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Journal of Archaeological Science 34 (2007) 335e343

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

The impact of manuring on nitrogen isotope ratios in


cereals: archaeological implications for reconstruction
of diet and crop management practices
A. Bogaard a,*, T.H.E. Heaton b, P. Poulton c, I. Merbach d
a

Department of Archaeology, University of Nottingham, University Park, Nottingham NG7 2RD, UK


b
NERC Isotope Geosciences Laboratory, British Geological Survey, UK
c
Rothamsted Research, UK
d
Bad Lauschstadt Experimental Station, Umweltforschungszentrum Leipzig-Halle, Germany
Received 22 February 2006; accepted 21 April 2006

Abstract
Recent archaeological studies of human diet have used stable nitrogen isotope ratios (d15N) from human bone collagen to infer the relative
importance of terrestrial plant and animal foods. This approach is based on widely observed enrichment of d15N up the food chain, plants having
distinctly lower values than the herbivores that consume them. Studies of early farming diets in Britain, Denmark and Germany have tended to
detect relatively high d15N values (e.g. c. 9&), interpreted as evidence of a diet largely based on animal products, though archaeobotanical
evidence for crop cultivation (e.g. carbonised cereal grain and chaff) is widespread. This paper investigates the impact of manuring on d15N
values in modern cereals, and of charring on these cereal values. The results from two long-term experiments demonstrate that manuring significantly raises d15N in cereal grain and chaff. Depending on manuring levels and frequency, it appears that human diets with a major component of such grain would conventionally be interpreted as indicating a largely animal-based diet or a mixed plant/animal diet. Moreover,
preliminary analyses of experimentally charred grain and chaff from manured and unmanured conditions are promising for the extraction of
reliable ancient d15N values from archaeobotanical cereal remains. The wider implications of these results, and the need for further work,
are discussed.
2006 Elsevier Ltd. All rights reserved.
Keywords: Nitrogen; Stable isotopes; Manuring; Cereals; Neolithic; Crop husbandry

1. Introduction
Stable carbon and nitrogen isotope ratios are now routinely
used to infer aspects of past human diets (e.g. [40,42,54,
55,62]). Carbon isotope ratios (d13C values) can potentially
distinguish between terrestrial and marine foods and/or C3
and C4 pathway plants [15,53,61]. Nitrogen isotope ratios
(d15N values) are used to infer the trophic level of food based

* Corresponding author. Tel.: 44 115 951 4815; fax: 44 115 951 4812.
E-mail address: amy.bogaard@nottingham.ac.uk (A. Bogaard).
0305-4403/$ - see front matter 2006 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jas.2006.04.009

on increases in d15N values up the food chain, herbivores having distinctly higher values than the plants that they eat (e.g.
[17,43,60,66]). In combination, carbon and nitrogen isotope
ratios have been used to reconstruct ancient human diets and
have, for example, played a key role in debate over the nature
of the MesolithiceNeolithic transition in north-west Europe
(e.g. [53,54,62,64]). The isotopic technique of reconstructing
ancient human diet is particularly attractive given the difficulty
of inferring the relative dietary contribution of animals and
plants from their remains on archaeological sites (e.g. [14]).
Recent applications of the isotopic method of dietary reconstruction in Britain [52,54,62], Denmark [55] and southern

336

A. Bogaard et al. / Journal of Archaeological Science 34 (2007) 335e343

Germany [20,21] have been interpreted to suggest the


following:
 Neolithic communities made little use of marine foods,
obtaining most of their protein from a combination of terrestrial animals and C3 plants
 d15N values are generally consistent with a largely animalbased diet, suggesting that C3 plants played a limited dietary role (despite widespread archaeobotanical evidence
for cereal use)
While there has been lively debate over the interpretation of
these results as they relate to marine versus terrestrial resources (e.g. [29,39,44]), the issue of terrestrial animal- versus
plant-based foods has received less attention. Durrwachter
et al. [21] have recently summarised a number of issues surrounding the interpretation of d15N as a reflection of the relative importance of plant- and animal-based foods. First,
isotope analysis of archaeological remains of mammals is
commonly undertaken on collagen separated from bone, and
the 15N-enrichment in bone collagen relative to dietary protein
values may be higher (e.g. up to 5&) than the commonly assumed c. 3& [5,19]. Second, there is a lack of information
available on d15N in ancient plant remains, largely, according
to Durrwachter et al. [21], because they are generally not adequately preserved to permit analysis. As a result, assumptions have to be made using generalised plant values and
archaeological herbivore bone collagen. Third, d15N may
tend to reflect the meat portion of the diet since animal foods
are generally richer in protein than plant foods, with the result
that d15N values are particularly insensitive to low or moderate consumption of plant foods [21]. Durrwachter et al. [21]
conclude, however, that d15N values can be used to distinguish heavy plant consumers from heavy animal consumers.
The aim of this paper is to begin exploring the potential
range of variability in plant d15N values, specifically those
for cultivated cereals, in order to set archaeological reconstructions of human diet and crop husbandry practices on
a firmer foundation. Cereal stable isotope values may be affected by alterations to the growing environment introduced
by ancient farmers; variation in d13C in cereals, for example,
has been related to water economy and irrigation in arid environments (e.g. [3,4,47]). A factor that could directly affect cereal d15N values is manuring e the application of animal dung
to cultivation plots in order to restore nutrients and enhance
crop yields. High d15N values in animal manure largely result
from the preferential loss of 14N in volatile gaseous ammonia,
leaving residual ammonium relatively enriched in 15N. This
ammonium is subsequently converted to nitrate with high
d15N values, which is taken up by plants [28,35,37]. Nitrates
are the major source of nitrogen used for the biosynthesis of
plant amino acids, which eventually end up in the bone collagen of consumers [58]. Previous studies suggest that the application of animal manure raises d15N values in soil and plants
(e.g. [10,16,63,68,70,71]). Van Klinken et al. [67] note that
a manuring effect on plant values would have a significant
impact on d15N in human consumers. These authors conclude,

Thus, there is a need to check for anthropogenic effects in the


archaeological food chain, which can be done by measuring
associated plant and animal remains [67, original italics].
In their study of Neolithic diet in central Europe, Durrwachter
et al. [21] have also stressed that isotopic information from ancient plant remains, especially crops, would serve to greatly
enhance the accuracy of human dietary reconstructions.
Chemical and soil micromorphological studies of ancient
soils (palaeosols) can provide direct evidence for manuring
(e.g. [12,13,22,24,63]). In agricultural landscapes cultivated
over many centuries, however, such evidence is rarely preserved and the practice can only be inferred indirectly, from
spreads of sherds and other inorganic inclusions across the
landscape (e.g. [1,68]), or from ecological characteristics of
arable weeds associated with crop remains in archaeological
deposits (e.g. [33,34]). An archaeobotanical study of weed assemblages and crop husbandry practices in Neolithic central
Europe by Bogaard [6] concluded that manuring was a likely
cause of frequent high fertility, and similar inferences have
been made for south-east Europe [26]. Indeed, one reading
of the Neolithic package of plant and animal domesticates
is that it was precisely such integration of plant and animal
(by-)products that enabled farming to successfully spread
across a range of environments [6,7,8]. From this perspective,
it is plausible that relatively high d15N values in Neolithic human remains from central Europe and elsewhere are due, at
least in part, to a manuring effect.
In order to assess the potential impact of manuring on the
reconstruction of human diet, this paper considers new data
on d15N in cereals grown under manured and unmanured conditions at two long-term experimental stations: Rothamsted,
Hertfordshire, England [23,57] and Bad Lauchstadt, LeipzigHalle, Germany [36]. These data on cereals grown under
known conditions can be used to assess how far the d15N
values in cereal grain and chaff are affected by manuring,
and hence the impact that manuring in the past would theoretically have on values in human bone collagen. This study is the
first of its kind to look at the effects of manuring on cereal
d15N values through time using long-term experimental data.
A second aim of this paper is to consider the impact of charring on d15N cereal values, since charred grains and chaff represent the most widespread form of archaeobotanical evidence
for cultivation.

2. Materials and methods


Two long-term experiments of over one hundred years duration, including one with archive cereal samples going back
to the first decade of the experiment, were selected in order
to assess the long-term effects of farmyard manure application
on d15N in cereals. Details of the two experiments are given in
Table 1.
The Broadbalk Wheat Experiment at Rothamsted, Hertfordshire studies winter wheat (Triticum aestivum L.) cultivation under various treatments. One plot (plot 22; previously
plot 2 or 2B) is treated only with farmyard manure. This

A. Bogaard et al. / Journal of Archaeological Science 34 (2007) 335e343

337

Table 1
Details of the experimental plots samples for isotopic analysis
Experiment

Location

Period

Cropping

Manuring treatment(s)

Soil type

References

Broadbalk,
Rothamsted

Hertfordshire,
England

1844present

Winter wheat

Cattle manure, 35
t/ha every year

[23,57]

Static fertilization
experiment,
Bad Lauchstadt

Leipzig-Halle,
Germany

1902present

Sugar beet-spring
barley-potatoeswinter wheat

Cattle manure, 20
and 30 t/ha every
second year

Chromic luvisol;
flinty-silty clay loam
over clay-with-flints
Haplic chernozem;
loess

plot receives annual dressings of cattle manure at the rate of


35 t (fresh weight) per hectare each year. Such a level of application is equivalent to the annual manure production of about
three cattle over one hectare [56, cf. 41]. A control plot (plot 3)
has received no farmyard manure or mineral fertilizers since
1844.
The Bad Lauchstadt experiment consists of a four-course
crop rotation (including winter wheat, T. aestivum L.) and
two different manuring levels: 20 or 30 t of farmyard manure
(Stalldung, fresh weight) per hectare every second year in
plots 12 and 6, respectively. A control plot (plot 18) receives
no inputs.
Samples of at least 20 whole wheat plants were collected
just before harvest time in 2004 from manured and control
plots in both experiments. Grain and rachis (the stalk segments to which spikelets are attached in the cereal ear) harvested from 20 plants were randomly subsampled (using
a riffle box) to provide bulk samples for analysis. All bulk samples analysed consisted of c. 20e30 grains or rachis segments.
The selection of rachis for measurement, in addition to grain,
is due to the fact that it is identifiable to species and is widely
preserved archaeologically (e.g. in charred form).
Archive samples of grain and rachis from Rothamsted for
the years 1852, 1895, 1935 and 1965 were also subsampled
to provide bulk samples for analysis. These years were chosen
to provide spread across the period covered by the archive
(1844e2005). Additionally, unpublished measurements of cereal grain from the manured and control plots in 1991 were included along with the new data (Smolinska, unpublished data
1991). The archives for Bad Lauchstadt consisted of grain
from recent years only; archive samples of wheat grain from
2001 to 2002 were subsampled to provide bulk samples for
analysis.
While the aim of bulk sampling was to explore variation
between treatments and variation through time, detailed sampling of individual plants was conducted to enable investigation of variation in d15N within a single cereal ear and
between plants. Samples of grain and rachis from individual
spikelets of four cereal plants e two ears from two different
plants harvested in the 2004 control plot, and two ears from
two different plants harvested in the 2004 manured plot at
Rothamsted e were taken for analysis. Table 2 summarises
the individual grain and rachis samples analysed.
Random subsamples of material harvested from Rothamsted
and Bad Lauchstadt in 2004 were also subjected to charring in
order to ascertain the impact of this process on d15N values.
Subsamples of wheat grain and rachis (c. 20e30 grains or

[2,36]

rachis segments) were charred in a low-oxygen atmosphere


(wrapped in aluminium foil) at 230  C for 2e24 h e conditions
that have proved suitable to reproduce undistorted archaeological charred grain and rachis [M. Charles and G. Jones pers
comm.; cf. 65]. Charring was carried out at the Department
of Archaeology, University of Nottingham, in a digitally controlled chamber furnace.
At the NERC Isotope Geosciences Laboratory, Keyworth,
all samples were homogenised using a freezer mill, weighed
into tin capsules, and combusted in an elemental analyser
(Thermo Finnigan Flash EA) on-line to an isotope ratio
mass spectrometer (Thermo Finnigan Delta XL). Sample
15
N/14N ratios were calculated as d15N values versus atmospheric N2, on the basis of comparison with samples of a laboratory plant standard whose d15N value was determined by
comparison with IAEA-N-1 (assuming d15N of IAEA-N1 0.4& [31]).
3. Results
3.1. Within- and between-plant variations in grain and
rachis d15N
The results for grain and rachis internodes from the individual spikelets of four wheat ears harvested in 2004 at Rothamsteds Broadbalk experiment (two ears from two plants
harvested in the control plot, and two ears from two plants harvested in the manured plot) are shown in Fig. 1. Values for
grain were reasonably constant. The rachis displayed greater
variation, typically showing a decline in d15N from the basal
spikelet to the terminal spikelet.
Variations between individual plants must also be expected.
Thus, from the data in Fig. 1, average grain d15N values differed by only 0.1& between the two ears from the plants harvested in the control plot (Plants 1 and 2), but by 1.3&
between the two plants harvested in the manured plot (Plants
Table 2
Summary of samples from individual plants harvested in 2004 at Rothamsted;
NIL control plot; FYM farmyard manure

Plant
Plant
Plant
Plant

1,
2,
3,
4,

ear
ear
ear
ear

1
2
3
4

(NIL)
(NIL)
(FYM)
(FYM)

Grain

Rachis

17
19
23
21

Not analysed
19 spikelets
23 spikelets
21 spikelets

spikelets
spikelets
spikelets
spikelets

Note: 1 grain and 1 rachis internode per spikelet were analyzed.

A. Bogaard et al. / Journal of Archaeological Science 34 (2007) 335e343

338

ROTHAMSTED
within-ear variation. NIL, plant 1

ROTHAMSTED
within-ear variation. NIL, plant 2

+1.0

+1.0

0.0

0.0

-1.0

-1.0

d15N vs AIR

d15N vs AIR

-2.0
-3.0

-2.0
-3.0
-4.0

-4.0
-5.0

-5.0
0

10

12

14

16

18

Spikelet position
Grain

ROTHAMSTED
within-ear variation. FYM, plant 3

+8.0

+7.0

+7.0

+6.0

+6.0

+5.0

+5.0

+4.0
+3.0
+2.0

+4.0
+3.0
+2.0

+1.0

+1.0

0.0

0.0
0

10

15

20

25

-1.0

Spikelet position
Grain

Rachis

ROTHAMSTED
within-ear variation. FYM, plant 4

+8.0

-1.0

10 12 14 16 18 20

Grain

d15N vs AIR

d15N vs AIR

Spikelet position

10

15

20

25

Spikelet position

Rachis

Grain

Rachis

Fig. 1. d15N values for grain and rachis internodes from the individual spikelets of four wheat ears: (a) plant 1, control plot at Rothamsted; (b) plant 2, control plot
at Rothamsted; (c) plant 3, manured plot at Rothamsted; and (d) plant 4, manured plot at Rothamsted. Rachis from plant 1 was not measured.

3 and 4). In another study, d15N values for whole wheat plants
(excluding roots) from plots treated with farmyard manure and
inorganically-fertilized plots ranged over 2& (1 SD 0.8 and
1.0& for two groups of n 5; Poulton unpublished data
1991).
3.2. Differences in grain and rachis d15N between
manured and control plots
Tables 3a and b and Fig. 2 summarise the results for bulk
samples per treatment and year. Elevated d15N values are associated with manured versus control treatments at Rothamsted
and Bad Lauchstadt, and these differences are visible in both
grain and rachis.
Several points are worth noting from these results. First,
a manuring effect is evident from the earliest archive samples available at Rothamsted (1852), less than 10 years after
the experiment began (in 1844). Second, manured and control
values in both experiments remain fairly constant through

time. Third, rachis d15N values are consistently lower than


those in grain. It can also be noted here that d15N values in cereal straw and leaves are also lower than those in grain (Smolinska, unpublished data 1991). Fourth, grain and rachis from
the lower level of manure application at Bad Lauchstadt
Table 3a
Results for the analysis of bulk samples from Rothamsted (na material not
available for analysis; nd not determined; NIL control plot; Fym farmyard manure)
Year

d15N

%N

Grain

1852
1895
1935
1965
1991
2004

Rachis

Grain

Rachis

Nil

FYM

Nil

FYM

Nil

FYM

Nil

FYM

2.7
2.9
4.0
0.7
0.6
0.8

5.8
7.8
8.3
7.4
8.6
6.6

na
0.6
0.4
2.5
na
2.6

na
5.0
7.1
5.3
na
3.4

1.8
1.9
1.8
2.0
1.5
1.5

2.0
1.8
nd
2.3
2.0
1.9

na
nd
0.6
0.3
na
nd

na
nd
0.6
0.8
na
nd

A. Bogaard et al. / Journal of Archaeological Science 34 (2007) 335e343

339

Table 3b
Results for the analysis of bulk samples from Bad Lauchstadt (na material not available for analysis; NIL control plot; FYM1 biennial 20 t/ha,
FYM2 biennial 30 t/ha)
Year

d15N

%N

Grain

2001
2002
2004

Rachis

Grain

Rachis

Nil

FYM1

FYM2

Nil

FYM1

FYM2

Nil

FYM1

FYM2

Nil

FYM1

FYM2

1.0
0.1
0.8

4.1
2.7
3.2

5.7
4.1
3.4

na
na
-1.3

na
na
0.9

na
na
2.2

1.2
1.4
1.1

1.1
1.4
1.3

1.3
1.5
1.3

na
na
0.2

na
na
0.3

na
na
0.3

(FYM1) is associated with lower d15N values than the higher


level (FYM2). The even higher rate of yearly manure application at Rothamsted is associated with the highest d15N values,
though it should be noted that the control plot values at Rothamsted also tend to be higher than those at Bad Lauchstadt.
Overall, it is clear that manuring has a distinct impact on
d15N values in both cereal grain and rachis. Moreover, the
low levels of natural variation within and between plants reviewed above would not obscure the manuring effect on
d15N values.
3.3. The impact of charring on d15N values in wheat
grain and rachis
Work by DeNiro and Hastorf [18] on charred plant remains
(seeds and tubers) from Peruvian archaeological sites showed
that d15N in charred plant material was similar to that in modern counterparts, suggesting that charring did not bias the signature. Fig. 3 shows the results of preliminary analyses on the
impact of charring, with d15N values in wheat grain (Fig. 3a)
and rachis (Fig. 3b) harvested in 2004 from the manured and
control plots at Rothamsted and Bad Lauchstadt, and charred
at 230  C under low-oxygen conditions for up to 24 h.
Fig. 3a indicates that charring causes only minor distortion
of d15N values in grain and that the manuring effect is not

obscured. The distortion of d15N values in rachis is more


marked (Fig. 3b), and the relative position of the control plots
at the two experiments is reversed after 2 h charring, but the
contrast between manured and control plots remains intact.
4. Discussion
With enrichment of c. 3& from one trophic level to the
next, the conventional wisdom is that bone collagen from humans having a largely plant-based diet would have d15N values
of c. 6& (assuming plant values of c. 3&), while a diet
based on herbivores should result in values of c. 9& (assuming herbivore values of c. 6&). A mixed diet in which both
plants and animals played a major role would lie between
6 and 9& (e.g. [52]). Neolithic values for human bone collagen recently reported from southern Germany, Denmark, the
west coast of Scotland and southern Britain (Table 4) have
been interpreted as evidence that diets were largely animalbased (southern German sites, Danish sites, western Scottish
sites, some Hambledon Hill samples, Parc le Breos) or, in
some cases in southern Britain (Hazleton, West Kennet,
some Hambledon Hill samples), a mixed diet of plant- and
animal-based foods.
The d15N values of Neolithic crops, however, could have
a major impact on the interpretation of such results. The

A
+10.0

+8.0

+8.0

+6.0

d15N vs AIR

+6.0

d15N vs AIR

+10.0

+4.0
+2.0

+4.0
+2.0

0.0

0.0

-2.0

-2.0

-4.0
1852

1895

1935

1965 1991 2004

years sampled
FYM gr
NIL gr

FYM ra
NIL ra

-4.0
2001

2002

2003

2004

years sampled
FYM2 gr
FYM1 gr
NIL gr

FYM2 ra
FYM1 ra
NIL ra

Fig. 2. d15N values for bulk samples of grain and rachis from (a) Rothamsted and (b) Bad Lauchstadt. NIL control plot; FYM farmyard manure (annual 35t/ha);
FYM1 biennial 20t/ha, FYM2 biennial 30t/ha.

A. Bogaard et al. / Journal of Archaeological Science 34 (2007) 335e343

340

B
+8.0

+6.0

+7.0

+5.0
+4.0

d15N vs AIR

d15N vs AIR

+6.0
+5.0
+4.0
+3.0

+3.0
+2.0
+1.0
0.0

+2.0

-1.0

+1.0

-2.0

0.0

-3.0
0

10

15

20

25

30

10

Time (hours)
Roth Nil
Roth FYM

15

20

25

30

Time (hours)
Roth Nil
Roth FYM

BadL FYM1
BadL FYM2

BadL NIL

BadL FYM1
BadL FYM2

BadL NIL

Fig. 3. The effect of charring at 230  C and its duration on d15N in grain and rachis (0 h uncharred): (a) grain; (b) rachis. Roth Rothamsted; BadL Bad
Lauchstadt.

data from modern experiments reported here suggest that


a diet largely based on manured cereals could result in Neolithic humans having fairly high d15N values e i.e. resembling
those resulting from a largely animal-based diet, or a mixed
plant- and animal-based diet. Thus, with trophic enrichment
of c. 3&, the bulk samples of manured cereal grain from Rothamsted (c. 6 to 8&) would be expected to yield values of
c. 9 to 11& in consumers (resembling a largely animalbased diet), with the lower levels of biennial manuring at
Bad Lauchstadt (grain values of c. 3 to 6&) yielding values
of c. 6 to 9& in consumers (resembling a mixed plant- and
animal-based diet).
If Neolithic farming in southern Germany, Denmark and
Britain was of the permanent, small-scale and intensive type
resembling gardening [6,7,8,26; cf. 9], long-term manuring

comparable to the applications at Rothamsted (equivalent to


the manure of c. 3 cattle per hectare each year) is not implausible. It is likely that manure inputs, along with other labourintensive measures, varied somewhat from year to year
depending on a range of factors (labour, livestock, weather,
etc.) [26,27; cf. 59]. Thus, variation among human d15N values
at well-sampled sites such as Hambledon Hill could reflect
variability in crop growing conditions through time or among
households, rather than diets ranging from mixed plant/animal
to largely animal-based [52].
The measurement of d15N in associated faunal remains is
widely accepted as a critical factor in the interpretation of
human d15N values. Values for domestic cattle and sheep from
the Neolithic sites in Germany and Britain are shown in Table 4.
Interpretation of some human d15N values as indicative of

Table 4
Summary of Neolithic d15N values from bone collagen of humans (adults), domestic cattle and sheep (excluding juvenile animals) in southern Germany, Denmark
and Britain; n number of samples
n

Cattle
d15N

9.9
9.7

21
40

7.0
5.7 (ave)

1
5

10.0

TRB
TRB

Site

Period

Human
d15N min

Human
d15N max

Herxheim, Germany
Trebur, Germany

LBK
HinkelsteinGrossgartach
TRB

7.8
8.5

12.1
10.5

9.9

10.0

Sheep
d15N

Interpretation

Reference

6.2 (ave)

Animal-based diet
Animal-based diet

[20,21]
[20,21]

Animal-based diet

[55]

8.2

Animal-based diet

[55]

8.0

Animal-based diet

[55]

Animal-based diet
Animal-based diet
Animal-based diet
Animal-based (10)
or mixed (7 to 9)
Animal-based diet
Mixed diet
Mixed diet

[55]
[62]
[62]
[52]

Human
d15N average

Ostrup, Store
mose, Denmark
A
Undlose, Store
mose, Denmark
A
Bodal, Store
mose, Denmark
A
Aldersro, Denmark
Carding Mill Bay
Crarae
Hambledon Hilla

TRB
Earlier Neolithic
Earlier Neolithic
Earlier Neolithic

7.5
8.8
9.1
7.0

9.3
10.0
9.5
10.5

8.4
9.3
9.2
9.5

6
9
3
56

Parc le Breos
Hazletonb
West Kennetb

Earlier Neolithic
Earlier Neolithic
Earlier Neolithic

8.9
7.3
8.1

10.4
8.4
8.5

9.7
7.9
8.3

8
5
3

a
b

5.5 (ave)

4.6 (ave)

Mostly adults according to Richards [52]; approximate values derived from scatter plot [52, Fig. 12.2].
Adult status of human remains uncertain [52].

5.0 (ave)

[52]
[52]
[52]

A. Bogaard et al. / Journal of Archaeological Science 34 (2007) 335e343

a largely animal-based diet rests on the fact the human samples


tend to appear one trophic level higher (c. 3& higher) than
associated herbivores (Table 4). The manuring of cereals in
an intensive cultivation regime, however, provides an alternative explanation for this discrepancy, assuming that humans
were the primary consumers of grain. Moreover, if arboreal
vegetation (leafy or branch hay) provided an important source
of fodder as has often been suggested for Neolithic central and
western Europe (e.g. [25,38,49,50,51]), the low d15N associated with forest ecosystems (e.g. [30,46,69]) would tend to distinguish plant consumption between livestock and humans in
terms of their d15N values. Furthermore, the results for rachis
values from Rothamsted and Bad Lauchstadt suggest that
d15N values in chaff tend to be lower and more variable than
in grain, as is the %N content of chaff. Measurements of
d15N in cereal straw and leaves likewise suggest that these
are low relative to grain (Smolinska, unpublished data 1991).
Even if livestock diets were supplemented with fodder consisting of the chaff by-products of manured cereals, therefore, the
impact on their d15N would be reduced in comparison with
human consumers of manured grain.
One methodological issue to be addressed concerns the
measurement of whole grain versus protein d15N values in manured and unmanured cereals. It has been assumed thus far
that changes in d15N values caused by manuring largely reflect
changes in protein N isotope ratios. While this assumption appears justified from previous work on d15N values in proteins
[11], further work is needed to confirm that whole grain and protein compound-specific d15N values show a similar manuring
effect. A related methodological point, raised by Durrwachter
et al. [21], is the need to characterise nitrogen stable isotope
values for individual amino acids from bone collagen, in order to narrow down their potential dietary sources. Finally,
the preliminary charring results are promising for the reliable
measurement of d15N values in charred archaeobotanical
material, but further work is needed to explore the full range
of relevant crops and charring conditions, as well as to
address issues of diagenesis and contamination in archaeological burial environments.

5. Conclusions
The results of the analyses presented here support previous
suggestions (e.g. [21,67]) that information on plant d15N
values e and in particular those of potential staples e is critical for accurate assessment of animal- and plant-based foods
in the human diet. The suggestions made here regarding alternative interpretations of Neolithic d15N values, however, must
remain speculative until reliable measurements of archaeological plant d15N values are available. Archaeological plant
values from Neolithic sites in north-west Europe would help
to resolve the problem of equifinality between a diet based
largely on animal products and one based on manured cereals.
Plant values may also be useful for the interpretation of similar
d15N values from human samples in later periods (e.g.
[32,45,48]).

341

Acknowledgements
This research was made possible by a grant-in-kind from
the NERC Isotope Geosciences Facilities Steering Committee.
We thank the Lawes Trust for access to the archived Rothamsted samples and Ursula Smolinska for the 1991 Broadbalk
data. Rothamsted Research receives grant-aided support from
the Biotechnology and Biological Sciences Research Council
of the UK. We thank the UFZ Centre for Environmental Research Leipzig-Halle for providing the plant samples of the
Static Fertilization Experiment Bad Lauchstadt and for allocation of archived grain samples. Finally, the authors are grateful
to Oliver Craig, Glynis Jones, Rick Schulting and two anonymous reviewers for insightful comments on the paper.

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