Biol. J. Linn. Soc., 3, pp. 303-312. With 2 plates and 1 figure
December 1971
Cotton seeds from the Neolithic in Egyptian Nubia and the
origin of Old World cotton
K. A. CHOWDHURY
AND
G. M. BUTH
Department of Botany, Aligarh Muslim University, Aligarh, India
Accepted for publication M a y 1971
During an excavation at Afyeh, Egyptian Nubia, in 1962, the Archaeological Survey of India
recovered some plant remains. These contained, in addition to charred cereals, a few cotton
seeds and dung of small animals. An anatomical study of the seed coat and hairs has revealed
that this ancient Nubian cotton is similar, in the characters examined, to Gossypium arboreum
var. soudunense and G. herbuceum var. ufricanum. Some hairs recovered from the dung indicate
that the animals were fed with this cotton. The significance of this find is that the Nubians
did not know at that time the potentiality of cotton hairs for making textile, but they apparently
recognized the nutritive value of cotton seeds for animal feed. The hairs show an early stage
of evolution from the wild to the cultivated cotton, having convolutions in the middle and no
convolution towards the ends.
CONTENTS
.
Introduction
Materials and methods
Results .
Nubian cotton seeds .
Seedcoat
.
Hairs .
Gossypium seeds from Africa .
Gossypium seeds from India .
Discussion
.
Archaeological significance
Botanical significance
Acknowledgements .
References
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305
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INTRODUCTION
In connection with the building of the High Aswan Dam in Egypt, funds were
collected, under the auspices of the UNESCO, to preserve ‘the heritage of thousands
of years of human suffering and triumph in the Nile Valley’ (Gerster, 1963). India was
one of the donors to this fund and was allowed to carry out excavation and study the
archaeological remains. In 1962, Mr B. B. Lal, the present Director-General of
Archaeology in India, carried out an excavation at a site near the village Afyeh, on the
west bank of the Nile, 15 km north of Anieba, the headquarters of Egyptian Nubia
21
303
K. A. CHOWDHURY AND G. M. BUTH
304
(Fig. 1). Amongst the materials collected there were some charred plant remains from
‘level ground adjacent outside a house’ (Ghosh, 1964). These were handed over to the
senior author for investigation. A preliminary report made by the archaeologist
said ‘amongst the cereals discovered at the site one may offhand identify the wheat,
barley, lentils, peas etc.’ We, however, recovered a mixture of wheat, barley, leguminous
r
Aswan
flediterronean
U A R
(Egypt)
I
/
I
Garf Husein A
0-
Statute
-
500
Miles
El Sibu
Abu Simbil
Mi,es---/
I
10
20
30
{
UNITED A R A B REPUBLIC
SUDAN
L - - _ _ _ - _ _ _ _ - _ _ _ _ _
40
f
FIGURE
1. After Gerster, (1963), showing site of excavation 0 .
seeds, rhizomes, cotton seeds and animal dung. In this paper only the last two items
will be dealt with. The cotton seeds and hairs have been matched with Gossypium
arboreum L. var. soudanense or Gossypium herbaceum L. var. africanum (Watt)
Hutchinson & Ghosh. The lint hairs recovered from the animal dung were similar to
those on the seeds mentioned above.
The site excavated, according to the Egyptian archaeologists, was inhabited by
‘A-Group’ people, who were in a stage of rural economy in which animals and plants
were being domesticated. The age of the site based on the plant materials, determined
by C14 method, is c. 2600-2400 B.C. (Ghosh, 1964).
COTTON SEEDS FROM EGYPTIAN NUBIA
305
According to Hutchinson (1959), the name cotton should preferably be applied to
only those species of Gossypium which supply seed hairs suitable for making textiles
etc. and not to the other species of Gossypium. In this paper we shall follow Hutchinson’s
definition.
MATERIALS AND METHODS
There were only four cotton seeds with hair for our investigations; one seed was
entire and the rest were partly damaged (Plate 1A). Dung pieces totalled about 200,
and all were separate.
For comparison we had fresh seeds of Gossypium from the Cotton Research Station,
Khartoum, Sudan. There were five samples of true cottons-three of Gossypium
arboreum var. soudanense and two of G. herbaceum var. africanum. There were also
single samples of three wild species, G . anomalum, G . somalense and G . stocksii. From
India we had four sets of seeds from different sources. These were G. arboreurn,
V-231R from Indian Agricultural Research Institute, Pusa, and K53-519 from
Indore, and G. herbaceurn, Digvijay from I.A.R.I., Pusa, and Jayathera from
Coimbatore.
The Nubian material had to be stored in a desiccator to save it from disintegration.
The first stage was to get rid of all dirt by treating repeatedly with 0-5-1% acetic
acid. A stage was reached when the long hairs could be removed easily; and further
treatment with acetic acid enabled us to remove the short hairs. Extreme caution was
necessary because the hairs were very brittle. Then small pieces of seed coat were taken
out and used for peels, section cutting and maceration. For peels Artschwager’s
method with some modifications was found helpful (Artschwager, 1930). The celloidinparaffin method used by Chowdhury & Ghosh (1954-55) was used for almost all the
material, and found suitable. For maceration Schultze’s method (Chamberlain, 1932)
was good for hard material, but for soft and delicate material treatment with 1-2%
acetic acid gave satisfactory results.
For animal dung a new process had to be developed. The first stage was treatment
for an hour in 1-2% acetic acid. Then hot water was poured on the mixture and it was
left for about half an hour. After centrifuging it was decanted in a flat-bottomed watchglass and the plant remains collected.
For fresh material, air was taken out by a vacuum pump till the seeds sank in water.
The same result could be obtained by keeping the seeds in vials with water and placing
the vials in an electric oven at between 50 and 60°C. Then the material was stored in
50% tap water and 50% commercial glycerine for five to six days. Small pieces of seed
coat were then removed from the centre of the seed and embedded in paraffin. Here
dehydration was done with the help of tertiary butyl alcohol. This made the material
fairly soft for cutting with a rotary microtome.
RESULTS
Nubian cotton seeds
The seeds are somewhat oval 6 x 5 mm in size. Under a binocular stereomicroscope,
two distinct types of hair are found attached to the seed coat: long white or whitish
hairs and short light to deep brown ones (Plate 1A).
K. A. CHOWDHURY AND G. M. BUTH
306
Seed coat
Under low power cross-sections of the seed coat show thick-walled epidermal
cells, which, due to their dark colour, are not individually distinct. Unevenly
distributed in an underlying pigmented layer are a few vascular bundles. Below this
is a colourless layer made up of single or double rows of cells (Plate lC,D), some of
which contain crystals of varying size and shape. Then comes the palisade tissue.
Cells are here long and arranged in a columnar fashion. When stained, the palisade
cells are individually distinct but not otherwise. Inside the palisade cells, at a distance
of about one third of their total height from the colourless layer, some remnants of
nuclear components are found. These components are laid down almost in a straight
line, somewhat resembling the light line found in some leguminous seed coats.
Hairs
The long hairs (lint) vary in length from 19-22 mm, but their width is almost
constant at 25 nm. They are thin-walled, flat and convoluted in the centre (Plate
2A) and thick-walled, round and without convolution towards the ends (Plate 2B).
Table 1. Seed coat characters of some genera of Malvaceae (from Reeves, 1936~)
~~
Name
Palisade tissue
Crystals in
colourless layer
Integumentary
bundles
Abutilon theophrasti Medic.
Modiola caroliniana L.
Callirhoe involucrata (Nun.) A. Gray
Sido rhombifolia L.
Malvwisnu drummondii Torr. & Gray
Abelmoschus esculentus L.
Paritium tiliaceum Juss.
Thespesia populnea L.
Gossypium species
Ingenhousia triloba Moc. & Sesse
present
present
present
present
absent
present
present
present
present
present
absent
absent
absent
absent
absent
absent
absent
present
present
present
absent
absent
absent
absent
absent
absent
absent
present
present
present
The short hairs (fuzz) vary in length from 1-5 mm and in width from 15-20 nm,
seldom showing any convolution (Plate 2C).
The dung pieces, when melted in liquid, yielded partly digested grass leaves and
dicotyledonous leaves and portions of seed hair with convolutions (Plate 2D).
The look of these seeds reminds one of the seeds of Malvaceae on which often seed
hairs are attached. Reeves (1936~)has made a comparative anatomical study of true
cotton seeds along with seeds of other Malvaceous plants. He has shown that, out of
ten genera, only Thespesia, Gossypium and Ingenhousiu have integumentary bundles
in the form of vascular tissue below the epidermis. Of these, Ingenhousiu has since been
placed in Gossypium (Hutchinson, Silow Sz Stephens, 1947). The remaining seven
show no vascular tissue (Table 1). The colourless layer is also absent in all except the
three genera mentioned above, and the palisade tissue is present in all except
Mulvuyiscw dmmmondii. Thus it will be seen that the Nubian material has all the
anatomical characters that these three genera possess. However, the genus Thespesiu
has much larger seeds and a very thick seed coat. The hairs are sparse and short, and
COTTON SEEDS FROM EGYPTIAN NUBIA
307
contain brown pigments. Again, the seeds of Ingenhousia are small and angular with
a thin seed coat. The hairs are scanty and inconvoluted, but have sharply angled
bends (Reeves, 1936~).These differences are so important that one can say that the
Nubian material cannot belong to these genera. We are therefore left with the genus
Gossypium, with which the Nubian material shows complete agreement in all anatomical
details. We are, therefore, justified in placing the Nubian material under Gossypium.
The genus Gossypium can be divided into two groups-the wild and the cultivated.
The wild species, which are rare and thought to be relics, are all diploid with 13 pairs of
chromosomes, except G . tomentosum from the Hawaiian Islands. The chromosomal
characteristics are so well marked in the genus that Beasley (1942) was able to place
them under four cytogenetic groups. There are four cultivated species, viz. G.
arboreum L. and G . herbaceurn L. of the Old World and G. barbadense L. and G.
hirsutum L. of the New World. The two Old World species are diploid with 13 pairs
of chromosomes and come under Beasley’s A group, whereas the two from New World
are polyhybrid species, having 26 pairs of chromosomes, and have been placed in the
A D group.
T h e wild and the cultivated species show some diagnostic characteristics in their
seed hairs. Hairs of wild species are short, unconvoluted, round in cross-section and
firmly attached to the seed coat, whereas those of the cultivated species are long as
well as short. T h e long hairs are convoluted, flat in cross-section and easily detachable.
T h e short hairs are firmly attached to the seed and are rarely convoluted (Hutchinson,
1959).
Since the ancient Nubian material comes from the Old World, it has been thought
advisable to confine the present study to only the main species now growing in the Old
World. For this purpose, fresh seeds, both cultivated and wild, were obtained from
Khartoum, Sudan. From Indian sources, four sets of cultivated cotton seeds were also
at our disposal. Our plan was first to make a comparative anatomical study of the
African Gossypium and the Nubian material, and then check up the result, in a general
way, with the data from Indian material.
Gossypium seeds from Africa
Morphologically, the seeds of five species from Africa can be placed under two
clear-cut categories. The true cotton seeds of G . arboreum and G. herbaceurn are fairly
large, ovoid and have thick seed coats. On the other hand, the wild species G. anomalum,
G. somalense and G . stocksii are comparatively small, longish and have thin seed coats.
Although the seed coat anatomy of all the species from African source shows common
features, yet there are differences in minute anatomy. For instance, there is no bulge
at the base of the epidermal cells in the wild species G. anomalum, G. somalense and G .
stocksii, but the bulge is quite frequent in the cultivated species G. arboreum and G .
herbaceum (Plate 1B). Fryxell(1964), while studying the epidermal cells of the genus
Gossypium, came across this bulge and called it a ‘foot’. For his comparative study
he made use of this structure. We have, however, thought it best not to utilize this
peculiarity of some epidermal cells for the present study, because the charred material
from Nubia did not always allow a clear view of the bulge,
K. A. CHOWDHURY AND G. M. BUTH
308
Below the epidermis, there is the outer pigmented zone (Plate 1B). Cells here are
horizontally elongated in the cross section of the seed coat and form a crushed tissue.
They are thick-walled and contain reddish to darkish brown deposits. In general,
this tissue is clearly visible in G. arboreurn and G. herbaceurn but not so in the rest of
the species. The vascular bundles are irregularly distributed in this tissue and easy
to detect.
Next to the pigmented zone is the colourless layer (Plate lB), made up of one to
two rows of cells in G. arboreurn and G. herbaceurn, but always in a single row in G.
anomalum, G. somalense and G. stocksii. Crystals present in the colourless layer also
Table 2. Anatomy of seed coat of Gossypium species
Source
North-east Africa
Fibre
Convolu- Round or
tion
flat
"Length
(mm)
*Width
(nm)
Seed coat
Layers of +Height of
colourless palisade
cells
cells (nm)
G. arboreum L. race soudanense
(Western Nubia)
present
flat
22
32
1-2
213rt8
present
flat
22
33
1-2
209*15
present
flat
19
35
1-2
214rt14
present
flat
20
30
1-2
270f22
present
absent
absent
absent
flat
round
round
round
18
8
7
10
28
13
13
18
1-2
1
1
1
270f19
158f8
125 f 14
209f 11
present
flat
21-
25
1-2
144f13
present
flat
-
33
-
-
G. arboreum L. race soudanense
(Upper Sudan)
G. arboreum L. race soudanense
(Kordofan)
G. herbaceum L. race africanum
(Mozambique, fc-8.1 9.)
G. herbaceurn L. race africanum
(Mozambique, fc-8.1)
G. anomalum Wawra & Peyr.
G. somalense Gurke
G. stockrii Masters
Ancient Nubian Cotton (from
plant remains)
Ancient Nubian Cotton (from
dung)
+
Mean of over 100 measurements of complete fibres.
Average of only two complete fibres.
++
show some variation. In G. arboreum, G. herbaceurn and G. anomalum, the percentage
of crystals in the colourless cells varies from 24 to 33, while in G. somahse and G.
stocksii it varies from 15 to 17.
Adjacent to the colourless layer, towards the inside, is the palisade tissue. Cells
here do not show any marked difference in shape but in height they exhibit some
characteristic dissimilarity. In G. arboreurn, G. herbaceurn and G. stocksii the height
varies from 209 to 270 nm, and in G. anomalum and G. somalense from 125 to 158
nm. All the cells here are extremely thick-walled and show some peculiar darkish
deposits resembling nuclear contents about one-third of the total height below the
colourless layer (Plate 1B).Below the palisade tissue occurs the inner pigmented zone,
which shows a structure similar to that described earlier for the outer pigmented zone.
COTTON SEEDS FROM EGYPTIAN NUBIA
309
T h e seed hairs of Gossypium species from the Sudan show considerable differences.
T h e true cottons, i.e. the cultivated species G . arboreum and G . herbaceum, produce
two kinds of hairs. T h e long hairs (lint) are flat in cross-section, convoluted throughout
and easily detachable. T h e short hairs (fuzz) are round, usually without any convolution
and firmly attached to the seed coat. T h e wild species, G. anomalum, G. somalense
and G . stocksii, have only one kind of hair which is round in cross-section, without
convolutions and difficult to detach. I n the cultivated species, the length of hair varies
from 18 to 28 mm and the width from 28 to 35 nm, whereas the length and width of
hair in wild species varies from 7 to 10 mm and 13 to 18 nm respectively.
T h e anatomical data given above have been summarized in Table 2. It will be seen
that the ancient cotton from Nubia is similar to the true cotton of commerce from
Africa in respect of length, width, convolution and flatness of the lint. T h e number
of cells in the colourless layer in the seed coat is also the same in both the ancient and
commercial species. Some difference in the height of palisade cells of the seed coat is
present, however. In the ancient cotton, the height is about 144 nm, whereas in the
commercial cotton it is about 212 nm. Similarities in all other anatomical details,
however, would appear to outweigh this difference. It can, therefore, be concluded
that the cotton from Nubia shows the greatest similarity to G . arboreum L. and G .
herbaceum L.-the old cultivated cottons of Africa.
Gossypium seeds from India
Study of seed coats and hairs from four sets of cultivated Gossypium, belonging to
G . arboreum and G. herbaceum, has enabled us to confirm the result obtained from the
seed from the Sudan. I n the Indian material the hairs are flat and convoluted and vary
in length from 20 to 22 mm and in width from 25 to 35 nm. T h e height of the palisade
cells is between 180-190 nm, and the colourless layer is one to two cells thick. Thus
there is a great deal of similarity in the anatomical structure of the seed coat and hair
between cottons grown in India and the Sudan, while the wild Gossypium species
from the Sudan are entirely different (Table 2).
DISCUSSION
Archaeological signijicance
Gossypium of commerce is characterized by having long, flat and convoluted seed
hairs. In fact these very characteristics have made the hairs suitable for spinning
and turning into yarn. T h e oldest written record on the use of cotton is from the
Hindu Rigveda, which dates from 1500-1000 B.C. Later, Manu also mentions cotton
thread in his religious books. But in 1928, Gulati & Turner reported on the use of
cotton for textiles from the material excavated from Mohenjo-Daro (Marshall, 193 l ) ,
which is now dated 2500-1700 B.C. (Wheeler, 1966). Gulati and Turner identified the
cotton hairs as belonging to Gossypium arboreum L.
Some recent reports from South America are interesting in this connection. From
an excavation at Huaca Prieta, north Peru, archaeologists recovered fishing net at
the level of 2400 B.c., lamp wick at 1700 B.C. and textile at 1600-1500 B.C. (Bird &
Mahler, 1951-52). All these were made of seed hairs of G . barbadense L. (Hutchinson,
310
K. A. CHOWDHURY AND G . M. BUTH
1959). These facts would lead one to think that the art of textile making was developed
separately in the Old World and the New World (Table 3).
Cotton lint with characteristics matching those of G . arboreum var. soudanense
has been reported from textiles recovered at Meroe in the Sudan from a period between
300 B.C. and A.D. 300, and at Karanog, Egyptian Nubia from the Roman Period
(Griffith & Crowfoot, 1934; Hutchinson, 1949, 1962). The latest Nubian find also
matches either G . arboreum var. soudanense or G. herbaceum var. africanum. Although
a true cotton was growing in Nubia 4500 years ago, yet the Nubians of that time do
not seem to have had any knowledge of its potentiality for making textiles. This is
so because, firstly, no evidence has so far been found from the Nubian excavation
Table 3. Showing the antiquity of the use of Gossypium of commerce
Species (putative)
~~
Site (country)
Age
Actual use
_ _ _ _ ~
Gossypium arboreum L.
Mohenjo-Daro
(Pakistan)
Gossypium barbadense L.
Huaca Prieta (North
Peru)
(a) c. 2400 B.C.
Fishing nets
(b) c. 1700 B.C.
Lamp wick
(c) c. 1600-1500 B.C. Textile
Gossypium arboreum L.
Meroe (Sudan)
300 B.C. to 300 A.D.
Textile
Gossypium arboreum L. race
soudanense
Karanog (Egyptian
Nubia)
Roman Period
Textile wrappings
2500 B.C.
Cotton seeds as animal
feed
Gossypium sp. cf. G.
herbaceum L. or G.
arboreum L.
1
Nubia (Egypt)
2500-1700
B.C.
Fragment of textile and
a string
of the use of textiles and, secondly, the earliest known use of textile by the Nubians
goes back only to the Roman Period (Griffith & Crowfoot, 1934; Nicholson, 1960).
All the same, the Nubians, from the present record, appear to be one of the earliest
people to use cotton seeds for animal feed. Although they did not have the knowledge
we have now of the chemical composition of cotton seed, yet they knew its nutritive
value as food for domestic animals.
Botanical significance
The genus Gossypium, because it contains the cottons of commerce, has received
considerable attention from botanists for more than half a century (Watt, 1907;
Beasley, 1942; Hutchinson, Silow & Stephens, 1947; Hutchinson, 1959; Saunders,
1961). These papers deal with many points which do not fall within the scope of this
contribution, where considerable attention has been paid to the anatomy of the seed
hairs and the seed coat. In 1928 Gulati & Turner reported on cotton fabrics from
Mohenjo-Daro, now dated as 2500-1700 B.C. (Wheeler, 1966), and ascribed them to
G. arboreum. They were not certain of the subspecies. They were, however, sure that
the seed hairs from the Indus Valley were not of G. herbaceum. At this stage it seems
proper to point out the limitations of anatomical studies. Although G. arboreum and
G . herbaceum are different from a taxonomic point of view, yet there is no difference
COTTON SEEDS FROM EGYPTIAN NUBIA
311
in the anatomy of their seed coats and hair. T h e organs of a plant, its tissues and cell
elements do not undergo synchronized evolution (Bailey, 1951; Chowdhury, 1954).
Thus there is nothing unusual in finding that no differentiation is possible on anatomical
characters. How Gulati and Turner came to the conclusion that they could determine
the Indus Valley cotton as G. arboreum is not clear from their paper.
Most of the publications mentioned earlier in this paper contain one important
theme, namely the origin of cultivated cottons; for instance Hutchinson (1959) says
‘By whom, and at what date, wild cottons were first brought into cultivation must
remain a matter of speculation’. In fact Watt, Hutchinson and Saunders have put
forward hypotheses indicating the likely wild Gossypium from which the cottons of
commerce might have originated in nature. Opinions on this matter have changed
with the advance of knowledge. Hutchinson et al. (1947) at first held the view that the
original domestication of Gossypium might have taken place in the Indus Valley;
but Hutchinson later accepted G. herbaceum var. africanum as truly wild in southern
Africa, and believed that from there the cultivated cottons of commerce might have
originated (Hutchinson, 1954). More recently he (Hutchinson, 1970) has acknowledged
the probability that G. arboreum is indigenous in India, and was originally wild there.
Thus the evidence now is that both Old World species, G. arboreum and G. herbaceurn
were originally wild lint-bearing species, and there is thus reason to suppose that they
were separately domesticated as a source of raw material for textiles.
The Nubian find will now be considered in the light of these hypotheses. Hutchinson
(1949), discussing the cotton fabrics from Meroe in the northern Sudan, suggested
that they were made from lint of G. arboreum var. soudanense. This cotton is now
found as an associate of man, in gardens and in waste places near habitations. He
therefore proposed that both the plant and the craft of spinning and weaving were
introduced, probably from India. The Nubian find shows conclusively that a cotton
anatomically indistinguishable from the Old World species was growing in Nubia
before the Nubians had acquired the textile craft. Moreover, the Nubian cotton bore
the most primitive lint hairs that have so far been observed. Thus, though it is not
possible to judge whether the plant was G. arboreum or G. herbaceum, it is possible
to assert that a cotton anatomically indistinguishable from the Old World species of
Gossypium was growing in Nubia before considerations of the needs of a textile craft
could have induced man to introduce it from elsewhere.
In summing up it may be pointed out that the 4500-year-old Nubian cotton supplies
actual evidence of the early stage of evolution from wild to lint-bearing Gossypium.
There is no doubt about the importance of this find, but at the same time it has raised
more controversial points than it has solved. This is probably due to the fact that the
archaeological Gossypium remains are too few at the present time to enable us to
reconstruct step by step the origin of cultivated cotton.
ACKNOWLEDGEMENTS
Dr H. K. Jain, Indian Agricultural Research Institute, Pusa, D r K. M. Simlote,
Cotton Specialist, Indore and Dr T. K. K. Kunniyan, Agricultural Institute, Coimbatore supplied us with authentically identified seeds. So did Dr J. H. Saunders of the
Cotton Research Station, Khartoum, Sudan. We are grateful to all of them, and
312
K. A. CHOWDHURY AND G. M. BUTH
especially to Dr Saunders, without whose help it would not have been possible for us
to carry out the investigation in the way that we have done here. Sir Joseph Hutchinson,
who happened to be in India during early 1970, was kind enough to go through some
of our microscopic preparations and the draft of this paper. We are very grateful to
him for this.
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EXPLANATION O F PLATES
PLATE1
A. Nubian cotton seeds, x6.
B. Cross-section of seed coat of G. herbaceum var. africanum from the Sudan, x 120.
C. Cross-section Nubian seed coat; note, crushed epidermis, vascular tissue and single row colourless
layer, ~ 6 0 0 .
D. Shows one to two rows of cells in colourless layer, x600.
Co.L., Colourless layer; Ep., epidermis; Int., inner integument; Pal., palisade cells; Pi.L., pigmented
layer; V.T., vascular tissue.
PLATE
2
A. Middle portion of a lint from Nubian seeds showing convolutions, ~ 4 5 0 .
B.End portion of the same fibre without convolution, x450.
C. Fuzz from Nubian seeds, x 300.
D. Portion of a lint from animal dung, x300.
C., Convolution; Fz., fuzz.