Towards the use of radiocarbon as a dietary

proxy: Establishing a first wide-ranging

radiocarbon reservoir effects baseline for
Germany
Ricardo Fernandes1,2,3 , Christoph Rinne 4, Marie-Jose Nadeau5, Pieter Grootes 1
1

Institute for Ecosystem Research, Christian-Albrechts-Universitt zu Kiel, Kiel, Germany, 2Leibniz Laboratory
for Radiometric Dating and Isotope Research, Christian-Albrechts-Universitt zu Kiel, Kiel, Germany,
3
McDonald Institute for Archaeological Research, University of Cambridge, Cambridge, UK, 4Institute of
Prehistoric and Protohistoric Archaeology, Christian-Albrechts-Universitt zu Kiel, Kiel, Germany, 5Graduate
School Human Development in Landscapes, Christian-Albrechts-Universitt zu Kiel, Kiel, Germany
Radiocarbon reservoir effects (RREs) are observed when the 14C concentration of aquatic reservoirs is lower
than the contemporary atmosphere. Within these reservoirs, aquatic species will also have a 14C depleted
signal, and humans feeding on these species will show a dietary RRE. Human dietary RREs are often
viewed as a problem for the establishment of reliable chronologies. However, they also represent an
opportunity to introduce radiocarbon as a dietary proxy when investigating possible past human
consumption of aquatic food groups. Here, a synthesis of previously published and new radiocarbon
dates on edible aquatic species from central and northern Germany is presented. The samples were
collected from modern and archaeological contexts. The goal was to provide an approximate RRE
baseline within Germany. The results show that within the German context, local RREs in edible aquatic
species are usually large and variable. The variability in local RREs implies that precise quantitative
human dietary estimates will most likely not be possible. However, the large values of local RREs allow the
use of 14C measured in human bone collagen as an extra dietary proxy that can aid in detecting the
consumption of aquatic food groups when traditional isotopic proxies (13C and 15N) do not provide
unambiguous estimates.
Keywords: Radiocarbon, Radiocarbon reservoir effects, Dietary proxies, Dietary tracers, Stable isotopes, Ancient diets

Introduction

14

An aquatic reservoir may show a C concentration

lower than the contemporary atmospheric value.
This is expressed as a radiocarbon reservoir effect
(RRE). In aquatic reservoirs, atmospheric CO2 is dissolved and in equilibrium with carbonic acid (H2CO3),
2
bicarbonate (HCO
3 ) and carbonate (CO3 ). Leached
carbonates from the geological background effectively
have a zero 14C concentration, and their contribution
to the carbonic acidcarbonate equilibrium lowers
the 14C concentration of the aquatic reservoir, an
effect also known as hard water effect (Geyh 2000).
Through underwater photosynthesis, plants and
algae acquire the 14C depleted signal which is then
transferred within aquatic food webs. A relationship
between local water alkalinity and the magnitude of

Correspondence to: Ricardo Fernandes, Institute for Ecosystem Research,

Christian-Albrechts-Universitt zu Kiel, Olshausenstrasse 75, D-24118
Kiel, Germany. Email: rfernandes@gshdl.uni-kiel.de; rf385@cam.ac.uk

Association for Environmental Archaeology 2014

DOI 10.1179/1749631414Y.0000000034

RREs in aquatic species has previously been reported

(Keaveney and Reimer 2012). However, significant
RREs were also observed in low alkalinity areas and
attributed to organic carbon inputs depleted in 14C
from the terrestrial catchment. Spatial and temporal
variations in hydrological patterns can also contribute
to variations in local RREs (Ascough et al. 2007),
including carbon contributions from 14C-depleted
groundwater in inland contexts (Landmeyer and
Stone 1995). Finally, within aquatic food webs, the
magnitude of local RREs will also depend on the
aquatic species themselves and their growth stage
(Fernandes et al. 2013).
Humans that have a significant intake of aquatic
food groups may also exhibit detectable dietary
RREs. Given the widespread use of radiocarbon
dating of human remains for the establishment of
chronologies in archaeological studies, the possible
presence of a dietary RRE is often viewed as a
major challenge. Several archaeological case studies

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Towards the use of radiocarbon as a dietary proxy

have shown that human dietary RREs are prevalent in

different historical periods, geographical areas and
human cultures (Lanting and van der Plicht 1998;
Cook et al. 2001; Shishlina et al. 2007, 2009; Lillie
et al. 2009). These include also recent examples
within Germany (Olsen et al. 2010; Fernandes et al.
2012a; Nadeau et al. 2012).
In secure and well-established archaeological contexts containing human remains, it may be possible
to establish an independent reference chronology
that does not rely on the radiocarbon dating of
human material. Different approaches can be taken
to obtain independent reference burial dates. These
include, the radiocarbon dating of contextually associated samples of terrestrial origin (e.g. animal or plant
remains), dendro-dated wood, precise typological
dating (e.g. coins or pottery), identifying the individual and its year of death from historical documents
or from collective graves associated with a well-dated
event (e.g. historical battle site, recorded natural catastrophe). In these cases, 14C measurements of human
remains can effectively be used as a dietary tracer or
proxy, provided that consumed aquatic food groups
had a significant RRE.
The main goal of this study is to provide a first, tentative 14C baseline of edible aquatic species for central
and northern Germany, relying on new and published
radiocarbon dates obtained from modern and archaeological contexts. The gathering of this type of data is
the first step necessary for the use of 14C as a dietary
proxy in the study of past human consumption of
aquatic food groups.

Material and Methods

Material
The data presented here were collected from published
radiocarbon data (Fernandes et al. 2012a, 2013;
Philippsen and Heinemeier 2013) and new radiocarbon and isotopic measurements. All sites are situated
within central and northern Germany and their
location is represented in Fig. 1.
New radiocarbon dates were obtained from modern
fish samples collected in the vicinity of the Neolithic
collective grave sites of Calden and Odagsen
(Raetzel-Fabian 2000; Rinne 2003). Three fish
samples were collected from the river Rebbe
(51792577N 9874202E), a tributary of the river
Leine, located in the vicinity of Einbeck. Another
three fish samples were collected from an artificial
lake, fed by a natural inflow, within the precinct of
Schloss Wilhelmsthal (51391095N 9421455E)
located 12 km northwest of Kassel. Samples from the
river Rebbe were collected in April 2013, whereas
samples from Schloss Wilhelmsthal were collected in
August 2013.

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All published and new samples obtained for this

study are listed in Table 1 and include only edible
aquatic species (water, plant and bird data of previous
studies were not included). The data represent a combination of modern and archaeological samples.
Archaeological data were only included when a welldefined and secure association between aquatic and
terrestrial sample could be established.

Methods
Fish flesh lipid removal
Lipids were removed from fish flesh to obtain a pure
protein isotopic signal. The extraction followed a
well-established method (Howland et al. 2003). A
sample of fish flesh was placed in a volume (20 ml/g)
of ultrapure solvent dichloromethane/methanol
(2:1). The samplesolvent mix was ultrasonicated for
10 minutes and then left to rest for 30 minutes. The
solvent was discarded and the sequence was repeated
twice. Finally, the flesh sample was dried in air at
room temperature.
Isotope Ratio Mass Spectrometry
Isotopic measurements were done at the Division of
Archaeological, Geographical and Environmental
Sciences, University of Bradford. Approximately
05 mg of fish flesh was weighed into tin capsules for
analysis on a Flash EA 1112 coupled to a Thermo
Delta Plus Mass Spectrometer. Stable isotope ratios
are expressed in the conventional delta notation
(13C and 15N) relative to VPDB (Vienna PeeDee
belemnite standard) and AIR (atmospheric nitrogen).
Precision was better than 02.
Radiocarbon dating
Fish flesh samples were combusted to CO2 in a sealed
quartz tube containing copper oxide and silver.
Reduction to graphite was done in a hydrogen atmosphere at 600C over an iron catalyst. The irongraphite
mix was pressed into a pellet and the 14C concentration measured in a 3 MV Tandetron from High
Voltage Engineering Europa (Nadeau et al. 1997,
1998).
The 14C concentration of a sample is expressed in
per cent of modern carbon ( pMC). From the 14C
sample concentration, a radiocarbon age (Age)
expressed in years (yr) can be calculated using
equation (1) and the conventional Libby 14C halflife value (5568 yr):
Age (yr) = 8033 ln( pMC/100)

(1)

By convention, radiocarbon ages are presented in

years before present (BP) where present (100 pMC)
is defined for the year 1950 AD (Stuiver and Polach
1977).

Fernandes et al.

Towards the use of radiocarbon as a dietary proxy

Figure 1 Distribution of sites within Germany for which local (archaeological and modern) RREs have been determined. Ranges
in RRE values refer only to measurements on fish or bivalve flesh/collagen, with the exception of Quern-Neukirchen for which
only shell values are available. The values from Schloss Wilhelmsthal are given, although the evidence points to a significant
terrestrial contribution to fish diet.

A RRE is defined as the difference in 14C concentration between the contemporary atmosphere and
the aquatic reservoir. This can be expressed in years
using equation (1) to calculate the difference between
the age of the atmosphere and that of the aquatic
sample. The atmospheric 14C concentration and, by
implication, its radiocarbon age can be determined
directly from collected atmospheric CO2 or indirectly
from terrestrial samples. For the archaeological
material listed in Table 1, terrestrial and aquatic
samples were analysed and the RRE is the radiocarbon age difference between the two. For modern
samples, atmospheric 14C concentrations at time of
collection are approximately known and listed in
Table 1 (Levin et al. 2013; I Levin personal communication). Atmospheric nuclear weapons tests, carried
out mostly between 1954 and 1963, have produced significant amounts of 14C. Thus, post 1954 AD atmospheric 14C concentrations are higher than 100 pMC
and radiocarbon ages have negative values.

Results
RREs for aquatic species collected within Germany
both from modern and archaeological contexts are
listed in Table 1. The data include both published
results (Fernandes et al. 2012a, 2013; Philippsen and
Heinemeier 2013) and new data obtained for this
study. Fig. 1 shows the geographical distribution and
the range in RRE values for sample flesh, except for
Quern-Neukirchen where only bivalve shell values
are available.
Quern-Neukirchen: The site is located approximately
200 m from the present day Baltic coast and is an
example of a shell midden site in use during the Late
Neolithic period in northern Germany, known under

the term Dolchzeit (cal 22001700 BC)

(Fernandes et al. 2012a). The RREs for QuernNeukirchen were defined by comparison of radiocarbon dates from mollusc shells of an unidentified
species (KIA-43951 and KIA-43952) with those from
collagen extracted from a bone of an unidentified herbivore (KIA-43919) and red deer antlers (KIA-43920).
Associated pairs mollusc-animal were collected in
close proximity to two well-defined contexts located
at different depths. The RRE values for both associated pairs are of similar magnitude (550 40 and
570 40 yr).
Kiel Fjord: Four modern samples of blue mussels
(Mytilus edulis) were collected from a beach location
in the Kiel Fjord. The following paired sample codes
specify flesh and shell of the same individual specimen: KIA-44379 and KIA-44383; KIA-44380 and
KIA-44384; KIA-44381 and KIA-44385; KIA-44382
and KIA-44386. Mean RRE values for blue mussel
flesh (605 yr) and shell (600 yr) are similar and also
statistically similar to the mean RRE value observed
at Quern-Neukirchen (560 yr).
Zauschwitz: The site of Zauschwitz is located in the
immediate vicinity of the Weie Elster River. The site
contains graves of the Salzmnde type (a variant of the
Neolithic Funnel Beaker culture). Large contemporaneous deposits of freshwater mussels were found,
suggesting a significant human consumption of
aquatic food groups in the past. Two individuals
buried with Salzmnde-type pottery were radiocarbon
dated significantly older than the established pottery
chronology and constitute examples of human
dietary RREs (Table 2). Two modern duck mussels
(Anodonta anatina) (KIA-40107-1 and KIA-40107-2)
were collected from the Weie Elster River and their

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Table 1 List of RREs for aquatic species collected from modern and archaeological contexts within Germany. References for
modern samples are the contemporary atmospheric values (data from Levin et al. 2013; I Levin personal communication).Three
archaeological aquatic samples are compared with terrestrial radiocarbon values from associated animal remains. The isotopic
values (13C and 15N) for aquatic samples are also listed, when available
Reference age (yr BP)

Aquatic sample

Site: Quern-Neukirchen (a)

3660 30*1
Mollusc (shell)
Mollusc (shell)
3650 25*2
Site: Kiel Fjord (a)
ca. 320
Blue mussel (flesh)
ca. 320
Blue mussel (shell)
ca. 320
Blue mussel (flesh)
ca. 320
Blue mussel (shell)
ca. 320
Blue mussel (flesh)
ca. 320
Blue mussel (shell)
ca. 320
Blue mussel (flesh)
ca. 320
Blue mussel (shell)
Site: Zauschwitz (a)
ca. 320
Duck mussel (flesh)
ca. 320
Duck mussel (shell)
ca. 320
Duck mussel (flesh)
ca. 320
Duck mussel (shell)
Bivalve shell
4460 40*3
Site: Lake Rosenfeld (a)
ca. 320
Swollen river mussel (flesh)
ca. 320
Swollen river mussel (shell)
Site: Lake Schwerin (b)
ca. 320
Pike (flesh)
ca. 320
Bream (flesh)
ca. 320
Eel (flesh)
ca. 320
Eel (flesh)
ca. 320
Zebra mussel flesh
ca. 320
Zebra mussel shell
Site: Lake Ostorf (b)
ca. 320
Zebra mussel flesh
ca. 320
Zebra mussel shell
Site: Alster (c)
ca. 440
Bivalve shell
ca. 440
Snail shell
ca. 440
Roach (bone collagen)
Site: Trave (c)
ca. 440
Roach (bone collagen)
ca. 440
Roach (bone collagen)
ca. 420
Spined loach (flesh)
ca. 420
Crayfish (flesh)
ca. 420
Roach (flesh)
Site: Rebbe (d)
ca. 240
Trout (flesh)
ca. 240
Roach (flesh)
ca. 240
Bream (flesh)
Site: SW (d)
ca. 240
Common rud (flesh)
ca. 240
Common rud (flesh)
ca. 240
Bream (flesh)

Lab code

Age (yr BP)

13C ()

KIA-43951
KIA-43952

4210 30
4220 30

KIA-44379
KIA-44383
KIA-44380
KIA-44384
KIA-44381
KIA-44385
KIA-44382
KIA-44386

290 30
350 40
260 30
200 20
280 30
275 20
315 25
300 30

221 02
14 02
226 02
11 02
219 02
15 02
223 02
06 02

68 02

74 02

75 02

73 02

610 35
670 45
580 35
520 30
600 35
595 30
635 30
620 35

1140 20
1495 20
1010 20
1195 20
6225 45

306 02
152 02
303 02
147 02

126 02

127 02

1460 30
1815 30
1330 30
1515 30
1765 60

KIA-44405
KIA-44404

575 20
410 30

285 02
79 02

94 02

895 30
730 35

KIA-46304
KIA-46305
KIA-46306
KIA-46307
KIA-46310
KIA-46310

270 20
190 30
260 25
625 25
230 20
210 30

197 02
194 02
147 02
206 02
271 02
36 02

132 02
115 02
128 02
170 02
81 02

590 30
510 35
580 30
945 30
550 30
530 35

KIA-46311
KIA-46311

1120 20
1060 30

278 02
75 02

146 02

1440 30
1380 35

AAR-11460
AAR-11461
AAR-11462

1214 34
433 32
223 29

132 01
154 01
255 01

122 03

1654 40
873 35
663 35

AAR-11394
AAR-11396
AAR-12875
AAR-12876
AAR-12878

285 32
244 28
1664 39
1365 40
67 32

259 01
242 01
272 01
279 05
223 01

149 04
153 10
156 03
119 20
149 02

725 35
684 35
2084 45
1785 45
487 35

KIA-49800
KIA-49801
KIA-49802

160 40
305 40
345 40

272 02
277 02
289 02

130 03
134 03
131 03

400 45
545 45
585 45

KIA-49820
KIA-49821
KIA-49822

260 40
120 40
135 40

277 02
275 02
306 02

79 03
79 03
55 03

20 45
120 45
105 45

KIA-40107-1
KIA-40107-1
KIA-40107-2
KIA-40107-2
KIA-41497

15N ()

RRE (yr)

550 40
570 40

Data were compiled from the following studies: (a) Fernandes et al. (2012a), (b) Fernandes et al. (2013), (c) Philippsen and
Heinemeier (2013) and (d) this study. Site SW stands for Schloss Wilhelmsthal. *13 archaeological collagen as reference sample
from: 1. KIA-43919, unidentified herbivore; 2. KIA-43920, red deer antler; and 3. KIA-41496 cattle bone.

shells showed large RRE values (1815 30 and

1515 30 yr). However, RREs of flesh for the same
specimens were significantly lower (1460 30 and
1330 30 yr, respectively). An archaeological cattle
bone (KIA-41496) was recovered from a shell
deposit at the same depth as a bivalve shell of an unidentified species (KIA-41497). Comparison of the
radiocarbon dates obtained from this pair gives a
RRE of 1765 30 yr, a value similar to those
observed for modern shells.

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Lake Rosenfeld: Lake Rosenfeld is a dammed lake

located ca. 5 km southeast of the Kiel Fjord. A
single specimen of swollen river mussel (Unio
tumidus) had a RRE value of 895 30 yr for the
flesh (KIA-44405) and 730 35 yr (KIA-44404) for
the shell.
Lake Ostorf: Within Lake Ostorf the small island of
Tannenwerder contains a Neolithic cemetery (so
defined by the presence of Funnel Beaker pottery)
where several radiocarbon dated individuals showed

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Towards the use of radiocarbon as a dietary proxy

Table 2 Examples of past humans showing significant dietary RREs. There are no 15N values available for the individuals listed
from the study by Lillie et al. (2009). The value between brackets represents an average of 15N values reported in the same study
for Neolithic individuals within the same region
Sample code/ID

Location

Study: Lanting and van der Plicht (1998)

Dirk VI
Low countries
Hadewig
Low countries
Gerlach
Low countries
Study: Lillie et al. (2009)
OxA-17495
Ukraine
OxA-17499
Ukraine
OxA-17502
Ukraine
Study: Higham et al. (2010)
OxA-9386
Russian North Caucasus
OxA-9387
Russian North Caucasus
OxA-14774
Russian North Caucasus
OxA-9390
Russian North Caucasus
OxA-9391
Russian North Caucasus
OxA-9388
Russian North Caucasus
OxA-9389
Russian North Caucasus
OxA-14777
Russian North Caucasus
Study: Ascough et al. (2012)
SUERC-2018
Iceland
SUERC-2016/SUERC 2660
Iceland
Study: Olsen et al. (2010)
UtC-8173
Ostorf
UtC-7447
Ostorf
UtC-7440
Ostorf
UtC-7445
Ostorf
UtC-7446
Ostorf
UtC-7448
Ostorf
UtC-8180
Ostorf
UtC-8181
Ostorf
AAR-9752
Ostorf
UtC-7449
Ostorf
UtC-8179
Ostorf
Study: Cook et al. (2001)
OxA-8502
Romania
OxA-8547
Romania
OxA-8581
Romania
OxA-8583
Romania
OxA-8548
Romania
Study: Fernandes et al. (2012a)
KIA-37858
Zauschwitz (Germany)
KIA-37859
Zauschwitz (Germany)

13C ()

15N ()

RRE (yr)

Medieval
Medieval
Medieval

189
198
196

106
109
110

340 32*
227 46*
282 32*

Neolithic
Neolithic
Eneolithic

225
188
208

(113)
(113)
(113)

251 49
472 49
111 60

Medieval
Medieval
Medieval
Medieval
Medieval
Medieval
Medieval
Medieval

168
176
173
167
165
173
131
152

99
106
117
113
103
103
93
114

313 48
140 46
85 40
334 54
205 56
96 47
108 47
52 48

Medieval
Medieval

193
191

100
90

120 49
212 35

Neolithic
Neolithic
Neolithic
Neolithic
Neolithic
Neolithic
Neolithic
Neolithic
Neolithic
Neolithic
Neolithic

206
22.0
208
208
209
20.0
199
206
198
194

128
139
134
137
142
13.0
133
132
152
153

168 52
264 61
288 54
31 56
103 59
327 80
435 51
397 51
401 57
835 65
552 60

Mesolithic
Mesolithic
Mesolithic
Mesolithic
Mesolithic

196
193
195
185
182

132
139
151
150
153

Neolithic
Neolithic

200
203

105
92

Time period

510 117
470 247
450 300
532 127
345 116
>300
>300

*Individual RRE values for the Lanting and van der Plicht (1998) study were established using the calibration curve Intcal 13 (Reimer
et al. 2013) and not the values reported in the cited study.

large dietary RREs (Olsen et al. 2010). A single specimen (KIA-46311) of zebra mussel (Dreissena polymorpha) collected from Lake Ostorf had a RRE of 1440
30 yr for the flesh and a similar value (1380 35 yr)
for the shell.
Lake Schwerin: Lake Schwerin is a large lake (area
ca. 62 km2) located approximately 15 km to the
northeast of Lake Ostorf. Four fish specimens (KIA46304, KIA-46305, KIA-46306 and KIA-46307) of
pike (Esox lucius), bream (Abramis brama) and eel
(Anguilla anguilla) collected from Lake Schwerin had
REE values between 510 30 and 945 35 yr. A
single specimen (KIA-46310) of zebra mussel (D. polymorpha) had RRE values of 550 30 yr for the flesh
and 530 35 yr for the shell.
River Alster: The Mesolithic Erteblle site of
Kayhude is located by the river Alster. For this site,

radiocarbon dating of food crusts, recovered from

ceramic shards, showed, in some instances, significant
RREs, indicating that these contained freshwater
ingredients (Philippsen and Heinemeier 2013). A
single fish specimen (AAR-11462), collected from
the river Alster, showed a RRE of 663 35 yr. An
unidentified freshwater snail shell (AAR-11461) had
a RRE of 873 35 yr, whereas an unidentified bivalve
shell (AAR-11460) had a RRE of 1654 40 yr. A
specimen of roach (Rutilus rutilus) had a RRE,
measured in bone collagen, of 663 35 yr.
River Trave: The Erteblle site of Schlamersdorf is
located by the Trave. From this site, radiocarbon
dates on pottery food crusts suggested the presence
of significant RREs (Philippsen and Heinemeier
2013). Four fish specimens (AAR-11394, AAR11396, AAR-12875 and AAR-12878) of roach (R.

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rutilus) and spined loach (Cobitis taenia) together with

a crayfish (AAR-12876), collected from the Trave, had
RREs between 487 35 and 2084 45 yr.
River Rebbe: The sample collection site is located at
12 km from the Funnel Beaker collective grave site of
Odagsen. Radiocarbon ages of some human remains
recovered from Odagsen may suggest possible dietary
RREs (Fernandes et al. 2012b). The three fish specimens (KIA-49800, KIA-49801 and KIA-49802) of
trout (Salmo trutta), roach (R. rutilus) and bream (A.
brama) collected from the river Rebbe had radiocarbon ages between 400 45 and 585 45 yr.
Schloss Wilhelmsthal: The site is located 21 km east
of the Neolithic collective grave site of Calden, where
fish remains were identified in the interior of the
monument (Raetzel-Fabian 2000). Three fish specimens (KIA-49820, KIA-49821 and KIA-49822) of
common rud (Scardinius erythropthalmus) and bream
(A. brama) were collected from an artificial lake
within Schloss Wilhelmsthal grounds. This lake is fed
by a natural stream. However, contrary to initial
reports, it was later established that the local fish were
being fed by humans and thus had a significant intake
of terrestrial food groups. RRE values for the three
specimens varied between 20 45 and 120 45 yr.

Discussion
RREs within Germany
The global average marine surface reservoir effect is
approximately 400 yr, while the radiocarbon results
listed in Table 1, for both coastal and inland locations
within Germany, show significantly larger RRE
values. Excluding the results for Schloss
Wilhelmsthal, that probably include a terrestrial diet
provided by humans, the lowest RRE was recorded
in the Rebbe (400 45 yr) whereas the largest REE
was recorded for the Trave (2084 45 yr). These
results confirm previous observations that showed
that for inland locations, local RREs can be both
very large and variable (Lanting and van der Plicht
1998; Keaveney and Reimer 2012). It is possible to
directly compare past and modern RREs for the
sites of Zauschwitz and Quern-Neukirchen. The
mean RRE value for Quern-Neukirchen, obtained
from archaeological samples, was 560 yr, a value
similar to the mean value observed for the modern
shell samples collected from the Baltic coastline at
the Kiel Fjord (606 yr). In the case of Zauschwitz, a
single bivalve archaeological shell had a RRE of
1765 35 yr, a value similar to those observed for
modern samples (1515 30 and 1815 30 yr). In
spite of this, some caution is necessary when inferring
past RREs from modern values. Temporal changes in
climatic or hydrological patterns and in catchment
areas will influence local RREs (Ingram and
Southon 1996; Stein et al. 2004; Ascough et al. 2007,

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2010). In addition to natural variability, human intervention in shaping the landscape and in modifying the
course of streams and rivers might also have caused
significant changes to the local RREs. Another issue
to consider in modern RREs is the possible contribution of groundwater having a 14C enriched signal
due to the nuclear tests of previous decades.
Archaeological samples might also present some difficulties in determining the relevant local RREs for
human dietary studies. The Zauschwitz and
Rosenfeld radiocarbon measurements on modern
bivalve samples show significant differences between
bivalve meat and shell. Bivalve shell is mainly precipitated from dissolved inorganic carbon (DIC) while
certain bivalve species metabolise dissolved organic
carbon (DOC) or particulate organic carbon (POC)
to build their flesh. These fractions (POC, DOC and
DIC) may show significantly different 14C concentrations and this justifies the observed radiocarbon
age differences between meat and shell for bivalves
from Lake Rosenfeld and Zauschwitz (Fernandes
et al. 2012a). No such differences were observed for
the zebra mussels collected from Lakes Ostorf and
Schwerin given that zebra mussels have a diet specialised in phytoplankton, which in turn obtains its
carbon in underwater photosynthesis from DIC
(Baker et al. 1998). Research is underway to verify
whether radiocarbon dates obtained from the protein
fraction in bivalve shells (conchiolin), measured as
an alternative to bivalve shell carbonate, provide a
radiocarbon age similar to bivalve flesh.

Radiocarbon as a Dietary Proxy

Aquatic species-dependent variability in dietary habits
and the temporal and spatial variability in local RREs
represent an important challenge for the use of 14C
measured in humans to obtain precise quantitative
estimates on the intake of aquatic food groups.
However, given the large RRE values observed,
especially for inland locations, 14C measured in
humans can be used to detect the consumption of
aquatic food groups when traditional isotopic analysis
cannot provide an unambiguous result.
Table 2 provides a list of examples of human dietary
RREs from different locations and time periods. Large
local RREs and a similar isotopic baseline of fish and
animal isotopic values (13C and 15N) are common to
all listed sites, with the exception of the Russian North
Caucasus due to the presence of C4-plants in the local
environment (Higham et al. 2010). Human dietary
RREs were established by comparison of radiocarbon
dates from human bone collagen with associated independent reference dates. In the case of the study by
Lanting and van der Plicht (1998) the time of death
was known from historical records. For the studies
by Cook et al. (2001), Lillie et al. (2009), Higham

Fernandes et al.

et al. (2010), Olsen et al. (2010) and Ascough et al.

(2012) individual human dietary RREs were determined by radiocarbon dating of associated animal
remains. The study by Higham et al. (2010) also
reported the chronology of buried artefacts (ceramics
and coins) that provided ages similar to animal radiocarbon dates. For the study by Fernandes et al. (2012a)
the reference date is based on ceramic typology and a
precise estimate cannot be provided.
Measurements of isotopic ratios, 13C and 15N, in
human bone collagen are often used as proxies to
provide dietary estimates. In predominantly C3-plant
environments, humans having a fully terrestrial diet
typically show 13Ccollagen values of ca. 20.
Human intake of marine aquatic foods can be detected
when 13Ccollagen values significantly higher than ca.
20 are observed (Schoeninger et al. 1983). For
instance, the comparison of 13C bone collagen
values in Neolithic Europeans with the preceding
Mesolithic populations shows clear differences
(Tauber 1981; Richards et al. 2003a), and these
results have been interpreted to signify a complete
shift towards a terrestrial-based diet. However, C4plants (e.g. millet and maize) have 13C signals
similar to marine foods and as such high 13Ccollagen
values may also be observed in some inland locations.
This is illustrated in Table 2 by the individuals from the
Russian North Caucasus (Higham et al. 2010) that all
show 13Ccollagen values above 18. Furthermore,
even when restricted to C3-type food groups, the isotopic difference between terrestrial and freshwater 13C
signals may be too small to offer the possibility of distinguishing a terrestrial from a freshwater contribution.
In some instances, the consumption of freshwater food
groups may be detected by 13Ccollagen values lower
than ca. 20 (Lanting and van der Plicht 1998).
Table 2 shows that the individual with code OxA17495 from a Neolithic site in the Dnieper basin
(Ukraine) has a 13Ccollagen value of 225. In spite
of this example, the majority of individuals listed in
Table 2 originating from predominantly C3 environments show 13Ccollagen values of ca. 20.
Consumption of freshwater food groups can, in
some instances, be established through the measurement of 15N in human bone collagen. A compilation
of published 15N data, with major outliers removed,
from Late Mesolithic and Neolithic central and northern European sites provides average values for animal
bone collagen of 15N = 59 14 (n = 268) and
15N = 88 15 for fishbone collagen (n = 27)
(Boric et al. 2004; Ogrinc and Budja 2005; Bsl
et al. 2006; Drrwchter et al. 2006; Bocherens et al.
2007; Hedges et al. 2008; Oelze et al. 2011; Stevens
et al. 2012). The higher 15N values observed for fish
compared with land animals constitute the basis for
the use of the 15Ncollagen proxy to detect the

Towards the use of radiocarbon as a dietary proxy

consumption of freshwater aquatic food groups.

Typical 15Ncollagen values for European Neolithic
populations are within the range 811
(Schoeninger et al. 1983; Richards et al. 2003b;
Fischer et al. 2007) with the lower limit interpreted
as dietary protein predominantly from plant foods
and the upper limit interpreted as dietary protein predominantly from animal foods (Richards and
Trinkaus 2009). Thus, human 15Ncollagen values
above 12 are often interpreted as representing a significant intake of aquatic foods. Pre-historic individuals listed in Table 2 from the studies by Olsen et al.
(2010) in Ostorf (Germany) and by Cook et al.
(2001) at the Iron Gates (Romania) all show
15Ncollagen values above 12. However, the remainder of the case studies listed in Table 2 all show individuals with 15Ncollagen values between 90 and
117 (Lanting and van der Plicht 1998; Higham
et al. 2010; Ascough et al. 2012; Fernandes et al.
2012a).
The interpretation of 15Ncollagen values needs to
consider that there are in fact multiple food groups
and that the isotopic signals of the different groups
may differ significantly. In addition, the real uncertainty that should be considered in the interpretation
of human 15Ncollagen values should not be the
typical experimental uncertainty (ca. 02) but the
natural inter- and inner-bone variability (ca. 1)
(DeNiro and Schoeninger 1983; Balasse et al. 1999;
Waters-Rist et al. 2011). Thus, there is not a simple
univocal correspondence between a certain isotopic
value and the intake of a particular food group.
Instead a single 15Ncollagen value should be viewed
as resulting from multiple possible mixture proportions each with an assigned probability. To illustrate that different dietary mixing proportions are
consistent with the same human 15Ncollagen signal, a
simple linear model is presented. This model takes as
isotopic baseline for protein (the source of dietary
nitrogen) 15N values from food groups reported
above that were available to Late Mesolithic and
Neolithic populations from central and northern
European sites (15Nanimal = 59; 15Nfish = 88).
As a reference for plant 15N, two possible values
were considered. A value of 2 represents a typical
15N value for edible European plants during the
Holocene (Richards and Trinkaus 2009). However,
recent research has shown that the practice of manuring in cereal crops was prevalent during the Neolithic
and a value of 45 should be used as reference for
manured plants (Bogaard et al. 2013). The linear
model considers the contribution of paired protein
sources weighed by percentage of intake (unmanured
plant protein vs. fish protein; manured plant protein
vs. fish protein; fish protein vs. animal protein). A
human diet-to-collagen offset of 55 was added to

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Fernandes et al.

Towards the use of radiocarbon as a dietary proxy

the calculated isotopic mix values. This value represents a consensus value derived from published
data on human diet-to-hair 15N enrichment values
(Minagawa et al. 1986, 1992; Schoeller et al. 1986;
Yoshinaga et al. 1996; Hedges et al. 2009;
Huelsemann et al. 2009) plus the isotopic offset (ca.
1) between human hair and bone collagen
(OConnell and Hedges 1999; OConnell et al. 2001;
Richards 2001). The chosen reference value is similar
to the reference value (ca. 6) derived from isotopic

values measured in human red blood cells during a

controlled feeding experiment (OConnell et al.
2012). A value of 55 for the diet-to-collagen offset
is higher than the traditionally used 34 range
(Hedges and Reynard 2007); however, it should be
noted that the higher value will, if incorrect, underestimate the contribution of fish to human diet.
Model outputs for human 15Ncollagen, shown in
Fig. 2, also include estimated human dietary RREs
for different radiocarbon ages of aquatic food groups
(500, 1000 and 1500 yr). To estimate the actual contribution of carbon from aquatic food sources towards
human bone collagen it is also necessary to consider
that part of the collagen carbon (ca. 25%) is routed
from dietary lipids and carbohydrates (Fernandes
et al. 2012c). Thus in Fig. 2 values given for human
dietary RREs, corresponding to 15Ncollagen = 12,
represent both a non-routed model and routed model
(between brackets). For the routed model it is
assumed that the only source of energy macronutrients
(carbohydrates and lipids) is terrestrial. The comparison of modelled results for the differently paired
food groups clearly shows that the same range of
15Ncollagen values, between 9 and 12 (grey area), is
compatible with significantly different dietary contributions of selected food groups. Furthermore, the
results show that significant human dietary RREs
may be observed with low protein contributions from
aquatic food groups. These results are in agreement
with the human dietary RREs listed in Table 2.
The German RRE baseline shows the widespread
presence of large local RREs. Although the variability
in local RREs is too large to allow for precise dietary
estimates, the use of 14C as an extra dietary proxy can
provide a valuable contribution in ancient human diet
reconstruction studies. The question of completeness
in the adoption of an agricultural subsistence strategy
and the total, or not, abandonment of fish consumption across the MesolithicNeolithic transition is a
prime example of such potential use.

Conclusion
15

Figure 2 Linear model results to estimate human Ncollagen

values (continuous green line) from the contributions of
paired food groups. Reference isotopic values were:
unmanured plant protein (15N = 20), manured plant
protein (15N = 45), animal protein (15N = 59) and fish
protein (15N = 88). A 55 diet-to-collagen offset was
added to calculate mix food isotopic values. The different
dashed lines represent modelled human dietary RREs for
different radiocarbon ages of aquatic food groups (500, 1000
and 1500 yr). The grey area defines the isotopic 15Ncollagen
range between 9 and 12. The vertical line defines the
intersect for 15Ncollagen = 12 and the adjacent values the
corresponding human dietary RREs for each value of aquatic
food RRE in non-routed and routed (between brackets)
dietary models.

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A first wide-ranging baseline of RREs has been established for Germany. Radiocarbon measurements on
aquatic food species edible for humans obtained
from modern and archaeological contexts show the
presence of large (>400 yr) and variable local RREs.
Observed variability does not allow the use of 14C
measured in human collagen to obtain precise quantitative estimates of dietary intake of aquatic food
groups. However, observed large local RREs,
especially in inland contexts, permit the use of 14C as
an extra dietary proxy that can detect the intake of
aquatic food groups when other standard isotopic
proxies (13C and 15N) fail to provide unambiguous
estimates.

Fernandes et al.

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