Towards the use of radiocarbon as a dietary proxy: Establishing a first wide-ranging radiocarbon...

8/21/2019 Towards the use of radiocarbon as a dietary proxy: Establishing a first wide-ranging radiocarbon reservoir effects

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Towards the use of radiocarbon as a dietaryproxy: Establishing a first wide-ranging

radiocarbon reservoir effects baseline forGermany

Ricardo Fernandes1,2,3, Christoph Rinne4, Marie-Jose Nadeau5, Pieter Grootes1

1Institute 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,3McDonald 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 lowerthan the contemporary atmosphere. Within these reservoirs, aquatic species will also have a 14C depletedsignal, and humans feeding on these species will show a dietary RRE. Human dietary RREs are oftenviewed as a problem for the establishment of reliable chronologies. However, they also represent anopportunity to introduce radiocarbon as a dietary proxy when investigating possible past humanconsumption 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 werecollected from modern and archaeological contexts. The goal was to provide an approximate RREbaseline within Germany. The results show that within the German context, local RREs in edible aquaticspecies 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 14

C measured in human bone collagen as an extra dietary proxy that can aid in detecting theconsumption of aquatic food groups when traditional isotopic proxies (13C and 15N) do not provideunambiguous estimates.

Keywords: Radiocarbon, Radiocarbon reservoir effects, Dietary proxies, Dietary tracers, Stable isotopes, Ancient diets

IntroductionAn aquatic reservoir may show a 14C concentrationlower than the contemporary atmospheric value.This is expressed as a radiocarbon reservoir effect(RRE). In aquatic reservoirs, atmospheric CO2is dis-solved and in equilibrium with carbonic acid (H2CO3),

bicarbonate (HCO3

) and carbonate (CO32

). Leachedcarbonates from the geological background effectivelyhave a zero 14C concentration, and their contributionto the carbonic acidcarbonate equilibrium lowersthe 14C concentration of the aquatic reservoir, aneffect also known as hard water effect(Geyh 2000).Through underwater photosynthesis, plants andalgae acquire the 14C depleted signal which is thentransferred within aquatic food webs. A relationshipbetween local water alkalinity and the magnitude of

RREs in aquatic species has previously been reported(Keaveney and Reimer 2012). However, significantRREs were also observed in low alkalinity areas andattributed to organic carbon inputs depleted in 14Cfrom the terrestrial catchment. Spatial and temporalvariations in hydrological patterns can also contribute

to variations in local RREs (Ascough et al. 2007),including carbon contributions from 14C-depletedgroundwater in inland contexts (Landmeyer andStone 1995). Finally, within aquatic food webs, themagnitude of local RREs will also depend on theaquatic species themselves and their growth stage(Fernandeset al. 2013).

Humans that have a significant intake of aquaticfood groups may also exhibit detectable dietaryRREs. Given the widespread use of radiocarbondating of human remains for the establishment ofchronologies in archaeological studies, the possiblepresence of a dietary RRE is often viewed as amajor challenge. Several archaeological case studies

Correspondence to: Ricardo Fernandes, Institute for Ecosystem Research,Christian-Albrechts-Universitt zu Kiel, Olshausenstrasse 75, D-24118Kiel, Germany. Email: [email protected]; [email protected]

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have shown that human dietary RREs are prevalent indifferent historical periods, geographical areas andhuman cultures (Lanting and van der Plicht 1998;Cook et al. 2001; Shishlina et al. 2007, 2009; Lillieet al . 2009). These include also recent exampleswithin Germany (Olsen et al. 2010;Fernandes et al.2012a;Nadeau et al. 2012).

In secure and well-established archaeological con-texts containing human remains, it may be possibleto establish an independent reference chronologythat does not rely on the radiocarbon dating ofhuman material. Different approaches can be takento obtain independent reference burial dates. Theseinclude, the radiocarbon dating of contextually associ-ated samples of terrestrial origin (e.g. animal or plantremains), dendro-dated wood, precise typologicaldating (e.g. coins or pottery), identifying the individ-ual and its year of death from historical documents

or from collective graves associated with a well-datedevent (e.g. historical battle site, recorded natural cata-strophe). In these cases, 14C measurements of humanremains can effectively be used as a dietary tracer orproxy, provided that consumed aquatic food groupshad a significant RRE.

The main goal of this study is to provide a first, ten-tative 14C baseline of edible aquatic species for centraland northern Germany, relying on new and publishedradiocarbon dates obtained from modern and archae-ological contexts. The gathering of this type of data is

the first step necessary for the use of

14

C as a dietaryproxy in the study of past human consumption ofaquatic food groups.

Material and MethodsMaterialThe data presented here were collected from publishedradiocarbon data (Fernandes et al . 2012a, 2013;Philippsen and Heinemeier 2013) and new radiocar-bon and isotopic measurements. All sites are situatedwithin 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 Neolithiccollective grave sites of Calden and Odagsen(Raetzel-Fabian 2000; Rinne 2003). Three fishsamples were collected from the river Rebbe(51792577N 9874202E), a tributary of the riverLeine, located in the vicinity of Einbeck. Anotherthree fish samples were collected from an artificiallake, fed by a natural inflow, within the precinct ofSchloss Wilhelmsthal (51391095N 9421455E)

located 12 km northwest of Kassel. Samples from theriver Rebbe were collected in April 2013, whereassamples from Schloss Wilhelmsthal were collected inAugust 2013.

All published and new samples obtained for thisstudy are listed in Table 1 and include only edibleaquatic species (water, plant and bird data of previousstudies were not included). The data represent a com-bination of modern and archaeological samples.Archaeological data were only included when a well-defined and secure association between aquatic andterrestrial sample could be established.

MethodsFish flesh lipid removal

Lipids were removed from fish flesh to obtain a pureprotein isotopic signal. The extraction followed awell-established method (Howland et al. 2003). Asample of fish flesh was placed in a volume (20 ml/g)of ultrapure solvent dichloromethane/methanol(2:1). The samplesolvent mix was ultrasonicated for10 minutes and then left to rest for 30 minutes. The

solvent was discarded and the sequence was repeatedtwice. Finally, the flesh sample was dried in air atroom temperature.

Isotope Ratio Mass Spectrometry

Isotopic measurements were done at the Division ofArchaeological, Geographical and EnvironmentalSciences, University of Bradford. Approximately05 mg of fish flesh was weighed into tin capsules foranalysis on a Flash EA 1112 coupled to a ThermoDelta Plus Mass Spectrometer. Stable isotope ratios

are expressed in the conventional delta notation(13C and 15N) relative to VPDB (Vienna PeeDeebelemnite standard) and AIR (atmospheric nitrogen).Precision was better than 02.

Radiocarbon dating

Fish flesh samples were combusted to CO2in a sealedquartz tube containing copper oxide and silver.Reduction to graphite was done in a hydrogen atmos-phere at 600C over an iron catalyst. The irongraphitemix was pressed into a pellet and the 14C concen-

tration measured in a 3 MV Tandetron from HighVoltage Engineering Europa (Nadeau et al. 1997,1998).

The 14C concentration of a sample is expressed inper cent of modern carbon (pMC). From the 14Csample concentration, a radiocarbon age (Age)expressed in years (yr) can be calculated usingequation (1) and the conventional Libby 14C half-life value (5568 yr):

Age (yr) = 8033ln(pMC/100) (1)

By convention, radiocarbon ages are presented inyears before present (BP) where present (100 pMC)is defined for the year 1950 AD (Stuiver and Polach1977).

Fernandes et al. Towards the use of radiocarbon as a dietary proxy

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A RRE is defined as the difference in 14C concen-tration between the contemporary atmosphere andthe aquatic reservoir. This can be expressed in yearsusing equation (1) to calculate the difference betweenthe age of the atmosphere and that of the aquaticsample. The atmospheric 14C concentration and, byimplication, its radiocarbon age can be determineddirectly from collected atmospheric CO2or indirectly

from terrestrial samples. For the archaeologicalmaterial listed in Table 1, terrestrial and aquaticsamples were analysed and the RRE is the radiocar-bon age difference between the two. For modernsamples, atmospheric 14C concentrations at time ofcollection are approximately known and listed inTable1(Levinet al. 2013; I Levin personal communi-cation). Atmospheric nuclear weapons tests, carriedout mostly between 1954 and 1963, have produced sig-nificant amounts of 14C. Thus, post 1954 AD atmos-pheric 14C concentrations are higher than 100 pMCand radiocarbon ageshave negative values.

ResultsRREs for aquatic species collected within Germanyboth from modern and archaeological contexts arelisted in Table 1. The data include both publishedresults (Fernandeset al. 2012a,2013;Philippsen andHeinemeier 2013) and new data obtained for thisstudy. Fig.1 shows the geographical distribution andthe range in RRE values for sample flesh, except forQuern-Neukirchen where only bivalve shell valuesare available.

Quern-Neukirchen : The site is located approximately200 m from the present day Baltic coast and is anexample of a shell midden site in use during the LateNeolithic period in northern Germany, known under

the term Dolchzeit (cal 22001700 BC)(Fernandes et al . 2012a). The RREs for Quern-Neukirchen were defined by comparison of radiocar-bon dates from mollusc shells of an unidentifiedspecies (KIA-43951 and KIA-43952) with those fromcollagen extracted from a bone of an unidentified her-bivore (KIA-43919) and red deer antlers (KIA-43920).Associated pairs mollusc-animal were collected in

close proximity to two well-defined contexts locatedat different depths. The RRE values for both associ-ated pairs are of similar magnitude (550 40 and570 40 yr).

Kiel Fjord: Four modern samples of blue mussels(Mytilus edulis) were collected from a beach locationin the Kiel Fjord. The following paired sample codesspecify flesh and shell of the same individual speci-men: KIA-44379 and KIA-44383; KIA-44380 andKIA-44384; KIA-44381 and KIA-44385; KIA-44382and KIA-44386. Mean RRE values for blue musselflesh (605 yr) and shell (600 yr) are similar and also

statistically similar to the mean RRE value observedat Quern-Neukirchen (560 yr).

Zauschwitz: The site of Zauschwitz is located in theimmediate vicinity of the Weie Elster River. The sitecontains graves of the Salzmnde type (a variant of theNeolithic Funnel Beaker culture). Large contempora-neous deposits of freshwater mussels were found,suggesting a significant human consumption ofaquatic food groups in the past. Two individualsburied with Salzmnde-type pottery were radiocarbondated significantly older than the established pottery

chronology and constitute examples of humandietary 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

Figure 1 Distributionof sites within Germany forwhichlocal (archaeological andmodern) 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.


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shells showed large RRE values (1815 30 and1515 30 yr). However, RREs of flesh for the samespecimens were significantly lower (1460 30 and1330 30 yr, respectively). An archaeological cattlebone (KIA-41496) was recovered from a shelldeposit at the same depth as a bivalve shell of an uni-

dentified species (KIA-41497). Comparison of theradiocarbon dates obtained from this pair gives aRRE of 1765 30 yr, a value similar to thoseobserved for modern shells.

Lake Rosenfeld: Lake Rosenfeld is a dammed lakelocated ca. 5 km southeast of the Kiel Fjord. Asingle specimen of swollen river mussel (Uniotumidus) had a RRE value of 895 30 yr for theflesh (KIA-44405) and 730 35 yr (KIA-44404) forthe shell.

Lake Ostorf: Within Lake Ostorf the small island ofTannenwerder contains a Neolithic cemetery (sodefined by the presence of Funnel Beaker pottery)where several radiocarbon dated individuals showed

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 fromLevin 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 Lab code Age ( yr BP) 13C () 15N () RRE (yr)

Site: Quern-Neukirchen (a)3660 30*1 Mollusc (shell) KIA-43951 4210 30 550 40

3650 25*2

Mollusc (shell) KIA-43952 4220 30

570 40Site: Kiel Fjord (a)

ca. 320 Blue mussel (flesh) KIA-44379 290 30 221 02 68 02 610 35ca. 320 Blue mussel (shell) KIA-44383 350 40 14 02 670 45ca. 320 Blue mussel (flesh) KIA-44380 260 30 226 02 74 02 580 35ca. 320 Blue mussel (shell) KIA-44384 200 20 11 02 520 30ca. 320 Blue mussel (flesh) KIA-44381 280 30 219 02 75 02 600 35ca. 320 Blue mussel (shell) KIA-44385 275 20 15 02 595 30ca. 320 Blue mussel (flesh) KIA-44382 315 25 223 02 73 02 635 30ca. 320 Blue mussel (shell) KIA-44386 300 30 06 02 620 35

Site: Zauschwitz (a)ca. 320 Duck mussel (flesh) KIA-40107-1 1140 20 306 02 126 02 1460 30ca. 320 Duck mussel (shell) KIA-40107-1 1495 20 152 02 1815 30ca. 320 Duck mussel (flesh) KIA-40107-2 1010 20 303 02 127 02 1330 30ca. 320 Duck mussel (shell) KIA-40107-2 1195 20 147 02 1515 304460 40*3 Bivalve shell KIA-41497 6225 45 1765 60

Site: Lake Rosenfeld (a)ca. 320 Swollen river mussel (flesh) KIA-44405 575 20 285 02 94 02 895 30ca. 320 Swollen river mussel (shell) KIA-44404 410 30 79 02 730 35

Site: Lake Schwerin (b)ca. 320 Pike (flesh) KIA-46304 270 20 197 02 132 02 590 30ca. 320 Bream (flesh) KIA-46305 190 30 194 02 115 02 510 35ca. 320 Eel (flesh) KIA-46306 260 25 147 02 128 02 580 30ca. 320 Eel (flesh) KIA-46307 625 25 206 02 170 02 945 30ca. 320 Zebra mussel flesh KIA-46310 230 20 271 02 81 02 550 30ca. 320 Zebra mussel shell KIA-46310 210 30 36 02 530 35

Site: Lake Ostorf (b)ca. 320 Zebra mussel flesh KIA-46311 1120 20 278 02 146 02 1440 30ca. 320 Zebra mussel shell KIA-46311 1060 30 75 02 1380 35

Site: Alster (c)ca. 440 Bivalve shell AAR-11460 1214 34 132 01 1654 40

ca.

440 Snail shell AAR-11461 433 32 154 01

873 35ca. 440 Roach (bone collagen) AAR-11462 223 29 255 01 122 03 663 35

Site: Trave (c)

ca. 440 Roach (bone collagen) AAR-11394 285 32 259 01 149 04 725 35ca. 440 Roach (bone collagen) AAR-11396 244 28 242 01 153 10 684 35ca. 420 Spined loach (flesh) AAR-12875 1664 39 272 01 156 03 2084 45ca. 420 Crayfish (flesh) AAR-12876 1365 40 279 05 119 20 1785 45ca. 420 Roach (flesh) AAR-12878 67 32 223 01 149 02 487 35

Site: Rebbe (d)ca. 240 Trout (flesh) KIA-49800 160 40 272 02 130 03 400 45

ca. 240 Roach (flesh) KIA-49801 305 40 277 02 134 03 545 45ca. 240 Bream (flesh) KIA-49802 345 40 289 02 131 03 585 45

Site: SW (d)ca. 240 Common rud (flesh) KIA-49820 260 40 277 02 79 03 20 45

ca. 240 Common rud (flesh) KIA-49821 120 40 275 02 79 03 120 45ca. 240 Bream (flesh) KIA-49822 135 40 306 02 55 03 105 45

Data were compiled from the following studies: (a) Fernandes et al. (2012a), (b) Fernandes et al. (2013), (c)Philippsen andHeinemeier (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.




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large dietary RREs (Olsenet al. 2010). A single speci-men (KIA-46311) of zebra mussel (Dreissena polymor-pha) collected from Lake Ostorf had a RRE of 144030 yr for the flesh and a similar value (1380 35 yr)for the shell.

Lake Schwerin: Lake Schwerin is a large lake (areaca. 62 km2) located approximately 15 km to thenortheast of Lake Ostorf. Four fish specimens (KIA-46304, KIA-46305, KIA-46306 and KIA-46307) ofpike (Esox lucius), bream (Abramis brama) and eel(Anguilla anguilla) collected from Lake Schwerin hadREE values between 510 30 and 945 35 yr. Asingle specimen (KIA-46310) of zebra mussel (D. poly-

morpha) had RRE values of 550 30 yr for the fleshand 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 fromceramic shards, showed, in some instances, significant

RREs, indicating that these contained freshwateringredients (Philippsen and Heinemeier 2013). Asingle fish specimen (AAR-11462), collected fromthe river Alster, showed a RRE of 663 35 yr. Anunidentified freshwater snail shell (AAR-11461) hada RRE of 873 35 yr, whereas an unidentified bivalveshell (AAR-11460) had a RRE of 1654 40 yr. Aspecimen of roach (Rutilus rutilus) had a RRE,measured in bone collagen, of 663 35 yr.

River Trave: The Erteblle site of Schlamersdorf islocated by the Trave. From this site, radiocarbon

dates on pottery food crusts suggested the presenceof significant RREs (Philippsen and Heinemeier2013). Four fish specimens (AAR-11394, AAR-11396, AAR-12875 and AAR-12878) of roach (R.

Table 2 Examples of past humans showing significant dietary RREs. There are no 15N valuesavailable for the individuals listed

from the study byLillie et al. (2009). The value between brackets represents an average of15N valuesreported in the same study

for Neolithic individuals within the same region

Sample code/ID Location Time period 13C () 15N () RRE (yr)

Study:Lanting and van der Plicht (1998)Dirk VI Low countries Medieval 189 106 340 32*Hadewig Low countries Medieval 198 109 227 46*

Gerlach Low countries Medieval 196 110 282 32*Study:Lillie et al. (2009)

OxA-17495 Ukraine Neolithic 225 (113) 251 49OxA-17499 Ukraine Neolithic 188 (113) 472 49OxA-17502 Ukraine Eneolithic 208 (113) 111 60

Study:Higham et al. (2010)OxA-9386 Russian North Caucasus Medieval 168 99 313 48OxA-9387 Russian North Caucasus Medieval 176 106 140 46OxA-14774 Russian North Caucasus Medieval 173 117 85 40OxA-9390 Russian North Caucasus Medieval 167 113 334 54OxA-9391 Russian North Caucasus Medieval 165 103 205 56OxA-9388 Russian North Caucasus Medieval 173 103 96 47OxA-9389 Russian North Caucasus Medieval 131 93 108 47OxA-14777 Russian North Caucasus Medieval 152 114 52 48

Study:Ascough et al. (2012)SUERC-2018 Iceland Medieval 193 100 120 49SUERC-2016/SUERC 2660 Iceland Medieval 191 90 212 35

Study:Olsen et al. (2010)UtC-8173 Ostorf Neolithic 168 52UtC-7447 Ostorf Neolithic 206 128 264 61UtC-7440 Ostorf Neolithic 22.0 139 288 54UtC-7445 Ostorf Neolithic 208 134 31 56UtC-7446 Ostorf Neolithic 208 137 103 59UtC-7448 Ostorf Neolithic 209 142 327 80UtC-8180 Ostorf Neolithic 20.0 13.0 435 51UtC-8181 Ostorf Neolithic 199 133 397 51AAR-9752 Ostorf Neolithic 206 132 401 57UtC-7449 Ostorf Neolithic 198 152 835 65UtC-8179 Ostorf Neolithic 194 153 552 60

Study:Cook et al. (2001)OxA-8502 Romania Mesolithic 196 132 510 117

OxA-8547 Romania Mesolithic 193 139 470 247OxA-8581 Romania Mesolithic 195 151 450 300OxA-8583 Romania Mesolithic 185 150 532 127

OxA-8548 Romania Mesolithic 182 153 345 116Study:Fernandes et al. (2012a)

KIA-37858 Zauschwitz (Germany) Neolithic 200 105 >300KIA-37859 Zauschwitz (Germany) Neolithic 203 92 >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.




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rutilus) and spined loach (Cobitis taenia) together witha crayfish (AAR-12876), collected from the Trave, hadRREs between 487 35 and 2084 45 yr.

River Rebbe: The sample collection site is located at12 km from the Funnel Beaker collective grave site ofOdagsen. Radiocarbon ages of some human remainsrecovered from Odagsen may suggest possible dietaryRREs (Fernandeset al. 2012b). The three fish speci-mens (KIA-49800, KIA-49801 and KIA-49802) oftrout (Salmo trutta), roach (R. rutilus) and bream (A.brama) collected from the river Rebbe had radiocar-bon ages between 400 45 and 585 45 yr.

Schloss Wilhelmsthal: The site is located 21 km eastof the Neolithic collective grave site of Calden, wherefish remains were identified in the interior of themonument (Raetzel-Fabian 2000). Three fish speci-mens (KIA-49820, KIA-49821 and KIA-49822) ofcommon rud (Scardinius erythropthalmus) and bream

(A. brama) were collected from an artificial lakewithin Schloss Wilhelmsthal grounds. This lake is fedby a natural stream. However, contrary to initialreports, it was later established that the local fish werebeing fed by humans and thus had a significant intakeof terrestrial food groups. RRE values for the threespecimens varied between 20 45 and 120 45 yr.

DiscussionRREs within Germany

The global average marine surface reservoir effect is

approximately 400 yr, while the radiocarbon resultslisted in Table1, for both coastal and inland locationswithin Germany, show significantly larger RREvalues. Excluding the results for SchlossWilhelmsthal, that probably include a terrestrial dietprovided by humans, the lowest RRE was recordedin the Rebbe (400 45 yr) whereas the largest REEwas recorded for the Trave (2084 45 yr). Theseresults confirm previous observations that showedthat for inland locations, local RREs can be bothvery large and variable (Lanting and van der Plicht1998; Keaveney and Reimer 2012). It is possible to

directly compare past and modern RREs for thesites of Zauschwitz and Quern-Neukirchen. Themean RRE value for Quern-Neukirchen, obtainedfrom archaeological samples, was 560 yr, a valuesimilar to the mean value observed for the modernshell samples collected from the Baltic coastline atthe Kiel Fjord (606 yr). In the case of Zauschwitz, asingle bivalve archaeological shell had a RRE of1765 35 yr, a value similar to those observed formodern samples (1515 30 and 1815 30 yr). Inspite of this, some caution is necessary when inferring

past RREs from modern values. Temporal changes inclimatic or hydrological patterns and in catchmentareas will influence local RREs (Ingram andSouthon 1996;Stein et al. 2004;Ascough et al. 2007,

2010). In addition to natural variability, human inter-vention in shaping the landscape and in modifying thecourse of streams and rivers might also have causedsignificant changes to the local RREs. Another issueto consider in modern RREs is the possible contri-bution of groundwater having a 14C enriched signaldue to the nuclear tests of previous decades.Archaeological samples might also present some diffi-culties in determining the relevant local RREs forhuman dietary studies. The Zauschwitz andRosenfeld radiocarbon measurements on modernbivalve samples show significant differences betweenbivalve meat and shell. Bivalve shell is mainly precipi-tated from dissolved inorganic carbon (DIC) whilecertain bivalve species metabolise dissolved organiccarbon (DOC) or particulate organic carbon (POC)to build their flesh. These fractions (POC, DOC andDIC) may show significantly different 14C concen-

trations and this justifies the observed radiocarbonage differences between meat and shell for bivalvesfrom Lake Rosenfeld and Zauschwitz (Fernandeset al. 2012a). No such differences were observed forthe zebra mussels collected from Lakes Ostorf andSchwerin given that zebra mussels have a diet special-ised in phytoplankton, which in turn obtains itscarbon in underwater photosynthesis from DIC(Baker et al. 1998). Research is underway to verifywhether radiocarbon dates obtained from the proteinfraction in bivalve shells (conchiolin), measured as

an alternative to bivalve shell carbonate, provide aradiocarbon age similar to bivalve flesh.

Radiocarbon as a Dietary Proxy

Aquatic species-dependent variability in dietary habitsand the temporal and spatial variability in local RREsrepresent an important challenge for the use of 14Cmeasured in humans to obtain precise quantitativeestimates on the intake of aquatic food groups.However, given the large RRE values observed,especially for inland locations, 14C measured inhumans can be used to detect the consumption of

aquatic food groups when traditional isotopic analysiscannot provide an unambiguous result.

Table2provides a list of examples of human dietaryRREs from different locations and time periods. Largelocal RREs and a similar isotopic baseline of fish andanimal isotopic values (13C and15N) are common toall listed sites, with the exception of the Russian NorthCaucasus due to the presence of C4-plants in the localenvironment (Higham et al. 2010). Human dietaryRREs were established by comparison of radiocarbondates from human bone collagen with associated inde-

pendent reference dates. In the case of the study byLanting and van der Plicht (1998)the time of deathwas known from historical records. For the studiesby Cook et al. (2001), Lillie et al. (2009), Higham




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et al. (2010),Olsen et al. (2010)andAscough et al.(2012) individual human dietary RREs were deter-mined by radiocarbon dating of associated animalremains. The study by Higham et al. (2010) alsoreported the chronology of buried artefacts (ceramicsand coins) that provided ages similar to animal radio-carbon dates. For the study by Fernandes et al. (2012a)the reference date is based on ceramic typology and aprecise estimate cannot be provided.

Measurements of isotopic ratios, 13C and 15N, inhuman bone collagen are often used as proxies toprovide dietary estimates. In predominantly C3-plantenvironments, humans having a fully terrestrial diettypically show 13Ccollagen values of ca. 20.Human intake of marine aquatic foods can be detectedwhen 13Ccollagenvalues significantly higher than ca.20 are observed (Schoeninger et al. 1983). Forinstance, the comparison of 13C bone collagen

values in Neolithic Europeans with the precedingMesolithic populations shows clear differences(Tauber 1981; Richards et al . 2003a), and theseresults have been interpreted to signify a completeshift towards a terrestrial-based diet. However, C4-plants (e.g. millet and maize) have 13C signalssimilar to marine foods and as such high 13Ccollagenvalues may also be observed in some inland locations.This is illustrated in Table 2 by the individuals from theRussian North Caucasus (Highamet al. 2010) that allshow 13Ccollagenvalues above 18. Furthermore,

even when restricted to C3-type food groups, the isoto-pic difference between terrestrial and freshwater 13Csignals may be too small to offer the possibility of dis-tinguishing a terrestrial from a freshwater contribution.In some instances, the consumption of freshwater foodgroups may be detected by 13Ccollagen values lowerthan ca. 20 (Lanting and van der Plicht 1998).Table 2 shows that the individual with code OxA-17495 from a Neolithic site in the Dnieper basin(Ukraine) has a 13Ccollagenvalue of225. In spiteof this example, the majority of individuals listed inTable2originating from predominantly C3 environ-

ments show13Ccollagenvalues of ca. 20.Consumption of freshwater food groups can, in

some instances, be established through the measure-ment of15N in human bone collagen. A compilationof published 15N data, with major outliers removed,from Late Mesolithic and Neolithic central and north-ern European sites provides average values for animalbone collagen of 15N= 59 14 (n= 268) and15N= 88 15 for fishbone collagen (n = 27)

(Boric et al. 2004; Ogrinc and Budja 2005; Bslet al. 2006;Drrwchteret al. 2006;Bocherens et al.

2007; Hedges et al. 2008;Oelze et al. 2011;Stevenset al. 2012). The higher 15N values observed for fishcompared with land animals constitute the basis forthe use of the 15Ncollagen proxy to detect the

consumption of freshwater aquatic food groups.Typical 15Ncollagen values for European Neolithicpopulations are within the range 811(Schoeninger et al . 1983; Richards et al . 2003b;Fischer et al. 2007) with the lower limit interpretedas dietary protein predominantly from plant foodsand the upper limit interpreted as dietary protein pre-dominantly from animal foods (Richards andTrinkaus 2009). Thus, human 15Ncollagen valuesabove 12are often interpreted as representing a sig-nificant intake of aquatic foods. Pre-historic individ-uals listed in Table2 from the studies byOlsen et al.(2010) in Ostorf (Germany) and by Cook et al .(2001) at the Iron Gates (Romania) all show15Ncollagenvalues above 12. However, the remain-

der of the case studies listed in Table 2all show indi-viduals with 15Ncollagen values between 90 and117 (Lanting and van der Plicht 1998; Higham

et al. 2010; Ascough et al. 2012; Fernandes et al.2012a).

The interpretation of 15Ncollagen values needs toconsider that there are in fact multiple food groupsand that the isotopic signals of the different groupsmay differ significantly. In addition, the real uncer-tainty that should be considered in the interpretationof human 15Ncollagen values should not be thetypical experimental uncertainty (ca. 02) but thenatural 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 simpleunivocal correspondence between a certain isotopicvalue and the intake of a particular food group.Instead a single 15Ncollagen value should be viewedas resulting from multiple possible mixture pro-portions each with an assigned probability. To illus-trate that different dietary mixing proportions areconsistent with the same human 15Ncollagensignal, asimple linear model is presented. This model takes asisotopic baseline for protein (the source of dietarynitrogen) 15N values from food groups reportedabove that were available to Late Mesolithic and

Neolithic populations from central and northernEuropean sites (15Nanimal= 59;

15Nfish= 88).As a reference for plant 15N, two possible valueswere considered. A value of 2 represents a typical15N value for edible European plants during the

Holocene (Richards and Trinkaus 2009). However,recent research has shown that the practice of manur-ing in cereal crops was prevalent during the Neolithicand a value of 45 should be used as reference formanured plants (Bogaard et al. 2013). The linearmodel considers the contribution of paired protein

sources weighed by percentage of intake (unmanuredplant protein vs. fish protein; manured plant proteinvs. fish protein; fish protein vs. animal protein). Ahuman diet-to-collagen offset of 55 was added to




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the calculated isotopic mix values. This value rep-resents a consensus value derived from publisheddata 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 similarto the reference value (ca. 6) derived from isotopic

values measured in human red blood cells during acontrolled feeding experiment (OConnell et al .2012). A value of 55for the diet-to-collagen offsetis higher than the traditionally used 34 range(Hedges and Reynard 2007); however, it should benoted that the higher value will, if incorrect, underes-timate the contribution of fish to human diet.

Model outputs for human 15Ncollagen, shown inFig.2, also include estimated human dietary RREsfor different radiocarbon ages of aquatic food groups(500, 1000 and 1500 yr). To estimate the actual contri-bution of carbon from aquatic food sources towardshuman bone collagen it is also necessary to considerthat part of the collagen carbon (ca. 25%) is routedfrom dietary lipids and carbohydrates (Fernandeset al. 2012c). Thus in Fig.2values given for humandietary RREs, corresponding to 15Ncollagen= 12,represent both a non-routed model and routed model

(between brackets). For the routed model it isassumed that the only source of energy macronutrients(carbohydrates and lipids) is terrestrial. The compari-son of modelled results for the differently pairedfood groups clearly shows that the same range of15Ncollagenvalues, between 9 and 12(grey area), is

compatible with significantly different dietary contri-butions of selected food groups. Furthermore, theresults show that significant human dietary RREsmay be observed with low protein contributions fromaquatic food groups. These results are in agreement

with the human dietary RREs listed in Table2.The German RRE baseline shows the widespreadpresence of large local RREs. Although the variabilityin local RREs is too large to allow for precise dietaryestimates, the use of14C as an extra dietary proxy canprovide a valuable contribution in ancient human dietreconstruction studies. The question of completenessin the adoption of an agricultural subsistence strategyand the total, or not, abandonment of fish consump-tion across the MesolithicNeolithic transition is aprime example of such potential use.

ConclusionA first wide-ranging baseline of RREs has been estab-lished for Germany. Radiocarbon measurements onaquatic food species edible for humans obtainedfrom modern and archaeological contexts show thepresence of large (>400 yr) and variable local RREs.Observed variability does not allow the use of 14Cmeasured in human collagen to obtain precise quanti-tative estimates of dietary intake of aquatic foodgroups. However, observed large local RREs,especially in inland contexts, permit the use of 14C as

an extra dietary proxy that can detect the intake ofaquatic food groups when other standard isotopicproxies (13C and 15N) fail to provide unambiguousestimates.

Figure 2 Linear model results to estimate human 15Ncollagen

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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