About 1 mg of AIF
was accurately weighed and then placed on the slide, six
drops of Entellan were added and a cover slip was lowered
carefully over the well-mixed suspension. At least 200 phy-
toliths were counted in each slide, when possible. Albert
and Weiner (2001) demonstrated that the counting of 194
phytoliths with consistent morphology gives an error margin
of 23%. We note that siliceous aggregates were not included
and silica skeletons were counted as one phytolith. This
could be a problem in calculations as silica skeletons are
usually more common in the plant phytolith records than
in the sediments. Our analysis of the sediments in the eth-
nographic environment as well as in the Neolithic site
shows that they are rich in silica skeletons and these are
not destroyed during sample preparation. The variability in
the degree of silicication in plants that could raise or lower
the number of silica skeletons is offset by the fact that the
reference collection is prepared in the same geographic re-
gion as the sediments analysed. The number of phytoliths
in the slide (consistent and variable morphology) was then
related to 1 g of dry plant material.
2.2. Phytolith classication
The phytoliths were classied morphologically according
to their anatomical origin and when this was not possible, ac-
cording to their geometrical shape. Following Albert et al.
(1999) they were separated into two major categories: those
with consistent morphologies that repeat themselves in more
than one sample and those with variable morphologies, with
irregular and unidentiable morphologies. Phytoliths with
consistent morphologies were classied and their relative pro-
portions determined in percent. The terminology used for con-
sistent morphology phytolith classication followed Albert
(2000), Brown (1984), Madella et al. (2005), Metcalfe
(1960), Piperno (1988) and Twiss et al. (1969).
3. Results and discussion
Fig. 2 shows the number of phytoliths per gram dry plant
material of all the plants analyzed in this study. The samples
were sub-divided into six categories in order to better evaluate
the contribution of each category to the archaeological record.
The overall range is from zero to 12 million phytoliths/g dry
plant material. It can readily be seen that the potential contri-
butions of different plants as well as different parts of plants to
the phytolith record, vary enormously. By knowing the phyto-
lith production per unit weight of each species, and in some
cases also the variations in phytolith production within a plant,
the quantities of different plant materials brought into an ar-
chaeological site can be inferred. This is certainly of much ar-
chaeological interest (see Albert and Weiner, 2001, for more
details). The following is a more detailed analysis of phytolith
production in different plant types following Fig. 2.
3.1. Wood and bark
Fig. 2a shows that many wood samples contain no phyto-
liths at all, and those that do, do not exceed 400 phytoliths/
gram dry wood. Bark samples contain an order of magnitude
more phytoliths per gram (Fig. 2b). Bark samples are however
prone to being contaminated by phytoliths from other plants
adhering strongly to the bark surface (see below). The most
prominent examples of contamination we found were Celtis
and Fagus bark samples, where the morphologies of most of
the phytoliths clearly show that they are derived from dicot
leaves and/or grasses. The average number of phytoliths/
gram in bark (excluding Celtis and Fagus) is 1300, compared
to 63, the average number of phytoliths per gram in wood. We
note that the weight proportion of wood and bark in small to
medium sized branches or trunks ranged from 4:1 to 9:1.
Wood and bark contain relatively few phytoliths per unit
weight of dry plant material. Furthermore, many of these
phytoliths have variable morphologies and cannot be uniquely
identied with wood and bark of specic taxa. There are how-
ever certain phytolith types (e.g., ellipsoids and spheroids
(globular) with smooth margins), that are indicative of having
been derived from wood and bark.
3.2. Edible parts of dicots (fruits, nuts, legumes)
The edible parts of trees (fruits, nuts), as well as legumes,
produce extremely small amounts of phytoliths (from a few
tens to 2000 phytoliths/g) (Fig. 2c). The exceptions are the
Table 1 (continued)
Latin name Family English name Provenance
Domesticated cereals
Avena sativa Gramineae Oats Thrace M
Hordeum vulgare Gramineae Barley Boeotia P
Secale cereale Gramineae Rye Thrace M
Triticum aestivum Gramineae Bread wheat Thrace M
Zea mays Gramineae Corn Thrace M
The provenance is shown: M, mountain; C, coastal plain; P, plain at the foot of the mountain.
1266 G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275
fruits of Ficus carica and the acorns of Quercus pubescens
(whole fruits) that produce 140,000 and 350,000 phytoliths/g
respectively. Thus, in general, the contribution of fruits to
the phytolith record will be minimal.
3.3. Leaves and inorescence of dicotyledonous
trees and shrubs
Fig. 2d shows that the leaves and owers of trees and
shrubs produce a huge range of phytoliths per gram of
dry material (from one hundred to millions/gram). The
widespread production of phytoliths in leaves of trees and
shrubs can therefore be most helpful for identifying plant
species in archaeological or geological sediments, provided
a reference collection of the surrounding area is available.
Of the 38 leaf samples analysed in this study, Celtis
australis and Urtica sp. contain the highest concentrations of
phytoliths. Both these plants are characterised anatomically
by hairy leaves reected in their rich phytolith record (52%
and 63% hair phytoliths respectively). Note too that 94% of
Table 2
Plant reference collections from Israel and Greece
Israel collection Greek collection
Species Provenance Plan part Phyt. gram
dry material
Species Provenance Plant part Phyt. gram
dry material
Avena sterilis Tel-Dor Leaves 7,000,000 Avena sterilis Thrace Culm/leaves 600,000
Avena sterilis Tel-Dor Stem 1,000,000
Avena sterilis Tel-Dor Inorescence 2,000,000 Avena sterilis Thrace Inorescence 2,000,000
Lagurus ovatus Tel-Dor Leaves 1,000,000 Lagurus ovatus Attika Culm/leaves 7,000,000
Lagurus ovatus Tel-Dor Stem 40,000
Lagurus ovatus Tel-Dor Inorescence 400,000 Lagurus ovatus Attika Inorescence 7,600,000
Hordeum vulgare Tel-Dor Whole plant 700,000 Hordeum vulgare Boeotia Culm/leaves 1,500,000
Hordeum vulgare Tel-Dor Inorescence 400,000 Hordeum vulgare Boeotia Inorescence 1,000,000
Triticum aestivum Tel-Dor Inorescence 400,000 Triticum aestivum Thrace Husk 4,000,000
Triticum aestivum Mt Carmel Whole plant 1,500,000 Triticum aestivum Thrace Culm/leaves 3,000,000
Cynodon dactylon (Northern Tanzania) Whole plant 5,000,000 Cynodon dactylon Thrace Whole plant 2,100,000
Arundo donax Mt Carmel Stem 500,000
Arundo donax Mt Carmel Sheath 2,000,000 Arundo sp. Boeotia Stem/sheath 1,500,000
Lens culinaris Mt Carmel Whole plant 1000 Lens culinaris Boeotia Whole plant 200
Salix acmophylla Mt Carmel Wood 5 Salix sp. Thrace Wood 0
Salix acmophylla Mt Carmel bark 1000 Salix sp. Thrace Bark 700
Salix acmophylla Mt Carmel Leaves 9000 Salix sp. Thrace Leaves 2000
Pistacia palaestina Mt Carmel Wood 4 Pistacia terebinthus Thrace Wood 1
Pistacia palaestina Mt Carmel Bark 71 Pistacia terebibthus Thrace Bark 800
Pistacia palaestina Mt Carmel Leaves 11,000 Pistacia terebinthus Thrace Leaves 11,000
Crataegus aronia Mt Carmel Wood 7 Crataegus sp. Thrace Wood 2
Crataegus aronia Mt Carmel Leaves 2000 Crataegus sp. Thrace Leaves 6000
Olea europaea Mt Carmel Wood 11 Olea europaea Thrace Wood 160
Olea europaea Mt Carmel Bark 54 Olea europaea Thrace Bark 200
Olea europaea Mt Carmel Leaves 1000 Olea europaea Thrace Leaves 40,000
Olea europaea Mt Carmel Fruits 8 Olea europaea Thrace Fruits (pulp) 20
Amygdalus communis Mt Carmel Wood 9 Prunus amygdalus Thrace Wood 100
Amygdalus communis Mt Carmel Bark 2000 Prunus amygdalus Thrace Bark 800
Amygdalus communis Mt Carmel Leaves 3000 Prunus amygdalus Thrace Leaves 60,000
Ephedra foemina Mt Carmel Branches 11,000 Ephedra altissima Thrace Whole plant 11,000
Ficus Carica Mt Carmel Leaves 1,000,000 Ficus carica Thrace Leaves 1,000,000
Ficus Carica Mt Carmel Fruits 100,000 Ficus carica Thrace Fruits 100,000
Quercus calliprinos Mt Carmel Wood 23 Quercus frainetto Thrace Wood 35
Quercus calliprinos Mt Carmel Bark 9000 Quercus frainetto Thrace Bark 3000
Quercus calliprinos Mt Carmel Leaves 11,000 Quercus frainetto Thrace Leaves 200,000
Quercus ithaburensis Mt Carmel Wood 100 Quercus pubescence Thrace Wood 3
Quercus ithaburensis Mt Carmel Bark 600 Quercus pubescence Thrace Bark 5000
Quercus ithaburensis Mt Carmel Leaves 12,000 Quercus pubescence Thrace Leaves 1,000,000
Quercus ithaburensis Mt Carmel Husk acorn 20 Quercus pubescence Thrace Acorns 300,000
Quercus ithaburensis Mt Carmel Caps 6000 Quercus coccifera Thrace Wood 9
Quercus coccifera Thrace Bark 200
Quercus coccifera Thrace Leaves 20,000
Quercus coccifera Thrace Acorns 2000
Quercus petraea Thrace Wood 100
Quercus petraea Thrace Bark 5000
Quercus petraea Thrace Leaves 500,000
The list contains the plants that belong to at least the same genus and the specic parts of the plants analysed, showing the number of phytoliths per gram dry
material.
1267 G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275
Fig. 2. Number of phytoliths per gram dry weight of all the plants analysed from Greece; (a) wood, (b) bark, (c) edible parts of dicots (fruits, nuts, legumes), (d)
leaves and owers of trees and herbaceous plants and (e) monocotyledons.
1268 G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275
the phytoliths produced by the leaves of Ficus carica are hair
phytoliths. The fruit of Ficus carica also produces hair phyto-
liths (62%). Hair phytoliths though have little taxonomic value
since they are produced in high amounts by both dicots and
monocots. Although there are studies suggesting that hairs
can be diagnostic (Theobald et al., 1979) especially for mono-
cots (Kaplan et al., 1992), this collection shows that hair phy-
toliths generally have a limited taxonomic value. On the other
hand, prickles can usually be assigned to grasses and hairs
attached to their bases can be classied to either dicotyledons
or monocotyledons. Nevertheless the hair phytoliths of Ficus
carica, Celtis australis and Urtica sp. in this collection are
similar and therefore cannot be used for plant identication.
Generally, the leaves of trees and shrubs produce phytoliths
characteristic of dicotyledons (mainly polyhedral phytoliths
and tracheary elements). One exception is the leaves of Juni-
perus oxycedrus (a conifer) that do not produce polyhedral
phytoliths, but a high proportion of cylindroids (76%). This
is a characteristic also noted in the conifers (Pinus halepensis
and Cupressus sempervirens) of the Mt. Carmel collection
(Albert, 2000).
A phytolith type of much interest is the jigsaw puzzle
phytolith. Bozarth (1992) and Geis (1973) suggested that
this type is produced by deciduous trees growing in humid
conditions. Albert (2000) did not nd this type of phytolith
in the Mt. Carmel collection. Cells of this shape were ob-
served in a thick section of leaves of Quercus calliprinos,
but they were not silicied. Quercus calliprinos is an ever-
green plant and Albert suggested that this cell type silicies
in non-deciduous trees over a long period of time. In con-
trast, jigsaw puzzle phytoliths are present in several decidu-
ous plants in this collection and especially in Sorbus
torminalis. The humid climate and high precipitation of the
Rhodope mountains thus seems to be the reason for their
formation. The different climatic conditions of the geo-
graphic areas in which the two collections were made, could
account for this difference. In this collection jigsaw puzzle
phytoliths were also found in several legumes and shrubs,
Lens culinaris (12%), Thymus capitatus (16%) and Pteridium
aquilinum (37%). It is interesting to note that the spodogram
of Vicia sativa showed the presence of this type of phytolith.
This plant was collected from a village in central Greece
(Boeotia). The climate in this region is not humid and the
bedrock is mainly carbonate. The Vicia sativa specimens an-
alysed were from an irrigated cultivated eld and are used
for animal fodder. Hence wet conditions exist and these
may indeed be responsible for jigsaw puzzle phytolith for-
mation. Thus the presence of this type of phytolith could in-
dicate either wet forest conditions around an archaeological
site or the use of irrigation.
Finally, the ower phytoliths of trees and shrubs cannot be
differentiated from those in the leaves. The fact that both
owers and leaves are associated with the small branches of
trees can provide information on the part of the tree (i.e.,
thin versus thick branches) used. The only exception is the in-
orescence of Vitex agnus castus (chaste tree) that produces
a specic phytolith type (thin platelets with stripes).
3.4. Monocotyledons
Monocotyledons are, as expected, the major phytolith
producers in this region. Almost all species analysed produce
over half a million phytoliths/gram. The stems of four species
(Avena sativa, Briza maxima, Phleum sp. and Secale cereale)
(Fig. 2e) produce no more than 700,000 phytoliths /g of dry
plant material. The largest amounts of phytoliths are present
in wild grasses (Polypogon monspeliensis, Phleum sp., Lagu-
rus ovatus, Stipa bromoides, Bromus sp.) that produce from
6 to 12 million phytoliths/g. The cereals that produce the larg-
est amounts of phytoliths are Triticum aestivum and Secale
cereale. Their inorescences contain approximately 4 million
phytoliths/g of dry plant.
Another archaeologically important parameter is the rela-
tive amounts of phytoliths in the inorescence of grasses
compared to the culm/leaves fraction of the same plant. Ino-
rescence produces more phytoliths than culms and leaves
(Piperno, 1988). In general, we observed the same pattern,
but with exceptions (Hordeum vulgare and Aegilops sp.)
(Fig. 3). We also noted that in some plants (Polypogon mon-
speliensis, Secale cereale, Arundo sp., Stipa bromoides,
Phleum sp.) the differences between the inorescence and
culm/leaves phytolith production are very large.
In general, the morphological classication of grass phyto-
liths in this collection is in agreement with published data. The
dominant types are the short-cells, long-cells, and groupings of
these within articulated phytoliths (silica skeletons), as was
noted also by Albert (2000) and Rosen (1992). The short cells
are distributed between the following sub-families according
to the classication by Twiss et al. (1969); rondels are pro-
duced by C3 festucoid grasses (cereals, Bromus sp., Melica
sp., etc.), saddles are produced by C4 chloridoid grasses (Cyn-
odon dactylon), bilobes by panicoid and arundinoid grasses
(Zea mays, Arundo sp.) and crosses are present in the only
panicoid grass of this collection (Zea mays).
There are however some common morphotypes, such as
crenates (trapeziforms sinuates and polylobates) that are
mainly produced in festucoid grasses but have been also found
in Cynodon dactylon (in lower amounts though, only 3%).
It is worth noting the predominance of C3 grasses in the
Rhodope mountains. This is consistent with the cool wet
conditions that are the favourable regions for these grasses
(Abrantes, 2003).
The inorescence of wild grasses and cereals has some
different phytolith morphological characteristics (Rosen,
1992). As far as silica skeletons from husks are concerned,
Rosen (1992) noticed that the size and width of the wave of
the long cells walls, as well as the frequency and the attributes
of the papillae and the shape of the cork cells, can be used to
identify a specic grass genus or even species. This is consis-
tent with observations made using this collection. The attri-
butes Rosen described for Triticum as well as Hordeum and
Avena are identical with those observed in the same species
in this collection, and they are distinct from the wild species.
Rye (Secale cereale), a cereal not included in her analysis,
shows great similarities with barley; thick cell walls, squarish
1269 G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275
waves, with even amplitude. The papillae though are slightly
different. They are bigger than barleys and domed in shape,
they have more than 12 pits on their periphery (barley has
7e9) and the cork cell is kidney-shaped but more elongated
and with round endings. It is obvious that for identifying bar-
ley or rye, a well-silicied and complete silica skeleton from
the plants husk is necessary.
The wild grasses differ a lot from the cereals. They showed
a narrow and erratic wave of the cell wall, high frequency of
small sized papillae, and many pits on their periphery (more
than 14). These attributes characterize all the wild species
analysed.
On the other hand stems produce long cell articulated silica
skeletons with psilate margins that are characteristic of stems,
but cannot be used to differentiate wild species from cereals.
Although Rosen (1992) has shown some special attributes of
cereal stem silica skeletons, this collection shows that both
wild grasses and cereals share these attributes and therefore
these attributes cannot be diagnostic. Furthermore, the stems
of many of the wild grasses and cereals produce relatively
few phytoliths compared to the inorescences.
3.5. Proportions of variable to consistent
morphology phytoliths
The variable to consistent morphology (v/c) ratio, together
with the information on phytolith production per gram dry
plant, can be indicative of the overall type of plant compo-
nent introduced into the site (Albert et al., 2000). Fig. 4
shows a plot of the average ratios obtained for different cat-
egories of plants. Grasses produce almost no variable mor-
phology phytoliths, and consequently have an average ratio
of 0.02. The average ratio of dicot leaves and inorescence
is also low (0.16 and 0.15 respectively). The herbaceous
plants including legumes have a higher ratio (average 0.6),
and wood and bark have the highest ratios (wood, 1.02;
wood and bark together, 0.95). Fruits also have a high ratio
(0.95).
3.6. Contamination
While studying the different parts of the plants we noted
that phytoliths from one plant type can contaminate other
plants. The fact that these phytoliths are intrusive and not an
integral part of the parent cell morphology was conrmed
by spodograms, i.e., a chemical preparation procedure
whereby the cytoplasmic content of plant material is removed
and epidermal layers are detached in order to observe the plant
epidermis with phytoliths in situ (Kaplan et al., 1992). Albert
(2000) adopted a similar approach using thin sections of the
inner bark of her tree collection to differentiate between
phytoliths formed in the bark from those derived from contam-
ination. Spodograms prepared for dicotyledonous leaves and
monocotyledons (Figs. 5, 6) clearly show which phytoliths
form in situ, such as the long cells in monocotyledons and
the polyhedral and tracheids in dicot leaves.
Fig. 7 is a plot of the average number of contaminating
phytoliths in each plant category. In estimating these values
we excluded the species for which we counted less than 50
consistent morphology phytoliths due to the lack of reliability
of the results. We noted that grasses show no contamination
from other plant material. In contrast, wood/bark shows the
highest contamination (49% are phytoliths characteristic of
grasses and dicot leaves), while dicot owers and leaves
have the lowest (0.5% and 2% respectively). Albert et al.
(2003) found that the contamination of wood/bark by grass
phytoliths was around 30%.
3.7. Comparison with the Albert and Weiner
(2001) reference collection
Table 2 shows the numbers of phytoliths per gram where
the same part of the plant from the same species or at least ge-
nus was analysed in the reference collection from Mt. Carmel
and the adjacent coastal area in Israel and the reference collec-
tion from central and northern Greece. We note that the num-
bers of phytoliths per gram of the woody taxa are similar in
Fig. 3. Amounts of phytoliths per gram dry weight in the inorescence and culm/leaves of grasses analysed from Greece.
1270 G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275
both collections even when they belong to different species.
Dicot leaves are also similar, except for Olea europaea and
Amygdalus communis, where small differences are observed.
Quercus species show bigger differences, but we should bear
in mind that different species are compared. Finally, grasses
also produce similar amounts of phytoliths in both regions
for most of the species analysed (Avena sterilis inorescence,
Cynodon dactylon whole plant (from Tanzania), Hordeum vul-
gare inorescence and Arundo donax stem and sheath), with
the only major exception being the Lagurus ovatus inores-
cence. We also note that within both collections where differ-
ent species of the same genus were analysed, there are also
such signicant differences.
As far as the same species are concerned, in only 4 out of
15 comparisons are there signicant differences. Based on this
rather small set of data, together with the more general obser-
vations above, we tentatively conclude that the major control
over the number of phytoliths per unit weight is genetic.
This is in agreement with Piperno (1988), Hodson et al.
(2005) and Piperno et al. (2002). The latter have shown the
phytolith formation in Curcubita fruits is genetically deter-
mined and is not affected by environmental variability. Zheng
et al. (2003) have made similar observations.
It is however possible that similar plant taxa from very
different geographical regions will differ both with respect
to amounts and types of phytoliths produced. Bamford et al.
(2006) noticed that the plants from East Africa produced
many more phytoliths than the same genera from Israel.
This may be due to the higher availability of silica in the
East African soils (Albert et al., 2006; Bamford et al.,
2006). It is also true when comparing silica yields in the leaves
of trees in Greece to those from the north central region of the
USA (Geis, 1973) (same genera but different species). The
absolute silica yields in the North American trees are consis-
tently lower than those analysed in the Rhodope Mountains,
despite the fact that the annual precipitation in the two regions
is about the same (800e1200 mm vs 700e1000 mm in the
Rhodope Mountains). The reason for this is not known but it
might also relate to the bedrock chemistry. Finally, Rosen
and Weiner (1994) noted large variations in the yields of phy-
toliths from wheat grown in northern Europe versus Israel, and
within Israel under different conditions. The signicance of
these observations is equivocal, as both the amount of rainfall
and the extent of irrigation vary.
In general the phytolith morphotypes in each plant or plant
part are similar in both collections. We note that in both
regions spheroids (globulars) are the predominant phytoliths
in wood and bark, as are polyhedrals and tracheids from dicot
leaves (e.g., in the Israel collection Prunus amygdalus pro-
duces 88% polyhedrals and tracheary elements and in the
Greek collection 72%). Lens culinaris on the other hand pro-
duces 65% tracheary elements in the Greek collection and
only 29% tracheary elements and 61% cylindroids in the Israel
collection. We noted above the presence of jigsaw puzzle phy-
toliths in Lens culinaris in the Greek collection (12%) and
their absence in the Israel collection. Grasses also show a sim-
ilar distribution of morphotypes. For example, in the inores-
cence of Avena sterilis short cells, long cells and silica
skeleton long cells are abundant and present in roughly the
same proportions in both collections. In the Greek collection
they constitute 39%, 25% and 25% respectively and in the
Fig. 4. Average ratios of variable to consistent morphology (v/c) phytoliths in the different plant categories analysed from Greece.
Fig. 5. Spodogram from the leaf of Pistacia terebinthus showing polyhedral
silica skeletons, mesophyl silica skeletons and tracheary elements.
1271 G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275
Israel collection 37%, 7% and 31% respectively. Similar distri-
butions are present in Cynodon dactylon; the short cells and
long cells prevail and constitute 48% and 24% respectively
in the Greek collection and 42% and 19% respectively in the
East African collection. We note though that the short cells
of Cynodon dactylon in the Greek collection are mostly of
the chloridoid type, whereas in the African collection they
are of the festucoid types.
Finally, a comparison of the v/c ratios obtained in this study
to those reported by Albert et al. (1999) shows that dicot
leaves and especially grasses have similar ratios. In Albert
et al.s collection the average is 0.22 and 0.09 respectively
and in this collection they are 0.16 and 0.02. Major differences
are present in fruits (Alberts average 2.18, while in this
collection it is 0.95) and especially in wood/bark (4.97 in
her collection and 0.95 in this one). The lower wood/bark ratio
obtained in this study can be attributed to the higher extent of
contamination (40e50% in this study vs about 30% in Albert
et al.s (2003) study).
4. General discussion
A quantitative reference collection of phytoliths from
modern plants is the basis for documenting which plants leave
their traces in the form of siliceous phytoliths in the sediments
of an archaeological site and to what extent. The quantitative
results presented here show that different plant species and
plant parts have vastly different phytolith concentrations and
hence their potential contributions vary enormously. This
needs to be taken into account when decoding the archaeolog-
ical or geological phytolith record. The following are some
additional strengths and weaknesses of the archaeological
phytolith record.
4.1. Plants that produce few phytoliths
The woody species are the most conspicuous and archaeo-
logically important group of low phytolith producers. This
broad survey of phytoliths in wood and bark conrms that
wood from many species does not contain any phytoliths,
and in those species that do contain phytoliths, the concentra-
tions are very low. In contrast, almost all the bark samples
analysed do contain phytoliths, although also in relatively
small concentrations. Furthermore a signicant proportion of
the bark phytoliths are clearly contaminants. An added prob-
lem is that many of the phytoliths produced by woody species
have irregular or variable morphologies. Using only phytoliths
to reconstruct the use of wood and bark by humans is clearly
fraught with difculties.
These severe weaknesses are offset to a certain extent by
the fact that many species of trees also produce a different
form of mineralized material mainly in their bark. This is an
aggregate of soil minerals embedded in a biologically pro-
duced matrix of silica that unlike phytoliths also contains
abundant ions such as iron, aluminium and potassium. These
were originally called silica aggregates by Sangster and
Parry (1981), but because the silica is so impure, Schiegl
et al. (1994) referred to them as siliceous aggregates. Sili-
ceous aggregates, like phytoliths, are relatively insoluble and
thus often persist in the archaeological record. Their presence
together with phytoliths characteristic of wood and bark (e.g.,
a high proportion of variable morphology phytoliths, spheroids
and ellipsoids) would be indicative of wood having been
brought to the site (Schiegl et al., 1994).
As wood is mainly used for fuel, the burning process can
also augment the phytolith record. If preservation conditions
are good, then the calcitic ash produced by burning of the
calcium oxalate minerals, which are usually abundant in
wood and bark, will also be preserved. Under poorer preserva-
tion conditions, the calcitic ash may convert into various more
insoluble phosphate minerals that can also be recognized as
having been derived from ash (Karkanas et al., 2000, 2002;
Schiegl et al., 1996). The refractive index of the phytoliths
changes as a result of exposure to high temperatures (Elbaum
et al., 2003). This too can be used to infer the presence of
wood fuel. Thus despite the fact that trees generally produce
few phytoliths, their unique characteristics and association
with siliceous aggregates, ash and charcoal, together with
the fact that often large quantities of wood were burnt, can
produce a reliable and informative record.
It has been noted that in certain prehistoric caves calcitic
ash is present, often in the form of hearths, but phytoliths
and siliceous aggregates indicative of wood and bark are
Fig. 7. Average percentages of contamination in each plant category analysed
from Greece.
Fig. 6. Spodogram from the stem of Juncus sp., showing rod and long cell
silica skeletons and stomata.
1272 G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275
unidentiable (Schiegl et al., 1996). This could be due to an
alkaline environment, such as is produced by calcite. At
high pH, phytoliths are less stable and may have dissolved
(Schiegl et al., 1996, 2004). Another possible reason for the
absence of wood-derived phytoliths is that the wood used
did not have associated bark, and as many species of trees pro-
duce wood without phytoliths or with very few phytoliths, no
record remains. Bark is often more susceptible to degradation,
and thus dry wood collected for fuel may be without its bark.
G.T. was informed by people living in northern Greece that
wood collected for fuel is often debarked in order to reduce
the weight of the load carried back to the village. Wood
used for construction is also debarked. Thus there are circum-
stances under which wood may not be represented in the
archaeological phytolith record at all, despite the fact that it
was commonly used.
When thin tree branches with proportionately more bark
and leaves were used as fuel or for other purposes (e.g., roof-
ing, wall constructions), it may be possible to detect this
human behaviour in the archaeological record, since bark
does have some characteristic phytoliths and leaves of most
plants do produce high amounts of characteristic phytoliths.
This was recognized in the Middle Paleolithic record of Hay-
onim Cave, but not in the overlying Natuan layers (Albert
et al., 2003).
Among the low phytolith producing plant parts are the
owers of trees and shrubs, apart from chaste tree (Vitex agnus
castus). This limits the possibility of obtaining information on
the season of occupation of a site. Fruits also cannot be used
in such a way since they produce no characteristic phytoliths.
Nevertheless, the presence of several indicators that could be at-
tributed to fruits (high ratio of variable to consistent morphology
phytoliths, high number of spheroids together with polyhedral
phytoliths) may in certain contexts be helpful for identifying
fruit despite the absence of characteristic phytoliths.
Fruits and wood/bark share similar characteristics; they both
showlowphytolith production, they have a high ratio of variable
to consistent morphology phytoliths and among the latter the
prevailing ones in both are the spheroids. Nevertheless since
fruits and wood/bark are used in very different ways, i.e.,
wood is expected to be found in hearths while fruits are usually
stored in vessels or other containers, it is unlikely that their
similarities will be misleading in the archaeological record.
Finally, among the monocotyledons analysed, only the
grains of Triticum, as is known (Lentfer et al., 1997), have al-
most no phytoliths. The lack of phytoliths in grains devoid of
glumes in general implies that cereal phytoliths will not be
found in large numbers in places where the grains were
cleaned well. Nevertheless often in antiquity, grains were
stored in their husks to reduce mildew and predation by
bugs. The presence of cereal husks should leave tangible proof
of their presence.
4.2. Plants that produce many phytoliths
The main use of phytoliths preserved in the archaeological
record to date is for identifying the use of specic plant taxa.
This is based on identifying phytoliths with morphologies
characteristic of the specic taxon. This approach is most
applicable in plant groups that produce large amounts of
phytoliths at many different anatomical locations. The well-
known examples are among the grasses (Brown, 1984;
Piperno, 1988; Twiss et al., 1969). The phytolith input of
each part (husk or stem) of cereals or wild grasses, can be
used for interpreting the use of a room or understanding
past cultural practices, such as stabling, storing or grain pro-
cessing (Rosen, 2000, 2005). The large difference in the
phytolith concentrations between stems and husks of grasses
can also be exploited. The recovery of large amounts of
stem phytoliths and no husk phytoliths in a room, for exam-
ple, can indicate that it was used as a barn for storing fodder
or a stable. In areas where dung accumulated, large amounts
of phytoliths can be expected, provided that the animals were
fed the whole plant. If only the stems of certain cereals and
wild grasses that contain relatively few phytoliths were used
as fodder, this would deplete the phytolith record signi-
cantly. In addition dung from goats, animals that prefer leaves
in their diet, may also contain small amounts of phytoliths
given that many dicot leaves do not contain large amounts
of phytoliths. This was the case in Arene Candide cave, where
stabling activities were investigated through micromorpholog-
ical analysis. It was found that leaf foddering produced almost
phytolith-free sediments (Macphail et al., 1997). Apart from
phytoliths, the identication of dung of certain herbivors
can be complemented with the identication of dung spheru-
lites produced in the stomach of some herbivors mainly rumi-
nants and deposited in the soil together with the dung (Canti,
1997, 1998, 1999). The lamination of the sediments observed
through micromorphological analyses, together with the iden-
tication of phytoliths types and spherulites, can be used to-
gether for recognizing dung remains (Shahack-Gross et al.,
2005).
4.3. Phytoliths with unique properties
A potentially very interesting archaeological application
that arises from this study is the possible identication of
change of climate (wet conditions) or ancient irrigation based
on the presence of jigsaw puzzle phytoliths. These were
observed in leaves of trees and shrubs in Greece, but not in Is-
rael, and could be related to climatic differences. They are also
observed in irrigated annual legumes. Therefore, the presence
of jigsaw puzzle-shaped phytoliths in sites where a palaeocli-
matic study shows that arid conditions prevailed, could be at-
tributed to irrigation of legumes.
The presence of characteristic phytoliths of Vitex agnus
castus inorescence imply that Vitex branches were used in
an archaeological site. It may also imply a certain cultural
behaviour, since chaste tree branches have been used from
antiquity to the present for mats and basketry due to their ex-
ibility (Bostock and Riley, 1855; Pollunin, 1969). The name
Vitex comes from the Latin verb vieo, which means to bend,
or to weave. Homer and other ancient Greek writers refer often
to the Vitex branches as constructive material for mats, baskets
1273 G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275
and other structures and imprints of it have been detected in
Greek prehistoric sites (Beloyanni, 1995).
Finally the unique characteristics of cereal husks serve as
indicators of the staples (wheat, barley, oats and rye) used in
ancient settlements and may constitute the key to understand-
ing past agricultural economies. A study of this kind has been
conducted in an Indian Neolithic site aimed at understanding
agricultural production and utilization of its products (Harvey
and Fuller, 2004).
4.4. Reconstructing the amounts of plant
types used at a site
Taking into account the variable extents of phytolith pro-
duction in plants, Albert et al. (2000, 2003) used the Israeli
reference collection to assess the relative amounts of grasses,
leaves and wood/bark that were brought into Hayonim and
Kebara Caves. This was done using not only the known re-
lations between dry plant weights and phytolith amounts,
but also by distributing the phytolith morphotypes that are
present in two or more of these groups according to their
known distributions in the modern reference collection.
Thus for example, the presence of one aliquot of ash of grass
phytoliths is roughly equivalent to the presence of 20 ali-
quots of ash of wood/bark phytoliths, based on their refer-
ence collection (Albert et al., 2000). In this more extensive
reference collection the relative proportions of phytoliths
from ash of grass compared to ash of wood/bark phytoliths
is around 40. Information on the relative amounts of grasses,
leaves and wood/bark used by the occupants of a given site,
can certainly shed light on human behaviour patterns partic-
ularly in the prehistoric periods.
4.5. Using phytoliths for regional studies
The comparison of the two reference collections from the
Mediterranean region shows that in general the amounts of
phytoliths produced by different plant taxa, are genetically
controlled. The inuence of climate can be detected, but
appears in this case to be secondary. This means that one quan-
titative reference collection may be useful for the entire
Mediterranean region. We thus propose that such regional
reference collections be produced. This would greatly facili-
tate phytolith studies in this archaeologically important region.
5. Conclusions
The quantitative and morphological analysis of phytoliths
from modern plants highlights some of the strengths and
weaknesses when decoding the archaeological phytolith re-
cord. In general, the phytolith record is the most robust and
informative source of information available on the botanical
archaeological record. This information is much enhanced
by the availability of a reference collection of modern plants
that includes information on the morphotypes as well as the
numbers of phytoliths per unit weight of the plant.
Acknowledgements
We thank Ali Sinap for his invaluable information on the
use of plants by the inhabitants of Rhodope mountain. We
also thank Ofer Bar-Yosef for his help with the initial sample
collection, as well as Gordon Hillman and Dorian Fuller for
their help in the identication of Hynaldia villosa and Stipa
bromoides. We thank Mr George Schwartzmann, Sarasota,
Florida, for his generous nancial support. We also thank
the Kimmel Center for Archaeological Science for their nan-
cial support. S.W. is the incumbent of the Dr Trude Burchardt
professorial chair of Structural Biology. G.T. conducted this
study during her fellowship in environmental studies by the
Wiener Laboratory at the American School of Classical
Studies at Athens. Part of this study was supported by the Brit-
ish School at Athens.
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