Journal of Archaeological Science 34 (2007) 1262e1275

http://www.elsevier.com/locate/jas

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. Considerable 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 significant amounts of phytoliths and grasses are prolific 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

* 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

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 quantities, 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

G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275

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 quantities produced per unit weight of plant material. It is also
necessary to differentiate between phytolith shapes that are
unique to a specific 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 introduced 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 reference collection may also reveal the use of a particular area of
the site for a specific 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 Natufian periods. Here we report the
results of a second, more comprehensive phytolith reference
collection mostly from northern Greece. These results, together 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 differentiating 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 exploitation of such plant types is often indicative of a particular
behaviour, such as fuel used for fires and fodder collected for
animals. There are however several inherent problems with
this approach. One is that the same phytolith types are sometimes 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 categories. See Albert et al. (2003) for more details. Another
problem is that many phytoliths have variable morphologies,
with irregular and not identifiable 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

1263

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 general plant categories such as grasses, dicots, cereals and legumes. Phytolith production is however also influenced 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 Sarakini, 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 environment and it is not an exhaustive analysis of the plants of northern 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 significance. 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 monocotyledons. 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 (gymnosperm), 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., Juglans 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 officinalis, 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

G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275

1264

Fig. 1. Map of Greece. The areas from which the plants of the reference collection have been collected are shown (Thrace, Argolid, Boeotia).

a specific plant part to the phytolith record. This is sometimes

most helpful, since different parts of plants may reflect various
human or animal habits and activities, including seasonal activities. Most of the grasses and all the cereals were separated
into two parts: inflorescence (including the grains) and the
leaves/stems together. Tree species were mechanically separated into five parts whenever possible: wood, bark, leaves,
flowers and fruits. Herbaceous plants were analyzed whole
and thus include stem, leaves and often the flowers. 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 consists 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 publication). Table 2 lists all the species analysed.
2.1. Phytolith extraction
The plants were cleaned extensively by washing in deionized 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 furnace at 500  for 4 h. The ash was treated with 1 N HCl
and centrifuged (3500 rpm for 10 min), resuspended in deionized 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 reflects also the yield of silica
(opal) in the plants. The AIF fractions were then dried under 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, anhydrite, 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 purified by centrifuging in 2.4 g/ml density heavy liquid (sodium polytungstate: Na6 (H2W12O40)$H2O). 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
filled with water for the final centrifugation. In so doing
the denser minerals were separated from the less dense
silica in four stages. The plants that were subject to the additional treatment included all the bark samples, and from
the wood samples Salix sp., Fagus sylvatica, Juglans regia,
Alnus glutinosa, Pistacia terebinthus, Quercus pubescens,

G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275

1265

Table 1
List of all the plants collected from Greece for phytolith analysis
Latin name

Family

English name

Provenance

Woody plants
Acer monospessulanum
Alnus glutinosa
Arbutus adrachne
Carpinus betulus
Celtis australis
Corylus avelana
Crataegus sp.
Fagus sylvatica
Ficus carica
Fraxinus ornus
Juglans regia
Juniperus oxycedrus
Olea europaea
Phillyrea latifolia
Pistacia terebinthus
Populus alba
Prunus amygdalus
Prunus avium
Pyrus amygdaliformis
Quercus coccifera
Quercus frainetto
Quercus petraea
Quercus pubescens
Salix sp.
Sambucus nigra
Sorbus torminalis
Tamarix sp.
Tilia sp.
Vitex agnus castus

Aceraceae
Betulaceae
Ericaceae
Betulaceae
Ulmaceae
Corylaceae
Rosaceae
Fagaceae
Moraceae
Oleaceae
Juglandaceae
Cupressaceae
Oleaceae
Oleaceae
Anacardiaceae
Salicaceae
Rosaceae
Rosaceae
Rosaceae
Fagaceae
Fagaceae
Fagaceae
Fagaceae
Salicaceae
Caprifoliaceae
Rosaceae
Tamaricaceae
Tiliaceae
Verbenaceae

Maple
Black alder
Strawberry tree
Hornbean
Nettle tree
Hazelnut
Hawthorn
Beech
Fig tree
Manna ash
Walnut
Juniper
Olive tree
Basswood
Terebinth
White poplar
Almond tree
Wild cherry
Wild pear
Kermes oak
Hungarian oak
Durmast oak
Downy oak
Willow
Common elder
Wild service tree
Tamarisk
Lime
Chaste tree

Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace

M
M
M
M
C
M
M
M
M
M
M
M
C
P
P
M
C
M
M
P
M
M
M
M
M
M
P
M
P

Herbaceous plants and shrubs

Cichorium sp.
Ephedra altissima
Euphorbia sp.
Nicotiana tabacum
Paliurus spina christi
Pteridium aquilinum
Rubus sanctus
Salvia officinalis
Thymus capitatus.
Urtica sp.
Vitis sylvestris

Cichoriaceae
Ephedraceae
Euphorbiaceae
Solanaceae
Rhamnaceae
Hypolepidaceae
Rosaceae
Labiatae
Labiatae
Urticaceae
Vitaceae

Chicory
Ephedra
Spurge
Tobacco
Christs thorn
Fern
Blackberry
Sage
Thyme
Stinging nettle
Grape-vine

Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace

M
C
M
M
P
M
M
M
M
M
M

Domesticated annual legumes

Cicer arietinum
Lens culinaris
Vicia faba

Leguminoseae
Leguminoseae
Leguminoseae

Chickpea
Lentil
Faba bean

Boeotia P
Boeotia P
Boeotia P

Monocotyledons
Aegilops sp.
Arundo sp.
Briza maxima
Bromus sp.
Cynodon dactylon
Dactylis glomerata
Hynaldia villosa
Lagurus ovatus
Lolium sp.
Melica sp.
Phleum sp.
Polypogon monspeliensis
Stipa bromoides
Juncus sp.

Gramineae
Gramineae
Gramineae
Gramineae
Gramineae
Gramineae
Gramineae
Gramineae
Gramineae
Gramineae
Gramineae
Gramineae
Gramineae
Juncaceae

Goat grass (wild wheat)

Reed
Quaking grass
Brome
Bermuda grass
Orchard grass

Thrace M
Boeotia P
Argolid P
Thrace M
Thrace M
Thrace M
Argolid P
Attika P
Thrace M
Thrace M
Thrace M
Thrace M
Thrace M
Argolid P

Hare tail
Ryegrass
Melick grass
Cats tail
Beard grass
Needle grass
Rush

(continued on next page)

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G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275

Table 1 (continued )
Latin name

Family

English name

Provenance

Domesticated cereals
Avena sativa
Hordeum vulgare
Secale cereale
Triticum aestivum
Zea mays

Gramineae
Gramineae
Gramineae
Gramineae
Gramineae

Oats
Barley
Rye
Bread wheat
Corn

Thrace M
Boeotia P
Thrace M
Thrace M
Thrace M

The provenance is shown: M, mountain; C, coastal plain; P, plain at the foot of the mountain.

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 further purified: Prunus avium, Rubus sp., Salix sp., Sambucus
nigra, Pyrus amygdaliformis, Tamarix sp., Crataegus sp., Vitis sylvestris, Vicia faba, Lens cullinaris, Salvia officinalis
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 phytoliths 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 ethnographic 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 silicification 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 region 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 classification

The phytoliths were classified morphologically according
to their anatomical origin and when this was not possible, according 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 unidentifiable morphologies. Phytoliths with
consistent morphologies were classified and their relative proportions determined in percent. The terminology used for consistent morphology phytolith classification 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 contributions of different plants as well as different parts of plants to
the phytolith record, vary enormously. By knowing the phytolith 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 archaeological site can be inferred. This is certainly of much archaeological 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 phytoliths 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
identified with wood and bark of specific taxa. There are however 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

G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275

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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
Avena sterilis
Avena sterilis
Lagurus ovatus
Lagurus ovatus
Lagurus ovatus
Hordeum vulgare
Hordeum vulgare
Triticum aestivum
Triticum aestivum
Cynodon dactylon
Arundo donax
Arundo donax
Lens culinaris
Salix acmophylla
Salix acmophylla
Salix acmophylla
Pistacia palaestina
Pistacia palaestina
Pistacia palaestina
Crataegus aronia
Crataegus aronia
Olea europaea
Olea europaea
Olea europaea
Olea europaea
Amygdalus communis
Amygdalus communis
Amygdalus communis
Ephedra foemina
Ficus Carica
Ficus Carica
Quercus calliprinos
Quercus calliprinos
Quercus calliprinos
Quercus ithaburensis
Quercus ithaburensis
Quercus ithaburensis
Quercus ithaburensis
Quercus ithaburensis

Tel-Dor
Tel-Dor
Tel-Dor
Tel-Dor
Tel-Dor
Tel-Dor
Tel-Dor
Tel-Dor
Tel-Dor
Mt Carmel
(Northern Tanzania)
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel
Mt Carmel

Leaves
Stem
Inflorescence
Leaves
Stem
Inflorescence
Whole plant
Inflorescence
Inflorescence
Whole plant
Whole plant
Stem
Sheath
Whole plant
Wood
bark
Leaves
Wood
Bark
Leaves
Wood
Leaves
Wood
Bark
Leaves
Fruits
Wood
Bark
Leaves
Branches
Leaves
Fruits
Wood
Bark
Leaves
Wood
Bark
Leaves
Husk acorn
Caps

7,000,000
1,000,000
2,000,000
1,000,000
40,000
400,000
700,000
400,000
400,000
1,500,000
5,000,000
500,000
2,000,000
1000
5
1000
9000
4
71
11,000
7
2000
11
54
1000
8
9
2000
3000
11,000
1,000,000
100,000
23
9000
11,000
100
600
12,000
20
6000

Avena sterilis

Thrace

Culm/leaves

Avena sterilis
Lagurus ovatus

Thrace
Attika

Inflorescence
Culm/leaves

2,000,000
7,000,000

Lagurus ovatus
Hordeum vulgare
Hordeum vulgare
Triticum aestivum
Triticum aestivum
Cynodon dactylon

Attika
Boeotia
Boeotia
Thrace
Thrace
Thrace

Inflorescence
Culm/leaves
Inflorescence
Husk
Culm/leaves
Whole plant

7,600,000
1,500,000
1,000,000
4,000,000
3,000,000
2,100,000

Arundo sp.
Lens culinaris
Salix sp.
Salix sp.
Salix sp.
Pistacia terebinthus
Pistacia terebibthus
Pistacia terebinthus
Crataegus sp.
Crataegus sp.
Olea europaea
Olea europaea
Olea europaea
Olea europaea
Prunus amygdalus
Prunus amygdalus
Prunus amygdalus
Ephedra altissima
Ficus carica
Ficus carica
Quercus frainetto
Quercus frainetto
Quercus frainetto
Quercus pubescence
Quercus pubescence
Quercus pubescence
Quercus pubescence
Quercus coccifera
Quercus coccifera
Quercus coccifera
Quercus coccifera
Quercus petraea
Quercus petraea
Quercus petraea

Boeotia
Boeotia
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace
Thrace

Stem/sheath
Whole plant
Wood
Bark
Leaves
Wood
Bark
Leaves
Wood
Leaves
Wood
Bark
Leaves
Fruits (pulp)
Wood
Bark
Leaves
Whole plant
Leaves
Fruits
Wood
Bark
Leaves
Wood
Bark
Leaves
Acorns
Wood
Bark
Leaves
Acorns
Wood
Bark
Leaves

1,500,000
200
0
700
2000
1
800
11,000
2
6000
160
200
40,000
20
100
800
60,000
11,000
1,000,000
100,000
35
3000
200,000
3
5000
1,000,000
300,000
9
200
20,000
2000
100
5000
500,000

600,000

The list contains the plants that belong to at least the same genus and the specific parts of the plants analysed, showing the number of phytoliths per gram dry
material.

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 inflorescence of dicotyledonous
trees and shrubs
Fig. 2d shows that the leaves and flowers 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 reflected in their rich phytolith record (52%
and 63% hair phytoliths respectively). Note too that 94% of

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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 flowers of trees and herbaceous plants and (e) monocotyledons.

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 phytoliths (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 monocots (Kaplan et al., 1992), this collection shows that hair phytoliths 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 classified 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 identification.
Generally, the leaves of trees and shrubs produce phytoliths
characteristic of dicotyledons (mainly polyhedral phytoliths
and tracheary elements). One exception is the leaves of Juniperus 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 find this type of phytolith
in the Mt. Carmel collection. Cells of this shape were observed in a thick section of leaves of Quercus calliprinos,
but they were not silicified. Quercus calliprinos is an evergreen plant and Albert suggested that this cell type silicifies
in non-deciduous trees over a long period of time. In contrast, jigsaw puzzle phytoliths are present in several deciduous 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 geographic 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 analysed were from an irrigated cultivated field and are used
for animal fodder. Hence wet conditions exist and these
may indeed be responsible for jigsaw puzzle phytolith formation. Thus the presence of this type of phytolith could indicate either wet forest conditions around an archaeological
site or the use of irrigation.
Finally, the flower phytoliths of trees and shrubs cannot be
differentiated from those in the leaves. The fact that both
flowers 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 inflorescence of Vitex agnus castus (chaste tree) that produces
a specific phytolith type (thin platelets with stripes).

1269

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., Lagurus ovatus, Stipa bromoides, Bromus sp.) that produce from
6 to 12 million phytoliths/g. The cereals that produce the largest amounts of phytoliths are Triticum aestivum and Secale
cereale. Their inflorescences contain approximately 4 million
phytoliths/g of dry plant.
Another archaeologically important parameter is the relative amounts of phytoliths in the inflorescence of grasses
compared to the culm/leaves fraction of the same plant. Inflorescence 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 monspeliensis, Secale cereale, Arundo sp., Stipa bromoides,
Phleum sp.) the differences between the inflorescence and
culm/leaves phytolith production are very large.
In general, the morphological classification of grass phytoliths 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 classification by Twiss et al. (1969); rondels are produced by C3 festucoid grasses (cereals, Bromus sp., Melica
sp., etc.), saddles are produced by C4 chloridoid grasses (Cynodon 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 inflorescence 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 specific grass genus or even species. This is consistent with observations made using this collection. The attributes 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

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G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275

Fig. 3. Amounts of phytoliths per gram dry weight in the inflorescence and culm/leaves of grasses analysed from Greece.

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 barley or rye, a well-silicified 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 inflorescences.
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 component introduced into the site (Albert et al., 2000). Fig. 4
shows a plot of the average ratios obtained for different categories of plants. Grasses produce almost no variable morphology phytoliths, and consequently have an average ratio
of 0.02. The average ratio of dicot leaves and inflorescence
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 confirmed
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 contamination. 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 flowers 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 genus was analysed in the reference collection from Mt. Carmel
and the adjacent coastal area in Israel and the reference collection from central and northern Greece. We note that the numbers of phytoliths per gram of the woody taxa are similar in

G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275

1271

Fig. 4. Average ratios of variable to consistent morphology (v/c) phytoliths in the different plant categories analysed from Greece.

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 inflorescence,
Cynodon dactylon whole plant (from Tanzania), Hordeum vulgare inflorescence and Arundo donax stem and sheath), with
the only major exception being the Lagurus ovatus inflorescence. We also note that within both collections where different species of the same genus were analysed, there are also
such significant differences.
As far as the same species are concerned, in only 4 out of
15 comparisons are there significant differences. Based on this
rather small set of data, together with the more general observations 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 determined 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 consistently 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 phytoliths from wheat grown in northern Europe versus Israel, and
within Israel under different conditions. The significance 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 produces 88% polyhedrals and tracheary elements and in the
Greek collection 72%). Lens culinaris on the other hand produces 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 phytoliths in Lens culinaris in the Greek collection (12%) and
their absence in the Israel collection. Grasses also show a similar distribution of morphotypes. For example, in the inflorescence 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. 5. Spodogram from the leaf of Pistacia terebinthus showing polyhedral

silica skeletons, mesophyl silica skeletons and tracheary elements.

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G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275

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 archaeological or geological phytolith record. The following are some
additional strengths and weaknesses of the archaeological
phytolith record.
4.1. Plants that produce few phytoliths

Fig. 6. Spodogram from the stem of Juncus sp., showing rod and long cell
silica skeletons and stomata.

Israel collection 37%, 7% and 31% respectively. Similar distributions 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

Fig. 7. Average percentages of contamination in each plant category analysed

from Greece.

The woody species are the most conspicuous and archaeologically important group of low phytolith producers. This
broad survey of phytoliths in wood and bark confirms that
wood from many species does not contain any phytoliths,
and in those species that do contain phytoliths, the concentrations are very low. In contrast, almost all the bark samples
analysed do contain phytoliths, although also in relatively
small concentrations. Furthermore a significant proportion of
the bark phytoliths are clearly contaminants. An added problem 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 difficulties.
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 produced 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. Siliceous 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 preservation 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

G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275

unidentifiable (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 produce 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 circumstances 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., roofing, 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 Hayonim Cave, but not in the overlying Natufian layers (Albert
et al., 2003).
Among the low phytolith producing plant parts are the
flowers 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 attributed 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
show low phytolith 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 almost 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 specific plant taxa.

1273

This is based on identifying phytoliths with morphologies

characteristic of the specific taxon. This approach is most
applicable in plant groups that produce large amounts of
phytoliths at many different anatomical locations. The wellknown 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 processing (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 example, 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 significantly. 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 micromorphological analysis. It was found that leaf foddering produced almost
phytolith-free sediments (Macphail et al., 1997). Apart from
phytoliths, the identification of dung of certain herbivors
can be complemented with the identification of dung spherulites produced in the stomach of some herbivors mainly ruminants 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 identification of phytoliths types and spherulites, can be used together 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 identification 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 Israel, 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 palaeoclimatic study shows that arid conditions prevailed, could be attributed to irrigation of legumes.
The presence of characteristic phytoliths of Vitex agnus
castus inflorescence 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 flexibility (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

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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 understanding 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 production 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 relations 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 aliquots of ash of wood/bark phytoliths, based on their reference 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 particularly 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 influence of climate can be detected, but
appears in this case to be secondary. This means that one quantitative reference collection may be useful for the entire
Mediterranean region. We thus propose that such regional
reference collections be produced. This would greatly facilitate 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 record. 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 identification of Hynaldia villosa and Stipa
bromoides. We thank Mr George Schwartzmann, Sarasota,
Florida, for his generous financial support. We also thank
the Kimmel Center for Archaeological Science for their financial 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 British School at Athens.

References
Abrantes, F., 2003. A 340,000 year continental climate record from tropical
Africa e news from opal phytoliths from the equatorial Atlantis. Earth
and Planetary Science Letters 209, 165e179.
Albert, R.M., 2000. Study of ash layers through phytolith analyses from the
Middle Palaeolithic levels of Kebara and Tabun Caves. PhD thesis, University of Barcelona, Barcelona.
Albert, R.M., Tsatskin, A., Ronen, A., Lavi, O., Estroff, L., Lev-Yadun, S.,
Weiner, S., 1999. Mode of occupation of Tabun Cave, Mt. Carmel Israel,
during the Mousterian period: a study of the sediments and the phytoliths.
Journal of Archaeological Science 26, 1249e1260.
Albert, R.M., Bar-Yosef, O., Meignen, L., Weiner, S., 2000. Phytoliths in the
Middle Paleolithic deposits of Kebara cave, Mt. Carmel, Israel: study of
the plant materials used for fuel and other purposes. Journal of Archaeological Science 27, 931e947.
Albert, R.M., Weiner, S., 2001. Study of phytoliths in prehistoric ash layers
using a quantitative approach. In: Meunier, J.D., Coline, F. (Eds.), Phytoliths: Applications in Earth Sciences and Human History. A.A. Balkema
Publishers, Netherlands, pp. 251e266.
Albert, R.M., Bar-Yosef, O., Meignen, L., Weiner, S., 2003. Phytolith and mineralogical study of hearths from the Middle Paleolithic levels of Hayonim
Cave (Galilee, Israel). Journal of Archaeological Science 30, 461e480.
Albert, R.M., Bamford, M.K., Cabanes, D., 2006. Taphonomy of phytoliths
and macroplants in different soils from Olduvai Gorge (Tanzania) and
the application to Plio-Pleistocene palaeoanthropological samples. Quaternary International 148, 78e94.
Albert, R.M., Shahack-Gross, R., Cabanes, D., Gilboa, A., Lev-Yadun, S.,
Portillo, M., Sharon, I., Boaretto, E., Weiner, S. Phytolith rich layers
from the Late Bronze and iron Ages at Tel-Dor (Israel): mode of formation
and archaeological significance. Journal of Archaeological Science, submitted for publication.
Ball, T., Gardner, J.S., Brotherson, J.D., 1996. Identifying phytoliths produced
by the inflorescence bracts of three species of wheat (Triticum monococcum
L., T. dicoccum Schrank., and T. aestivum L.) using computer-assisted image
and statistical analyses. Journal of Archaeological Science 23, 619e632.
Ball, T.B., Gardner, J.S., Anderson, N., 1999. Identifying inflorescence phytoliths from selected species of wheat (Triticum monococcum, T. dicoccon, T.
dicoccoides, and T. aestivum) and barley (Hordeum vulgare and H. spontaneum). American Journal of Botany 86 (11), 1615e1623.
Bamford, M.K., Albert, R.M., Cabanes, D., 2006. Assessment of the Lowermost Bed II Plio-Pleistocene vegetation in the eastern palaeolake margin
of Olduvai Gorge (Tanzania) and preliminary results from macroplant
and phytoliths. Quaternary International 148, 95e112.
Beloyanni, M.P., 1995. Mat impressions on prehistoric pottery from Tsoungiza. Annals of Archaeology XXII 1989, 171e182 (in Greek).

G. Tsartsidou et al. / Journal of Archaeological Science 34 (2007) 1262e1275

Berlin, A., Ball, T., Thompson, R., Kittleson, D., Herbert, S.C., 2003. Ptolemaic agriculture, Syrian wheat, and Triticum aestivum. Journal of
Archaeological Science 30, 115e121.
Bostock, J., Riley, H.T. (Eds.), 1855. Pliny the Elder, the Natural History. Taylor and Francis, London. book 24, chapter 38.
Bozarth, S.R., 1992. Classification of phytoliths formed in selected dicotyledons native to the Great Plains. In: Rapp, G., Mulholand, S.C. (Eds.), Phytolith Systematics. Plenum Press, New York, pp. 193e214.
Brown, D.A., 1984. Prospects and limits of a phytolith key for grasses in the
Central United States. Journal of Archaeological Science 11, 345e368.
Canti, M., 1997. An investigation of microscopic calcareous spherulites from
herbivore dungs. Journal of Archaeological Science 24, 219e231.
Canti, M., 1998. The micromorphological identification of faecal spherulites
from archaeological and modern materials. Journal of Archaeological
Science 25, 434e444.
Canti, M., 1999. The production and preservation of faecal spherulites:
animals, environment, and taphonomy. Journal of Archaeological Science
26, 251e258.
Elbaum, R., Albert, R.M., Elbaum, M., Weiner, S., 2003. Detection of burning
of plant materials in the archaeological record by a change in the refractive
indices of siliceous phytoliths. Journal of Archaeological Science 30,
217e226.
Geis, J.W., 1973. Biogenic silica in selected species of deciduous angiosperms.
Soil Science 116, 113e130.
Harvey, E.L., Fuller, D.Q., 2004. Investigating crop processing using phytolith
analysis: the example of rice and millets. Journal of Archaeological
Science 32, 739e752.
Hodson, M.J., White, P.J., Mead, A., Broadley, M.R., 2005. Phylogenetic
variation in the silicon composition of plants. Annals of Botany 96,
1027e1046.
Jones, L.H.P., Handreck, K.A., 1965. Studies of silica in the oat plant III. Uptake of silica from soils by the plant. Plant and Soil 23, 79e96.
Jones, J.G., Bryant Jr., V.M., 1992. Phytolith taxonomy in selected species of
Texas cacti. In: Rapp, G., Mulholand, S.C. (Eds.), Phytolith Systematics.
Plenum Press, New York, pp. 215e238.
Kaplan, L., Smith, M.B., Sneddon, L.A., 1992. Cereal grain phytoliths of
Southwest Asia and Europe. In: Rapp, G., Mulholand, S.C. (Eds.), Phytolith Systematics. Plenum Press, New York, pp. 149e174.
Karkanas, P., Bar-Yosef, O., Goldberg, P., Weiner, S., 2000. Diagenesis in
prehistoric caves: the use of minerals that form in situ to assess the completeness of the archaeological record. Journal of Archaeological Science
23, 915e929.
Karkanas, P., Rigaud, J.P., Simek, J.F., Albert, R.M., Weiner, S., 2002. Ash,
bones and guano: a study of the minerals and phytoliths in the sediments
of Grotte XVI, Dordogne, France. Journal of Archaeological Science 29,
721e732.
Lentfer, C., Boyd, W.E., Gojak, D., 1997. Hope farm windmill: phytolith
analysis of cereals in Early Colonial Australia. Journal of Archaeological
Science 24, 841e856.
Macphail, R.I., Courty, M.A., Harther, J., Wattez, J., 1997. The soil micromorphological evidence of domestic occupation and stabling activities. In:
Maggi, R. (Ed.), Arene Candide: a Functional and Environmental Assessment of the Holocene Sequences Excavated by Bernado Brea (1940e
1950), vol. V. Memorie dell Istituto di Paleontologia, Umana, pp. 53e88.
Madella, M., Jones, M.K., Goldberg, P., Goren, Y., Hovers, E., 2002. The
exploitation of plant resources by Neanderthals in Amud cave (Israel):
the evidence from phytolith studies. Journal of Archaeological Science
29, 703e719.
Madella, M., Alexandre, A., Ball, T., 2005. International code for phytolith
nomenclature. Annals of Botany 96, 253e260.
Metcalfe, C.R., 1960. Anatomy of the Monocotyledons I. Oxford University
Press, London. Gramineae.
Ollendorf, A.L., Mulholand, S.C., Rapp, G., 1988. Phytolith analysis as
a means of plant identification: Arundo donax and Phragmites communis.
Annals of Botany 61, 209e214.

1275

Pearsall, D.M., Piperno, D.R., Dinan, E.H., Umlauf, M., Zhao, Z.,
Benefer, R.A., 1995. Distinguishing rice (Oryza sativa Poaceae) from
wild Oryza species through phytolith analysis: results of preliminary
research. Economic Botany 49, 183e196.
Piperno, D.R., 1984. A comparison and differentiation of phytoliths from
maize and wild grasses: use of morphological criteria. American Antiquity
49, 361e383.
Piperno, D.R., 1988. Phytolith Analysis: An Archaeological and Geological
Perspective. Academic Press, New York.
Piperno, D., Holst, I., Wessel-Beaver, L., Andres, T.C., 2002. Evidence for the
control of phytolith formation in Cucurbita fruits by the hard rind (Hr)
genetic locus: archaeological and ecological implications. Proceedings of
the National Academy of Science of the United States of America 99,
10923e10928.
Pollunin, O., 1969. Flowers of Europe, a Field Guide. Oxford University Press,
London.
Rosen, A.M., 1992. Preliminary identification of silica skeletons from Near
Eastern archaeological sites: an anatomical approach. In: Rapp, G.,
Mulholand, S.C. (Eds.), Phytolith Systematics. Plenum Press, New York,
pp. 129e148.
Rosen, A.M., 2000. Phytolith analysis in Near Eastern archaeology. In:
Pike, S., Gitin, S. (Eds.), The Practical Impact of Science on Near
Eastern and Aegean Archaeology. Archaetype Publications, London,
pp. 9e15.
Rosen, A.M., 2005. Phytolith indicators of plant and land use at C
atalhoyuk.
In: Hodder, I. (Ed.), Inhabiting C
atalhoyuk: Reports from the 1995e1999
Seasons. C
atalhoyuk Project, Vol. 4, Part A. McDonald Institute Monographs, Cambridge, pp. 203e212.
Rosen, A.M., Weiner, S., 1994. Identifying ancient irrigation: a new method
using opaline phytoliths from emmer wheat. Journal of Archaeological
Science 21, 132e135.
Sangster, A.G., Parry, D.W., 1981. Ultrastructure of silica deposits in
higher plants. In: Simpson, T.L., Volcani, B.E. (Eds.), Silicon and Siliceous Structures in Biological Systems. Springer-Verlag, New York,
pp. 383e407.
Schiegl, S., Lev-Yadun, S., Bar-Yosef, O., El Goresy, A., Weiner, S., 1994.
Siliceous aggregates from prehistoric wood ash: a major component of
sediments in Kebara and Hayonim caves (Israel). Israel Journal of Earth
Science 43, 267e278.
Schiegl, S., Goldberg, P., Bar-Yosef, O., Weiner, S., 1996. Ash deposits in
Hayonim and Kebara caves, Israel: macroscopic microscopic and mineralogical observations and their archaeological implications. Journal of
Archaeological Science 23, 763e781.
Schiegl, S., Stockhammer, P., Scott, C., Wadley, L., 2004. A mineralogical and
phytolith study of the Middle Stone Age hearths in Sibudu cave, KwazuluNatal South Africa. South African Journal of Science 100, 185e194.
Scott Cummings, L., 1992. Illustrated phytoliths from assorted food plants. In:
Rapp, G., Mulholand, S.C. (Eds.), Phytolith Systematics. Plenum Press,
New York, pp. 175e192.
Shahack-Gross, R., Albert, R.M., Gilboa, A., Nagar-Hilman, O., Sharon, I.,
Weiner, S., 2005. Geoarchaeology in an urban context: the uses of space
in a Phoenician monumental building at Tel Dor (Israel). Journal of
Archaeological Science 32, 1417e1431.
Theobald, W.L., Krahulik, J.L., Rollins, R.C., 1979. Trichome description and
classification. In: Metcalfe, C.R., Chalk, L. (Eds.)Anatomy of Dicotyledons, second ed., vol. 1. Clarendon Press, Oxford, pp. 40e53.
Twiss, P.C., Erwin, S., Smith, R.M., 1969. Morphological classification of
grass phytoliths. Soil Science Society of America Proceedings 33,
109e115.
Zhao, Z., Pearsall, D.M., Benefer, A.B., Piperno, D.R., 1998. Distinguishing
rice (Oryza sativa Poaceae) from wild Oryza species through phytolith
analysis, II: finalized method. Economic Botany 52, 134e135.
Zheng, Y., Dong, Y., Matsui, A., Udatsu, T., Fujiwara, H., 2003. Molecular
genetic basis of determining subspecies of ancient rice using the shape
of phytoliths. Journal of Archaeological Science 30, 1215e1221.