Stable isotope insights ( δ 18 O, δ 13 C) into cattle and sheep husbandry at Bercy (Paris, France,...

Stable isotope insights (d18O, d13C) into cattleand sheep husbandry at Bercy (Paris, France,4th millennium BC): birth seasonality andwinter leaf foddering

Marie Balasse, Loıc Boury, Joel Ughetto-Monfrin and Anne Tresset

Bercy is a prehistoric village sited by the Seine river (Paris, France), whose main period of

occupation was dated to the very beginning of the 4th millennium BC. The animal subsistence

economy relied heavily on cattle husbandry, complemented by other species including sheep.

Cattle and sheep isotopic history (d13C, d18O) was investigated at the seasonal scale, through

sequential sampling in tooth enamel, providing new insights into seasonality of birth and diet.

Sheep were lambing in mid-spring, only slightly later than expected from what is observed

nowadays in temperate Europe at similar latitude. Cattle were born over a period spanning

approximately six months, which was an unexpected result compared with a two to three months

calving period in free-ranging cattle populations. The extension of the calving period might have

been related to seasonal food supplementation. Some cattle and some sheep fed on a 13C-

depleted resource in winter, potentially leafy fodder. A direct consequence of an extended

calving period would be the availability of cow milk, which would have covered the whole year at

Bercy. This is important information in a context where the exploitation of cattle milk by the human

community was highly suspected from the demographic management of the herd.

Keywords: cattle, sheep, carbon and oxygen stable isotope ratios, tooth enamel, birth seasonality, fodder

Introduction

Bercy is a large prehistoric village site sited on a bank

of the Seine river (Paris, France), excavated during

the 1990s under the direction of P. Marquis and Y.

Lanchon (Marquis 1991; Lanchon et al. 2000). Its

main occupation, attributed to the Chasseen septen-

trional culture, was dated to the very beginning of the

4th millennium BC. Excavation of the adjacent pa-

leochannel produced rich assemblages of faunal and

botanical remains. A pluridisciplinary research was

conducted in the 1990s in order to investigate the

environmental context of this agro-pastoral settlement

and evaluate its impact on the milieu. From paly-

nological (Leroyer 1998), anthracological (Pernaud

1997) and carpological (Dietsch 1996; Dietsch-

Sellami 2001) data, at the time of the Chasseen occu-

pation, the river bank was characterised by a riverine

forest dominated by the alder. A wet meadow had

developed on the marshy soils of the valley floor,

with a high frequency of ruderal species suggesting

that it was used for pasture. Alluvial terraces were

covered with a dense oak tree forest exploited by the

human community for fire wood and partly cleared

for agricultural purpose.

The animal subsistence economy (Tresset 1996;

1997) relied heavily on cattle husbandry, complemen-

ted by the raising of pig, sheep and goat. The hunted

wild fauna, dominated by red deer, also included roe

deer, aurochs, wild pig and small furry game.

Documented on the basis of tooth eruption and wear

stages, the mortality profiles of domestic animals

Marie Balasse (corresponding author), Joel Ughetto-Monfrin and AnneTresset, Museum national d’Histoire naturelle USM 303/CNRS UMR 7209‘Arche ozoologie, Arche obotanique: Socie t e s, Prat iques etEnvironnements’, Departement Ecologie et Gestion de la Biodiversite,case postale 56, 55 rue Buffon, 75005 Paris, France, e-mail: [email protected]; Loıc Boury, Universite de Strasbourg/CNRS UMR 7044‘Etude des civilisations de l’Antiquite’, ANTEA Archeologie, 11 rue deZurich, 68440 Habsheim, France.

� Association for Environmental Archaeology 2012Published by ManeyReceived July 2011; revised manuscript accepted January 2012DOI 10.1179/1461410312Z.0000000003 Environmental Archaeology 2012 VOL 17 NO 1 29

show that pigs and small stock were culled mainly

when reaching a good weight and not maintained to

older ages, suggesting a strategy oriented essentially

towards meat production. Cattle were raised for meat

but also for milk, as suggested by the presence of old

individuals, presumably cast milk cows, and a very

high mortality of young individuals reflecting prob-

ably a post-lactation slaughter (Tresset 1996; 1997;

Balasse et al. 1997; 2000; Balasse and Tresset 2002).

Previous stable isotope analyses of bone collagen

of large herbivores from Bercy provided a first

glimpse of the occupation of the landscape by wild

and domestic herbivores. From the d13C values

measured in bone collagen from these animals,

potentially reflecting the feeding on closed forest

understorey vegetation, it was suggested that domes-

tic cattle were raised in open pastures whereas coeval

aurochs fed within the dense forest (Balasse et al.

2000; Drucker et al. 2008). Red deer and roe deer had

a more opportunistic feeding strategy along the forest

edge (Drucker et al. 2008). Prolonging this effort in

the study of the stable isotope ecology of the

Neolithic settlement of Bercy, the present study

focuses on domestic cattle and sheep. Their isotopic

history (d13C, d18O) was investigated at the seasonal

scale, through sequential sampling in tooth enamel,

providing new insights into seasonality of birth and

diet.

Investigating birth seasonality in Neolithicdomestic stock

The onset and the length of the birth period in

domestic stock influences the annual cycle of pastoral

tasks. In mammals, synchronisation of the breeding

cycle to the most appropriate time for survival of

the young is driven by environmental variables. In

domestic sheep, within the period of fertility defined

by the photoperiodic cycle (Karsch et al. 1984),

the onset and the length of the breeding period

are determined by climatic, environmental (weather

conditions, food availability) and genetic parameters

(Hafez 1952; Lincoln et al. 1990; Rosa and Bryant

2003). Cattle do not experience anoestrus and are

fertile all the year round. However, in extensive

conditions their breeding cycle is driven by environ-

mental variables including the vegetation annual

cycle (Lecomte and Le Neveu 1986; Reinhardt et al.

1986). Although it is most likely that in Neolithic

temperate Europe the births of domestic cattle and

sheep were seasonal, how restricted the birth period

was still needed to be defined. Calving also coincides

with the initiation of lactation and this parameter

might have been especially important in a context of

dairying, as strongly suggested for cattle at Bercy

(Tresset 1996; 1997; Balasse et al. 1997; 2000).

Birth seasonality can be investigated from tooth

remains using analysis of enamel oxygen isotope

composition (Bryant et al. 1996a; 1996b). The oxygen

isotope composition (d18O) of the skeleton calcium

phosphate (bioapatite) is mainly linked to that of

ingested water, i.e. indirectly meteoric water (Land

et al. 1980; Longinelli 1984; Luz et al. 1984; D’Angela

and Longinelli 1990; Delgado Huertas et al. 1995). At

high and middle latitudes, the d18O of precipitation

varies seasonally with ambient temperature (Gat

1980). The seasonal variations are recorded in tooth

mineral during development and are not remodelled

once enamel mineralisation is completed. Recon-

struction of the seasonal cycle is therefore possible by

performing a sequential d18O analysis along the tooth

growth axis (Koch et al. 1989; 1995; Fricke and

O’Neil 1996; Stuart-Williams and Schwarcz 1997;

Kohn et al. 1998; Sharp and Cerling 1998; Sharma

et al. 2004; Nelson 2005; van Dam and Reichart 2009;

Bernard et al. 2009). Because the timing of tooth

development is fixed within a species, seasonality of

birth can be investigated through the study of inter-

individual variability in d18O changes along tooth

crown (Balasse et al. 2003; Nelson 2005; Balasse and

Tresset 2007; Britton et al. 2009; Henton et al. 2010;

Towers et al. 2011).

Investigating leaf foddering in Neolithictemperate Europe

Sequential sampling of tooth enamel also permits to

investigate animal diet at the seasonal scale, from the

analysis of carbon stable isotope composition (d13C)

of bioapatite. The potential of stable carbon isotope

analysis to investigate seasonality of diet is not wide,

especially in North-western Europe where C4 plants

are very rare (today: up to 1?2% of species, including

up to a quarter of invasive species originating in other

continents and mostly recently introduced to Europe;

Pyankov et al. 2010). However, it can still be used to

investigate seasonal consumption of plants from

closed areas, where the canopy effect would result

in lower d13C values (van der Merwe and Medina

1991), inherited in animals feeding on them.

The use of tree leaves to feed domestic cattle and

small stock was widespread in Europe until recent

history from Norway to the Mediterranean area, to

compensate for restricted availability of grass during

dry summers or snowy winters and/or over periods

of stalling (see Troels-Smith 1960; Rasmussen 1990;

Halstead and Tierney 1998; Thiebault 2005 for

archaeobotanical evidence and Lucas 1989 and

Balasse et al. Birth seasonality and winter leaf foddering

30 Environmental Archaeology 2012 VOL 17 NO 1

Kelly 1997 for historical accounts). The practice

was directly evidenced at middle Neolithic sites in

Switzerland, where botanical remains were retrieved

in cattle and small stock dung levels, attesting the use

of twigs and leaves from various tree species, some of

which might have been pollarded especially for the

purpose (Rasmussen 1989; 1990; 1993; Akeret et al.

1999). The common tree taxa recovered at these sites

(Corylus, Alnus, Betula, Salix, Fraxinus, Tilia) are

typical of open forest and their consumption might

not be detected through a d13C analysis. Never-

theless, Hedera helix L. and Ilex aquifolium L., men-

tioned in the early Irish literature (Kelly 1997) could

be detected provided they have been cut in deeper

forest, which is likely. The use of deciduous oak trees

as leafy fodder is also reported in recent history

(Halstead and Tierney 1998) and its over-representa-

tion in the anthracological assemblages of the early

and middle Neolithic horizons of sheepfold rock-

shelters from the alpine and circum alpine area in

France was interpreted as possibly reflecting the

harvesting of leafy fodder to feed caprines (Thiebault

2001; 2005; Delhon et al. 2008). When harvested in

a dense forest, leafy fodder might be marked by

the canopy effect. Although the use of leafy fodder

and the choice of the tree species are submitted to

cultural choices and environmental availability, this

possibility could be addressed at Bercy, where d13C

measured in bone collagen from wild large herbivores

(Drucker et al. 2008) suggested the presence of a

closed canopy in the surrounding environment, while

the presence of a dense oak forest was evidenced from

pollen remains (Leroyer 1998). Although previous

bone collagen d13C analysis suggested the feeding

of domestic stock in open areas, sequential stable

isotope analysis in teeth would permit the question of

leaf foddering to be investigated at a seasonal scale.

Material and methods

Five hemi-mandibles from sheep and ten hemi-

mandibles from bovines were selected, belonging to

different individuals. Sheep present at Bercy are

securely domestic, as the species has no wild ancestor

in Europe (Poplin 1979; Tresset et al. 2009). Within

the caprine tooth rows, they were distinguished from

goats from morphological criteria (Balasse and

Ambrose 2005; Payne 1985; Helmer 2000; Halstead

et al. 2002). Both wild and domestic bovines occur

in the faunal assemblage of Bercy; this has been

suspected from the osteometric analysis (Tresset

1997) but also demonstrated on molecular grounds

(Edwards et al. 2004; Bollongino et al. pers. comm.).

Assessing the wild or domestic status of a bovine

from a tooth row basing on osteometric criteria only

is problematic. An attempt was made to use size

criteria on the third molar. The length and the width

of the crown were measured on 47 lower third molars

(M3) from Bercy. Results are reported in Fig. 1. The

M3 lengths range from 30?5 to 45?6 mm and the

widths from 14?5 to 19?1 mm. The five M3, the length

of which is less than 34 mm, are characterised by the

atrophy of the third lobe. BQSBos2M3 is one of

them. All other M3 included in the stable isotope

study have a length comprised between 35 and 40 mm

and a width comprised between 15 and 17 mm

(Fig. 1), except for BQSBos11M3, which is the largest

of the whole assemblage with a 45?6 mm length and a

19?1 mm width. It is the most likely candidate for a

wild status, although we are fully aware that the size

criteria might sometimes be misleading for the dis-

tinction between wild and domestic bovines (Vigne

et al. 2007; Tresset et al. 2009).

The sampling was performed on the posterior lobe

of the M2 in sheep and on the anterior lobe of the M3

in cattle. Enamel surfaces were cleaned with a gentle

brush and the external surface removed by drilling

with a tungsten drill bit. A sequential sampling was

performed by drilling with diamond burr on the

buccal side of the teeth, perpendicularly to the crown

growth axis, going through the whole thickness of the

Figure 1 Results from measurements of the length and

width of cattle lower third molars. Bos11M3 is

the largest tooth from the assemblage Figure 2 Enamel sampling procedure in cattle (a) and

sheep (b) third and second molars respectively


Environmental Archaeology 2012 VOL 17 NO 1 31

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http://www.maneyonline.com/action/showImage?doi=10.1179/1461410312Z.0000000003&iName=master.img-001.jpg&w=227&h=127

enamel layer (Fig. 2). Because of the complexity

of the enamel deposition and maturation processes,

this sampling procedure does not permit isolation of

discrete growth increments and time segments, but

the sequence measured was shown to be chronolo-

gical in sheep and cattle teeth, as evidenced by

unambiguous seasonal cycles in the oxygen isotope

values (Balasse 2003; Balasse et al. 2003). Enamel

samples weighed between 4?5 mg and 8 mg. The

number of samples taken from one tooth varies from

18 to 20 on the cattle third molars, 13 to 18 on the

sheep second molars, for a total number of 267

samples.

Enamel powder was treated for bioapatite extrac-

tion as described in Balasse et al. (2002). Enamel was

treated with sodium hypochlorite 2–3% (24 h) to

remove organic matter, and then with 0?1 M acetic

acid (4 h, 0?1 ml/mg) to remove exogenous carbonate.

Bioapatite samples weighing around 600 mg were

reacted with 100% phosphoric acid at 70uC in

individual vessels in an automated cryogenic distilla-

tion system (Kiel IV device) interfaced with a Delta V

Advantage isotope ratio mass spectrometer. Over the

period of analysis of these bioapatite samples, 155

analyses of our laboratory internal carbonate stan-

dard (Marbre LM) gave a mean d13C value of

2?13¡0?02 % (1s) (theoretical value normalised to

NBS 19 5 2?13 %) and a mean d18O value of

21?79¡0?07% (1s) (theoretical value normalised to

NBS 19 5 21?83 %). The analytical precision within

each run, calculated from 5 to 12 measurements of

the standard Marbre LM, varies from 0?01 to 0?03%for d13C and from 0?03 to 0?07% for d18O.

Results

Results are reported in Tables 1 and 2 and Figs 3 and

4. Stable oxygen isotope values vary between 26?9

and 21?9% in cattle and between 27?6 and 21?2%in sheep. In both species, the d18O values show

sinusoidal variations with time along tooth crown,

with amplitude of variation of 1?5 to 4?0% in cattle

and 2?5 to 4?8% in sheep. They most likely reflect the

seasonal cycle in ambient temperature. A whole

annual cycle is recorded over approximately 40 to

50 mm in the cattle M3 (Fig. 3). In sheep, the M2

crowns were worn up to a height of approximately 25

mm or less. The cycle measured over this length was

incomplete; half an annual cycle was measured over

approximately 15 mm (Fig. 4). Stable carbon isotope

values vary between 214?7 and 211?6% in cattle and

between 215?3 and 210?3% in sheep. Within a

tooth, the amplitude of variation is between 0?8 and

2?0% in cattle and between 1?5 and 2?7% in sheep.

When seasonal changes are recorded in cattle teeth,

the highest d13C values occur in the summer and the

lowest in the winter. The same trend is observed in

the sheep teeth.

DiscussionCattle birth seasonality

Birth seasonality is assessed from inter-individual

variability in the position in the tooth crown where

the lowest and highest d18O are measured. In order to

identify these positions while erasing analytical noise,

the measured data sequences were modelled using the

following equation mainly based on a cosine function

(Balasse et al. 2011):

d18Om5A. cos (2P? (x-x0)/X)zM

where:

d18Om is the modelled d18O;

x is the distance from enamel-root junction;

X is the period (in mm), corresponding to the

length of tooth crown potentially formed over a

whole annual cycle;

A is the amplitude (5 (max-min)/2) (in %);

x0 is the delay (mm), which depends on the time of

the year when tooth growth started; d18O attains

maximum value when x 5 x0;

M is the mean (5 (maxzmin)/2) expressed in %.

Fig. 5 shows an example of the model for

BQSBos1M3 detailing the parameters. The best fit

of the model parameters to the measured data was

determined using an iterative method and a mini-

misation of the sum of the square of the difference

between the model and the measurements (method of

least squares). Results from the calculation of the best

fit for combined variations of X, A, x0 and M are

shown in Table 3 and in Fig. 3. The fitting of the

model to the dataset is estimated using the Pearson’s

correlation coefficient. This coefficient was always

>0?94 (Table 3), demonstrating that the model is in

each case very close to the dataset. The period X

varies from 39?4 to 44?7 mm for most individuals.

BQSBos5M3 and BQSBos7M3 differ from the others

with respectively shorter (31?0 mm) and larger (57?3

mm) periods, although results from the modelling of

BQSBos7M3 may not be accurately reflecting the

cycle (the first three samples had to be excluded from

the calculation because they would force the model

to unsuitable solutions). Variability in the modelled

period reflects variability in growth rate, related to

tooth size. It is necessary to remove this variability

before comparing the position in tooth crown where

the lowest and highest d18O values were measured in

the different specimens. Because the period represents

always the same duration of time (one year) in all



Tab

le1

Sta

ble

carb

on

(d13C

)an

do

xyg

en

(d18O

)is

oto

pe

valu

es

measu

red

into

oth

en

am

el

bio

ap

ati

tefr

om

catt

lelo

wer

thir

dm

ola

rs(M

3).

d13C

an

dd1

8O

valu

es

are

exp

ressed

in%

.S

am

ple

sa

relo

ca

ted

ine

ac

hto

oth

cro

wn

by

the

dis

tan

ce

fro

mth

ee

na

me

l–

roo

tju

nc

tio

n(m

m).

D5

intr

a-t

oo

tha

mp

litu

de

of

va

ria

tio

n

Sam

ple

mm

d13C

VP

DB

d18O

VP

DB

Sam

ple

mm

d13C

VP

DB

d18O

VP

DB

Sam

ple

mm

d13C

VP

DB

d18O

VP

DB

Sam

ple

mm

d13C

VP

DB

d18O

VP

DB

sam

ple

mm

d13C

VP

DB

d18O

VP

DB

BQ

SB

os1

M3-1

49

?52

12

?82

5?7

BQ

SB

os2

M3-1

47

?42

13

?72

5?5

BQ

SB

os3

M3-1

45

?72

12

?72

4?6

BQ

SB

os4

M3-1

46

?92

12

?62

5?4

BQ

SB

os

5M

3-1

46

?92

12

?62

5?4

BQ

SB

os1

M3-2

47

?32

13

?02

5?8

BQ

SB

os2

M3-2

45

?02

13

?72

5?8

BQ

SB

os3

M3-2

43

?62

12

?52

5?1

BQ

SB

os4

M3-2

44

?62

12

?72

5?5

BQ

SB

os

5M

3-2

44

?62

12

?72

5?5

BQ

SB

os1

M3-3

45

?22

13

?72

6?1

BQ

SB

os2

M3-3

42

?92

13

?92

5?7

BQ

SB

os3

M3-3

41

?32

12

?32

5?4

BQ

SB

os4

M3-3

42

?32

12

?72

5?6

BQ

SB

os

5M

3-3

42

?32

12

?72

5?6

BQ

SB

os1

M3-4

42

?92

13

?82

6?5

BQ

SB

os2

M3-4

40

?52

13

?82

6?1

BQ

SB

os3

M3-4

38

?92

12

?32

5?4

BQ

SB

os4

M3-4

39

?92

12

?62

5?4

BQ

SB

os

5M

3-4

39

?92

12

?62

5?4

BQ

SB

os1

M3-5

40

?82

14

?12

6?5

BQ

SB

os2

M3-5

38

?12

13

?92

6?1

BQ

SB

os3

M3-5

36

?72

12

?42

5?5

BQ

SB

os4

M3-5

37

?52

12

?52

4?8

BQ

SB

os

5M

3-5

37

?52

12

?52

4?8

BQ

SB

os1

M3-6

38

?42

14

?22

6?9

BQ

SB

os2

M3-6

35

?82

14

?12

6?9

BQ

SB

os3

M3-6

34

?22

12

?52

5?7

BQ

SB

os4

M3-6

35

?12

12

?42

4?5

BQ

SB

os

5M

3-6

35

?12

12

?42

4?5

BQ

SB

os1

M3-7

36

?32

14

?42

6?5

BQ

SB

os2

M3-7

33

?32

14

?22

6?4

BQ

SB

os3

M3-7

32

?02

12

?52

5?8

BQ

SB

os4

M3-7

32

?82

12

?32

4?1

BQ

SB

os

5M

3-7

32

?82

12

?32

4?1

BQ

SB

os1

M3-8

34

?22

14

?32

6?8

BQ

SB

os2

M3-8

30

?92

14

?42

6?0

BQ

SB

os3

M3-8

29

?72

12

?62

6?0

BQ

SB

os4

M3-8

30

?62

12

?02

3?7

BQ

SB

os

5M

3-8

30

?62

12

?02

3?7

BQ

SB

os1

M3-9

32

?02

14

?32

6?4

BQ

SB

os2

M3-9

28

?22

14

?02

6?4

BQ

SB

os3

M3-9

27

?42

12

?62

6?5

BQ

SB

os4

M3-9

28

?22

11

?92

3?5

BQ

SB

os

5M

3-9

28

?22

11

?92

3?5

BQ

SB

os1

M3-1

029

?92

14

?12

5?8

BQ

SB

os2

M3-1

025

?82

13

?92

5?6

BQ

SB

os3

M3-1

025

?22

12

?72

6?5

BQ

SB

os4

M3-1

026

?22

12

?02

3?7

BQ

SB

os

5M

3-1

026

?22

12

?02

3?7

BQ

SB

os1

M3-1

128

?02

13

?52

5?2

BQ

SB

os2

M3-1

123

?42

13

?42

4?5

BQ

SB

os3

M3-1

123

?22

12

?72

6?8

BQ

SB

os4

M3-1

123

?92

12

?12

3?5

BQ

SB

os

5M

3-1

123

?92

12

?12

3?5

BQ

SB

os1

M3-1

225

?52

13

?62

4?6

BQ

SB

os2

M3-1

221

?22

13

?32

3?9

BQ

SB

os3

M3-1

220

?82

12

?92

6?9

BQ

SB

os4

M3-1

221

?72

12

?12

3?7

BQ

SB

os

5M

3-1

221

?72

12

?12

3?7

BQ

SB

os1

M3-1

323

?42

13

?42

4?1

BQ

SB

os2

M3-1

318

?92

13

?22

3?6

BQ

SB

os3

M3-1

318

?82

12

?72

6?5

BQ

SB

os4

M3-1

319

?82

12

?02

3?9

BQ

SB

os

5M

3-1

319

?82

12

?02

3?9

BQ

SB

os1

M3-1

421

?52

13

?22

3?7

BQ

SB

os2

M3-1

416

?72

12

?92

3?4

BQ

SB

os3

M3-1

416

?52

13

?12

6?1

BQ

SB

os4

M3-1

417

?82

12

?12

3?9

BQ

SB

os

5M

3-1

417

?82

12

?12

3?9

BQ

SB

os1

M3-1

519

?52

13

?02

3?7

BQ

SB

os2

M3-1

514

?32

12

?92

3?5

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D5

2?5

3?4

D5

2?2

4?1



individuals, it was used to normalise the measured

distances. In Fig. 6, the location in the tooth crown

where the highest d18O values occur for each

individual was normalised with their respective

period X (x0/X). Values for x0/X are evenly distrib-

uted between 0?12 and 0?62 (Table 3 and Fig. 6a),

meaning that the highest d18O values were measured

at 12 to 62% of the end of the recorded periodic cycle,

the end being defined as when the cycle reaches the

enamel-root junction. At a monthly scale (a whole

period represents 12 months), a delay of approxi-

mately six months over the annual cycle separates the

measure of the highest d18O values in BQS Bos3M3

and BQS Bos9M3 (Fig. 6b), meaning that if the

Figure 3 Results from sequential analysis of stable carbon (d13C, black symbols) and oxygen (d18O, white symbols) iso-

tope ratios in cattle tooth enamel bioapatite. Samples are located in each tooth crown by their distance from

the enamel-root junction (erj). The grey line shows the modelled d18O values




population analysed in this study is representative of

the herd, cattle births were occurring over approxi-

mately six months.

The dataset acquired at Bercy was then compared

to results obtained following the same analytical

procedure on cattle teeth from the Neolithic farm-

stead of the Knap of Howar (Orkney, ca. 3600 BC;

Ritchie 1983), using the dataset published in Balasse

and Tresset (2007). Although only five individuals

could be analysed from this assemblage, interpreta-

tion of the raw data suggested a strong seasonality of

birth over a very restricted time of the annual cycle

(Balasse and Tresset 2007). From the model, the

period calculated for the cattle M3 from the Knap of

Howar varies from 34?2 to 46?8 mm, reflecting again

significant variability in tooth size and growth rate

(Table 3). Fig. 6c compares individuals after normal-

isation of x0 with the period X calculated for each

individual. At the Knap of Howar, the highest d18O

values in cattle M3 were measured at 45 to 56% of the

end of the periodic cycle recorded in the tooth. Over

the annual cycle, the birth of these five individuals

was distributed over less than two months (Fig. 6d).

Compared to the Knap of Howar, the Bercy

dataset therefore appears to reflect a less restricted

period of birth for cattle. As discussed elsewhere

(McCormick 1998; Balasse and Tresset 2007), con-

straining climatic conditions might explain the very

restricted birth period observed at the Knap of

Howar, which might either have actually led to a

restricted period of births and/or might have resulted

in the death of the calves born unseasonably: the

analyses performed on the third molar only reflect the

birth season of cattle who survived their second year,

over which the tooth is formed (Brown et al. 1960).

Observations on primitive cattle breeds free ranging

Figure 4 Results from sequential analysis of stable carbon (d13C, black symbols) and oxygen (d18O, white symbols) iso-

tope ratios in sheep tooth enamel bioapatite. Samples are located in each tooth crown by their distance from

the enamel-root junction (erj). The grey line shows the modelled d18O values

Figure 5 Application of the model to the d18O dataset

measured in BQS Bos1M3. M 5 mean; A: ampli-

tude; X: period; x0: delay





in extensive conditions with no control of breeding

in Northern France (Normandie: Lecomte and Le

Neveu 1986) and in Scotland (Reinhard et al. 1986)

show that calving occur over two to three months in

these populations, from May to July in Northern

France, from May to June in Scotland. In this regard,

a calving period spanning approximately six months

at Bercy seems relatively long, and might have been

related to the domestic status of the herd. A direct

consequence of a calving period extended over six

months would be availability of cattle milk all the

year round if a 6 to 7 months lactation is assumed for

Neolithic primitive domestic females (Peske 1994;

Tresset 1996).

Sheep birth seasonality

The same modelling procedure was applied to the

d18O values measured in sheep teeth. Results from the

calculation of the best fit for combined variations of

X, A, x0 and M are shown in Table 3 and in Fig. 4.

Values for x0/X vary from 0.19 to 0.43 (Table 3 and

Fig. 7a) and average 0?31. This figure is very similar

to the pattern observed on the second molar of a

dataset of ten modern sheep from a Shetland cross-

breed, born between the end of April and the

beginning of May, for which x0/X averages 0?28

(Balasse et al. 2012; Fig. 7b). This suggests that the

births of the analysed sheep from Bercy were centred

around that time. This is only slightly later than what

is observed nowadays in temperate climate and at

very close latitude: the birth period of free rang-

ing Shetland sheep in the nature reserve of Ma-

rais Vernier (Normandie, Northern France) occurs

between mid March and the end of April (Lecomte

and Le Neveu 1986).

Seasonal changes in cattle and sheep diet d13C

Stable carbon isotope values as low as 214?7% in

cattle and 215?3% in sheep were measured season-

ally in tooth enamel. Considering a 14?1% 13C-

enrichment (e*) between diet and bioapatite (Cerling

and Harris 1999), such values would correspond to

d13C values of 228?4% and 229?0% respectively for

diet. These end-members may be underestimated due

to a dampening effect caused by a delay in enamel

mineralisation (Balasse 2002; 2003; Passey and Cer-

ling 2002). This is considerably lower than the modal

d13C values of 225?6% expected for pre-industrial C3

plants in open areas (using a modal value of 227%for modern C3 plants: O’Leary 1988; Tieszen and

Boutton 1988; corrected by z1.4 % for the fossil fuel

effect: Freyer and Belacy 1983; Stuiver et al. 1984;

Friedli et al. 1986; Leavitt and Long 1986). In

temperate environments, low d13C values may be

attributed to the canopy effect, i.e. consumption of

vegetation grown in a dense forest. Kohn (2010)

recently recommended the d13C value of 231?5% as a

cut-off indicating the understories of closed-canopy

forests. However, d13C values measured on modern

roe deer bone collagen in the Dourdan deciduous

forest (France) average 224?7% (n 5 18, Rodiere

et al. 1996), from which, considering a 5% 13C-

enrichment between diet and collagen (Ambrose

1993; van der Merwe 1989), an average value of

229?7% can be estimated for diet. This is slightly

higher than the cut-off values recommended by

Table 3 Results from the calculation of the best fit (method of least squares) for combined variation of X (period), A(amplitude), x0 (delay) and M (mean) when the four-parameters model is applied to the sheep and cattle d18Odata sets. BQS 5 teeth from Bercy; KH: teeth from the Knap of Howar. R (Pearson): Pearson’s correlationcoefficient; r 5 cov (d18O, d18Om)/(s (d18O) * s (d18Om)); when | r | 5 1, the series are identical

Specimen X (mm) A (%) x0 (mm) M (%) x0/X r (Pearson)

BQS Bos1M3 41?8 1?5 18?1 25?3 0?43 0?98BQS Bos2M3 43?3 1?5 14?6 25?1 0?34 0?96BQS Bos3M3 39?4 0?9 4?6 25?8 0?12 0?94BQS Bos4M3 40?9 1?1 24?8 24?5 0?61 0?99BQS Bos5M3 31?0 0?7 17?8 25?4 0?57 0?95BQS Bos6M3 44?7 1?4 9?9 23?7 0?22 0?97BQS Bos7M3 57?3 2?1 26?9 24?1 0?47 0?99BQS Bos8M3 41?3 0?8 14?1 23?6 0?34 0?96BQS Bos9M3 43?0 1?0 26?8 24?0 0?62 0?94KH Bos1M3 37?6 1?2 17?0 25?1 0?45 0?98KH Bos2M3 39?6 1?3 21?5 24?9 0?54 0?99KH Bos3M3 46?8 1?1 26?4 25?1 0?57 0?98KH Bos4M3 42?1 1?3 22?0 25?5 0?52 0?99KH Bos5M3 34?2 0?7 19?2 25?2 0?56 0?98BQS Ovis1M2 33?4 1?7 6?2 25?8 0?19 0?98BQS Ovis2M2 22?8 2?0 7?6 23?8 0?33 0?99BQS Ovis3M2 29?2 2?1 12?5 23?5 0?43 0?98BQS Ovis4M2 19?2 1?1 7?2 24?3 0?38 0?96BQS Ovis5M2 36?7 2?8 7?9 25?1 0?21 0?99



Figure 6 Results from normalisation of the location in each cattle tooth crown where the highest d18O is measured (x0)

with the period (X) calculated for each tooth at Bercy (a) and the Knap of Howar (c). On graph b and d, on the

Y axis the period (1 5 a whole year) is divided into 12 units (months)

Figure 7 Results from normalisation of the location in each sheep tooth crown where the highest d18O is measured (x0)

with the period (X) calculated for each tooth at Bercy (a). Comparison with the Rousay sheep (b)





Kohn (2010), defined in tropical forests. Cor-

rected for the fossil fuel effect, this would lead to

an approximate value of 228?3 % in pre-industrial

ecosystems.

At Bercy, d13C values varying from 224?0% to

223?1% (n 5 6) were measured in aurochs bone

collagen (Balasse et al. 2000; Drucker et al. 2008)

corresponding to d13C values of 229?0 % to 228?1

% for diet. They were interpreted as reflecting feeding

in the understories of a dense forest. Fig. 8 compares

previous d13C values measured in bone collagen from

wild and domestic bovines (Balasse et al. 2000;

Drucker et al. 2008) with the range of intra-individual

variation of d13C values measured in cattle and sheep

tooth enamel (this study). To allow comparison, all

measured d13C values were translated to estimated

diet d13C values (see figure caption for an explana-

tion). For BQSBos1M3 to BQSBos9M3, the indivi-

dual mean d13C value measured in tooth enamel,

corresponding approximately to an annual mean

given that the sequence spans approximately a year,

falls within the range of variation of previous data

for domestic bovines from Bercy (reflecting a pluri-

annual mean). For BQSBos11M3 the mean d13C

value falls within the range of variation of previous

data for wild bovines, although only a short part of

the annual cycle is preserved in this tooth (Fig. 3).

When the range of intra-tooth variation is observed,

d13C values measured in BQSBos2, Bos5, Bos7 and

Bos8 fall seasonally within the range of variation

measured in aurochs, possibly reflecting feeding on

plants grown in similar ecological niches. From the

co-variation of d13C and d18O values in tooth enamel,

the lowest d13C values measured in the cattle M3

occur when d18O values are the lowest, in the winter.

The same pattern is observed in two sheep teeth

(BQSOvis2M2 and BQSOvis3M2). For both species,

seasonal consumption of plants from the neighbour-

ing oak forest (Leroyer 1998) can be envisaged.

Domestic stock could have been feeding directly in

the forest floor or lower part of the canopy as

mentioned for example in numerous accounts from

late Medieval and Renaissance Ireland (Lucas 1989),

or were provided with fodder collected in the forest

understories (see above).

Lynch et al. (2008) provided as an alternate

explanation for the lower d13C values measured in

aurochs from England compared with contempora-

neous domestic bovines, a preference for marshy

habitats for the aurochs. However, there is no evidence

for 13C-depletion in plants occurring naturally in

freshwater marshes and in the surroundings of rivers

and lakes (DeLaune 1986; Boutton 1991; Chmura and

Aharon 1995). Consequently, in the present state, the

canopy effect hypothesis is favoured.

In their stable isotope study of aurochs and early

cattle from southern Scandinavia, Noe-Nygaard et al.

(2005a and b) found no obvious evidence for leaf-

foddering in early domestic cattle (4000 to 3800

cal BC), with d13C values ranging from 222?5 to

221?4%, while d13C values of native aurochs from

the late Boreal–early Atlantic ranged from 224?2 to

Figure 8 Stable carbon isotope ratios (d13C) of the diet of cattle and sheep from Bercy, estimated from values measured

in tooth enamel (this study; mean and amplitude of variation) and in bone collagen: *BXXX were measured in

long bones (Drucker et al. 2008) and MB XXX were measured in mandibles (Balasse et al. 2000). The calculation

considered a 14?1% enrichment (e) between tooth enamel bioapatite and diet d13C (Cerling and Harris 1999) and

a 5% spacing between collagen and diet d13C. Shaded area: range of variation of d13C values estimated from

aurochs bone collagen d13C. B2201C, B6100C, B2001, B5500, B5600C and B6200C, were identified as aurochs

from morphological criteria (Drucker et al. 2008)




222?4%, reflecting a diet gathered in the dense

Atlantic forest. This comparison was based on bone

collagen values, reflecting pluri-annual means. At

Bercy, the annual mean estimated from the yearly

d13C sequences measured in the cattle M3 is

(213?4% in individuals seasonally feeding on low

d13C resources (Fig. 8). This would correspond to an

approximate value of 222?1% for collagen d13C

(using a 14?1% 13C-enrichment e between bioapatite

and diet, and a 5% spacing between collagen and diet

d13C). From this, it can be concluded that the

seasonal contribution of arboreal fodder to the diet

of early domestic cattle from southern Scandinavia

can not be definitely ruled out at least for three

individuals with collagen d13C values respectively of

222?26%, 222?16% and 222?44% (Noe-Nygaard

et al. 2005a, table 2). These estimations do not take

into account the precision around the determination

of the 214?1% enrichment between bioapatite and

diet (0?05%, Cerling and Harris 1999).

In temperate Europe, archaeological evidence of

tree foddering in the Middle Neolithic comes from

botanical remains found in sheepfolds or byres iden-

tified as such by ruminant dung layers and/or the

presence of shed deciduous teeth and high concentra-

tion of spheroliths. Research has focused on well-

preserved assemblages from lake shore settlements in

Switzerland (Akeret et al. 1999; Akeret and Jacomet

1997; Rasmussen 1989; 1993) and rockshelters from

southern France (Dehlon et al. 2088; Thiebault 2001;

2005). When all other pieces of evidence deal with the

foddering of caprines, at the Weier settlement in

Switzerland tree fodder was fed to cattle (Rasmussen

1989). Although most of the time the findings include

leaves or leaf bearing branches, the use of leafless

twigs was reported at Egolzwil 3 (Switzerland) asso-

ciated with small stock faeces (Rasmussen 1993).

Indeed, many descriptions from historical times attest

the harvesting of branches throughout the winter,

while deciduous trees are not bearing leaves, in

Scandinavia (Rasmussen 1993).

It may be important to note that the use of leafless

twigs as fodder might be more difficult to detect from

stable carbon isotopes. Although twigs are submitted

to the canopy effect as well as leaves, non-photo-

synthetic plant tissues in general, including twigs,

tend to be 13C-enriched by 1 to 3% compared with

leaves (Gebauer and Shulze 1991; Cernusak et al.

2009). Seasonal feeding on a diet gathered from the

forest understories would therefore be more difficult

to detect — compared with the feeding in open

pastures — if the consumed part is twigs rather than

leaves. For this reason the hypothesis of a leafy

resource should be favoured for the sheep and cattle

from Bercy.

Conclusion

Complementing pre-existing data from osteoarch-

aeology and bone collagen stable isotope analyses,

the present stable isotope study on tooth enamel

provides a more concrete picture of cattle and sheep

husbandry at Bercy, at the very beginning of the 4th

millennium BC. Sheep were lambing over a period

centred in the end April/beginning of May, only

slightly later than expected from what is observed

nowadays in temperate Europe at similar latitude.

Cattle were born over a period spanning approxi-

mately six months. This was unexpected compared

with a two to three months calving period in free-

ranging cattle populations. The extension of the calv-

ing period might have been related to the domestic

status of the herd, and possibly with seasonal food

supplementation. Some cattle and some sheep from

Bercy fed on a 13C-depleted resource in the winter,

potentially leafy fodder, although it is not possible

to say whether it was gathered in the forest by the

animals themselves, or harvested by the herders.

This practice did not happen every year and/or only

involved part of the herd. The effect on the timing

and duration of the reproduction cycle would be

less in sheep than in cattle, the breeding pattern of

the former being more directly linked to the pho-

toperiodic cycle. A direct consequence of an ex-

tended calving period would be on the annual

availability of cow milk, which would have covered

the whole year at Bercy. This is important informa-

tion in a context where the exploitation of milk by

the human community was demonstrated by the

presence of dairy product residues in ceramic vessels

(Regert 2007) and highly suspected from the demo-

graphic management of the cattle herd (Tresset

1996; 1997).

Acknowledgements

Yves Lanchon gave access to the faunal material for

stable isotope analysis. This research received finan-

cial support from the Region Ile de France, a CNRS

INSU program ECLIPSE II 2005–2007 (Climatic

constraints and development of Neolithic husbandry in

Western Europe at the Atlantic/Sub-Boreal boundary

(4th millennium BC), dir. M. Balasse and A. Tresset)

and a ERC Starting Grant GA 202881 SIANHE (dir.

M. Balasse). Stephanie Brehard provided us with her

expertise for the determination of sheep teeth. We

would like to thank Allowen Evin (UMR 7209



CNRS) and Gael Obein (Laboratoire Commun de

Metrologie, LNE-CNAM) for advice on the statis-

tical treatment of data.

ReferencesAkeret, O., Haas, J. N., Leuzinger, U. and Jacomet, S. 1999. Plant

macrofossils and pollen in goat/sheep faeces from the Neolithic

settlement Arbon Bleiche 3, Switzerland. The Holocene 9, 175–

82.

Akeret, O. and Jacomet, S. 1997. Analysis of plant macrofossils in goat/

sheep faeces from the Neolithic lake shore settlement of Horgen

Scheller — an indication of prehistoric transhumance? Vegetation

History and Archaeobotany 6, 235–39.

Ambrose, S. H. 1993. Isotopic analysis of paleodiets: methodological

and interpretive considerations, pp. 59–130 in Standford, M. K.

(ed.), Investigations of Ancient Human Tissue, Chemical Analyses in

Anthropology. Langhorne: Gordon and Breach Science Publishers.

Balasse, M. 2002. Reconstructing dietary and environmental history

from enamel isotopic analysis: time resolution of intra-tooth

sequential sampling. International Journal of Osteoarchaeology 12,

155–65.

Balasse, M. 2003. Potential biases in sampling design and interpreta-

tion of intra-tooth isotope analysis. International Journal of

Osteoarchaeology 13, 3–10.

Balasse, M. and Ambrose, S. H. 2005. Mobilite altitudinale des

pasteurs neolithiques dans la vallee du Rift (Kenya): premiers

indices de l’analyse du d13C de l’email dentaire du cheptel

domestique. Anthropozoologica 40(1), 147–66.

Balasse, M., Ambrose, S. H., Smith, A. B. and Price, T. D. 2002. The

seasonal mobility model for prehistoric herders in the south-

western Cape of South Africa assessed by isotopic analysis of

sheep tooth enamel. Journal of Archaeological Science 29, 917–

32.

Balasse, M., Bocherens, H., Tresset, A., Mariotti, A. and Vigne, J.-D.

1997. Emergence de la production laitiere au Neolithique?

Contribution de l’analyse isotopique d’ossements de bovins

archeologiques. C. R. Acad. Sci. Paris, Sciences de la Terre et

des Planetes 235, 1005–10.

Balasse, M., Obein, G., Ughetto-Monfrin, J. and Mainland, I. 2012.

Investigating seasonality and season of birth in past herds: a

reference set of sheep enamel stable oxygen isotope ratios.

Archaeometry 54(2), 349–368.

Balasse, M., Smith, A. B., Ambrose, S. H. and Leigh, S. R. 2003.

Determining sheep birth seasonality by analysis of tooth enamel

oxygen isotope ratios: the Late Stone Age site of Kasteelberg

(South Africa). Journal of Archaeological Science 3, 205–15.

Balasse, M. and Tresset, A. 2002. Early weaning of Neolithic domestic

cattle (Bercy, France) revealed by intra-tooth variation in nitrogen

isotope ratios. Journal of Archaeological Science 29, 853–59.

Balasse, M. and Tresset, A. 2007. Environmental constraints on

reproductive activity of domestic sheep and cattle: what latitude

for the herder? Anthropozoologica 42(2), 71–88.

Balasse, M., Tresset, A., Bocherens, H., Mariotti, A. and Vigne, J.-D.

2000. Un abattage ‘post-lactation’ sur des bovins domestiques

neolithiques. Etude isotopique des restes osseux du site de Bercy

(Paris, France). Anthropozoologica 31/Ibex J. Mountain Ecol. 5,

39–48.

Bernard, A., Daux, V., Lecuyer, C., Brugal, J.-P., Genty, D., Wainer,

K., Gardien, V., Fourel, F. and Jaubert, J., 2009. Pleistocene

seasonal temperature variations recorded in the d18O of Bison

priscus teeth. Earth and Planetary Science Letters 283, 133–43.

Boutton, T. W. 1991. Stable carbon isotope ratios of natural materials:

II. Atmospheric, Terrestrial, Marine and freshwater environments,

pp. 173–85 in Coleman, D. C. and Fry, B. (eds), Carbon Isotope

Techniques. San Diego: Academic Press, Inc.

Britton, K., Grimes, V., Dau, J. and Richards, M. P., 2009.

Reconstructing faunal migrations using intra-tooth sampling and

strontium and oxygen isotope analyses: a case study of modern

caribou (Rangifer tarandus granti). Journal of Archaeological

Science 36, 1163–72.

Brown, W. A. B., Christofferson, P. V., Massler, M and Weiss, M. B.

1960. Postnatal tooth development in cattle. American Journal of

Veterinary Research, 21(80), 7–34.

Bryant, J. D., Froelich, P. N., Showers, W. J., Genna, B. J. 1996a. A

tale of two quarries: biologic and taphonomic signatures in the

oxygen isotope composition of tooth enamel phosphate from

modern and Miocene equids. Palaios 11, 397–408.

Bryant, J. D., Froelich, P. N., Showers, W. J., Genna, B. J. 1996b.

Biologic and climatic signals in the oxygen isotopic composition of

Eocene-Oligocene equid enamel phosphate. Palaeogeography,

Palaeoclimatology, Palaeoecology 126, 75–90.

Cerling, T. E. and Harris, J. M. 1999. Carbon isotope fractionation

between diet and bioapatite in ungulate mammals and implica-

tions for ecological and paleoecological studies. Oecologia 120,

347–63.

Cernusak, L. A., Tcherkez, G., Keitel, C., Cornwell, W. K., Santiago,

L. S., Knohl, A., Barbour, M. M., Williams, D. G., Reich, P. B.,

Ellsworth, D. S., Dawson, T. E., Griffiths, H. G., Farquhar, G. D.

and Wright, I. J. 2009. Why are non-photosynthetic tissues

generally 13C enriched compared with leaves in C3 plants? Review

and synthesis of current hypotheses. Functional Plant Biology 36,

199–213.

Chmura, G. L. and Aharon, P. 1995. Stable carbon isotope signatures

of sedimentary carbon in coastal wetlands as indicators of salinity

regimes. Journal of Coastal Research 11, 124–35.

D’Angela, D. and Longinelli, A., 1990. Oxygen isotopes in living

mammal’s bone phosphate: further results. Chemical Geology 86,

75–82.

DeLaune, R. D. 1986. The use of d13C signature of C3 and C4 plants in

determining past depositional environments in rapidly accreting

marshes of the Mississippi river deltaic plain, Louisiana, USA.

Chemical Geology (Isotope Geosciences Section) 59, 315–20.

Delgado Huertas, A., Iacumin, P., Stenni, B., Sanchez, Chillon B. and

Longinelli, A., 1995. Oxygen isotope variations of phosphate in

mammalian bone and tooth enamel. Geochimica et Cosmochimica

Acta 59, 4299–4305.

Delhon, C., Martin, L., Argant, J. and Thiebault, S. 2008. Shepherds

and plants in the Alps: multi-proxy archaeobotanical analysis of

Neolithic dung from ‘La Grance Rivoire’ (Isere, France). Journal

of Archaeological Science 35, 2937–52.

Dietsch, M.-F. 1996. Gathered fruits and cultivated plants at Bercy

(Paris), a Neolithic village in a fluvial context. Vegetation History

and Archaeobotany 5, 89–97.

Dietsch-Sellami, M.-F. 2001. Les plantes sauvages au menu des

chasseens de Bercy: l’apport de la carpologie, pp. 9–16 in Actes

des Journees Archeologiques d’Ile-de-France, 9–10 decembre 2000.

Saint-Denis: Service regional de l’archeologie.

Drucker, D. G., Bridault, A., Hobson, K. A., Szuma, E. and

Bocherens, H. 2008. Can carbon-13 in large herbivores reflect

the canopy effect in temperate and boreal ecosystems? Evidence

from modern and ancient ungulates. Palaeogeography, Palaeo-

climatology, Palaeoecology 266, 69–82.

Edwards, C. J., MacHugh, D. E., Dobney, K. M., Martin, L., Russell,

N., Horwitz, L. K., McIntosh, S. K., MacDonald, K. C., Helmer,

D., Tresset, A., Vigne, J.-D. and Bradley, D. G. 2004. Ancient

DNA analysis of 101 cattle remains: limits and prospects. Journal

of Archaeological Science 31, 695–710.

Freyer, H. D. and Belacy, N. 1983. 13C/12C records in Northern

hemisphere trees during the past 500 years. Anthropogenic impact

and climatic superposition. Journal of Geophysical Research 88,

6844–52.

Fricke, H. C. and O’Neil, J. R., 1996. Inter- and intra-tooth variation in

the oxygen isotope composition of mammalian tooth enamel:

some implications for paleoclimatological and paleobiological

research. Palaeogeography, Palaeoclimatology, Palaeoecology 126,

91–100.

Friedli, H., Lotscher, H., Oeschger, H., Siegenthaler, U. and Stauffer,

B. 1986. Ice core record of the 13C/12C ratio of atmospheric CO2 in

the past two centuries. Nature 324, 237–38.

Gat, J. R., 1980. The isotopes of hydrogen and oxygen in precipitation,

pp. 21–42 in Fritz, P. and Fontes, J.-C. (eds), Handbook of

Environmental Isotope Geochemistry, vol. 1. The Terrestrial

Environment, vol. 1. The Terrestrial Environment. Amsterdam:

Elsevier.

Gebauer, G. and Schulze, E.-D. 1991. Carbon and nitrogen isotope

ratios in different compartments of a healthy and a declining Picea

abies forest in the Fichtelgebirge, NE Bavaria. Oecologia 87, 198–

207.



Hafez, E. S. E. 1952. Studies on the breeding season and reproduction

of the ewe. Journal of Agricultural Science 42, 189–231.

Halstead, P., Collins, P. and Isaakidou, V. 2002. Sorting the sheep from

the goats: morphological distinction between the mandibles and

mandibular teeth of adult Ovis and Capra. Journal of

Archaeological Science 29, 545–53.

Halstead, P. and Tierney, J. 1998. Leafy hay: an ethnoarchaeological

study in NW Greece. Environmental Archaeology 1, 71–80.

Helmer, D. 2000. Discrimination des genres Ovis et Capra a l’aide des

premolaires inferieures 3 et 4 et interpretation des ages d’abattage:

l’exemple de Tikili Tash (Grece). Anthropozoologica 31/Ibex J.

Mountain Ecol. 5, 29–38.

Henton, E., Meier-Augenstein, W. and Kemp, H. F., 2010. The use of

oxygen isotopes in sheep molars to investigate past herding

practices at the Neolithic settlement of Catalhoyuk, central

Anatolia. Archaeometry 52, 429–49.

Karsch, F. J., Bittman, E. L., Foster, D. L., Goodman, R. L., Legan,

S. J. and Robinson, J. E. 1984. Neuroendocrinal basis of seasonal

reproduction. Recent Progress in Hormone Research 40, 185–

232.

Kelly, F., 1997. Early Irish Farming. Vol. IV. Early Irish Law Series.

Dublin.

Koch, P. L., Fisher, D. C. and Dettman, D., 1989. Oxygen isotope

variation in the tusks of extinct proboscideans: a measure of

season of death and seasonality. Geology 17, 515–19.

Koch, P. L., Heisinger, J., Moss, C., Carlson, R. W., Fogel, M. L. and

Behrensmeyer, A. K., 1995. Isotopic tracking of change of diet and

habitat use in African elephants. Science 267, 1340–43.

Kohn, M. 2010. Carbon isotope compositions of terrestrial C3 plants as

indicators of (paleo)ecology and (paleo)climate. Proceedings of the

National Academy of Sciences 107(46), 19691–95.

Kohn, M., Schoeninger, M. J. and Valley, J. W., 1998. Variability in

oxygen compositions of herbivore teeth: reflections of seasonality

or developmental physiology? Chemical Geology 152, 97–112.

Lanchon, Y., Marquis, P. and Roblin-Jouve A., 2000. Le Premier

Village de Paris, il y a 6000 ans. Les Decouvertes Archeologiques de

Bercy. Paris: Paris-Musees

Land, L. S., Lundelius, E. L. and Valastro, S. 1980. Isotopic ecology of

deer bones. Palaeogeography, Palaeoclimatology, Palaeoecology

32, 143–51.

Leavitt, S. W. and Long, A. 1982. Evidence for 13C/12C fractionation

between tree leaves and wood. Nature 298, 742–43.

Lecomte, T. and Le Neveu, C. 1986. Le Marais Vernier. Contribution a

l’Etude et a la Gestion d’une Zone Humide. Unpublished PhD

thesis, Universite de Rouen Haute-Normandie.

Leroyer, C. 1998. Evolution de la vegetation et emprise de l’homme sur

le milieu a Bercy (Paris), pp. 407–15 in Gutherz, X. and

Joussaume, R. (eds), Le Neolithique du Centre-Ouest de la

France. Actes du XXIe Colloque Inter-regional sur le Neolithique,

Poitiers, 14–16 octobre 1994. Poitiers: Ministere de la Culture et de

la Communication, Association des Archeologues de Poitou-

Charentes.

Lincoln, G. A., Lincoln, C. E., McNeilly, A. S. 1990. Seasonal cycles

in the blood plasma concentration of FSH, inhibin and

testosterone, and testicular size in rams of wild, feral and do-

mesticated breeds of sheep. Journal of Reproduction and Fertility

88, 623–33.

Longinelli, A. 1984. Oxygen isotopes in mammal bone phosphate: a

new tool for paleohydrological and paleoclimatological research.

Geochimica et Cosmochimica Acta 48, 385–90.

Lucas, A. T. 1989. Cattle in Ancient Ireland. Irish Series. Kilkenny:

Boethius Press.

Luz, B., Kolodny, Y. and Horowitz, M. 1984. Fractionation of

oxygen isotopes between mammalian bone-phosphate and

environmental drinking water. Geochimica et Cosmochimica

Acta 48, 1689–93.

Lynch A. H., Hamilton, J. and Hedges, R. E. M. 2008. Where the wild

things are: aurochs and cattle in England. Antiquity 82, 1025–

39.

Marquis, P. 1991. Le site neolithique de Bercy (Paris XIIe). Bulletin de

la Societe prehistorique francaise 88, 239.

McCormick, F. 1998. Calf slaughter as a response to marginality, pp.

49–51 in Mills, C. M. and Coles, G. (eds), Life on the Edge: Human

Settlement and Marginality Oxford: Oxbow Monograph 10.

Nelson, S. V. 2005. Paleoseasonality inferred from equid teeth and

intra-tooth isotopic variability. Palaeogeography, Palaeoclima-

tology, Palaeoecology 222, 122–44.

Noe-Nygaard, N., Price, T. D. and Hede, S. U. 2005a. Diet of aurochs

and early cattle in southern Scandinavia: evidence from 15N

and 13C stable isotopes. Journal of Archaological Science 32, 855–

71.

Noe-Nygaard, N., Price, T. D. and Hede, S. U. 2005b. Corrigendum to

‘Diet of aurochs and early cattle in southern Scandinavia: evidence

from 15N and 13C stable isotopes’ [J. Archaeol. Sci. 32 (2005)

855e871]. Journal of Archaological Science 32, 1432.

O’Leary, M. H. 1988. Carbon isotopes in photosynthesis. Bioscience 38,

328–35.

Passey, B. H. and Cerling, T. E. 2002. Tooth enamel mineralization in

ungulates: implications for recovering a primary isotopic time-

series. Geochimica et Cosmochimica Acta 66, 3225–34.

Payne, S. 1985. Morphological distinction between the mandibular

teeth of young sheep, Ovis, and goats, Capra. Journal of

Archaological Science 12, 139–47.

Pernaud, J.-M. 1997. Paleoenvironnements Vegetaux et Societes a

l’Holocene dans le Nord du Bassin Parisien. Anthraco-analyses de

Sites Archeologiques d’Ile de France et de Picardie: Methodologie

et Paleoecologie. Unpublished Phd thesis, Universite de

Paris I.

Peske, L. 1994. Contribution to the beginning of milking in Prehistory.

Archeologishe rozhledy 46, 97–104.

Poplin, F. 1979. Origine du Mouflon de Corse dans une nouvelle

perspective paleontologique: par marronage. Annales de Genetique

et de Selection animale 11(2), 133–43.

Pyankov, V. I., Ziegler, H., Akhani, H., Deigele, C. and Luttge, U.

2010. European plants with C4 photosynthesis: geographical and

taxonomic distribution and relations to climate parameters.

Botanical Journal of the Linnean Society 163, 283–304.

Rasmussen, P. 1989. Leaf-foddering of livestock in the Neolithic:

archaeobotanical evidence from Weier, Switzerland. Journal of

Danish Archaeology 8, 51–71.

Rasmussen, P. 1990. Leaf foddering in the earliest Neolithic

agriculture. Evidence from Switzerland and Denmark. Acta

Archaeologica 60, 71–86.

Rasmussen, P. 1993. Analysis of goat/sheep faeces from Egolzwil 3,

Switzerland: evidence for branch and twig foddering of

livestock in the Neolithic. Journal of Archaeological Science

20, 479–502.

Regert, M., 2007. Elucidating pottery function using a multi-step

analytical methodology combining infrared spectroscopy, mass

spectrometry and chromatographic procedures. British

Archaeological Reports S1650, 61–76.

Reinhardt, C., Reinhardt, A. and Reinhardt, V. 1986. Social behaviour

and reproductive performance in semi-wild Scottish Highland

cattle. Applied Animal Behaviour Science 15, 125–36.

Ritchie, A. 1983. Excavation of a Neolithic farmstead at Knap of

Howar, Papa Westray, Orkney. Proceedings of the Society of

Antiquaries of Scotland 113, 40–121.

Rodiere, E., Bocherens, H., Angibault, J.-M. and Mariotti, A.1996.

Particularites de l’azote chez le chevreuil (Capreolus capreolus L.):

implications pour les reconstitutions paleoenvironnementales.

Comptes Rendus de l’Academie des Sciences, Paris 323(IIa), 179–

85.

Rosa, H. J. D. and Bryant, M. J. 2003. Seasonality of reproduction in

sheep. Small Ruminant Research 48, 155–71.

Sharma, S., Joachimski, M. M., Tobschall, H. J., Singh, I. B., Tewari,

D. P. and Tewari, R. 2004. Oxygen isotopes of bovid teeth as

archives of paleoclimatic variations in archaeological deposits of

the Ganga plain, India. Quaternary Research 62, 19–28.

Sharp, Z. D. and Cerling, T. E., 1998. Fossil records of seasonal climate and

ecology: straight from the horse’s mouth. Geology 26, 219–

22.

Stuart-Williams, H. L. Q. and Schwarcz, H. P., 1997. Oxygen isotopic

determination of climatic variation using phosphate from beaver

bone, tooth enamel and dentine. Geochimica et Cosmochimica

Acta 61, 2539–2550.

Stuiver, M., Burk, R. L. and Quay, P. D. 1984. 13C/12C ratios in tree

rings and the transfer of biospheric carbon to the atmosphere.

Journal of Geophysical Research 89, 11731–48.



Thiebault, S. 2001. Anthracoanalyse des etablissements neolithiques de

la region liguro-provencale. Bulletin de la Societe Prehistorique

Francaise 98, 399–409.

Thiebault, S. 2005. L’apport du fourrage d’arbre dans l’elevage depuis

le Neolithique. Anthropozoologica 40(1), 95–108.

Tieszen, L. L. and Boutton, T. W. 1988. Stable carbon isotopes in

terrestrial ecosystem research, pp. 167–95 in Rundel, P. W.,

Ehleringer, J. R. and Nagy, K. A. (eds), Stable Isotopes in

Ecological Research.. New York: Springer-Verlag.

Towers, J., Jay, M., Mainland, I., Nehlich, O. and Montgomery, J.

2011. A calf for all seasons? The potential of stable isotope

analysis to investigate prehistoric husbandry practices. Journal of


Tresset, A. 1996. Le Ro le des Relations Homme/Animal

dans l’Evolution Economique et Culturelle des Societes des

V-IVe Millenaires en Bassin Parisien. PhD thesis, Universite

Paris 1.

Tresset, A. 1997. L’approvisionnement carne cerny dans le contexte

neolithique du Bassin Parisien, in la culture de Cerny: nouvelle

economie, nouvelle societe au Neolithique. Actes du colloque de

Nemours 1994. Memoires du Musee de Prehistoire d’Ile de France

6, 299–314.

Tresset, A., Bollongino, R., Edwards, C. J., Hughes, S. and Vigne, J.-D.

2009. Early diffusion of domestic Bovids in Europe. An indicator

for human contacts, exchanges and migrations?, pp. 69–90 in

d’Errico F. and Hombert J.-M. (eds), Becoming Eloquent.

Amsterdam/Philadelphia: John Benjamins.

Troels-Smith, J. 1960. Ivy, mistletoe and elm climate indicators: a

contribution to the interpretation of the pollen zone border VII–

VIII. Danmarks Geologiske Undersogelse IV Raekke, 4(4), 4–32.

van Dam, J. A. and Reichart, G. J., 2009. Oxygen and carbon isotope

signatures in late Neogene horse teeth from Spain and application

as temperature and seasonality proxies. Palaeogeography,

Palaeoclimatology, Palaeoecology 274, 64–81.

van der Merwe, N. J., 1989. Natural variation in 13C concentration and

its effect on environmental reconstruction using 13C/12C ratios in

animal bones, pp. 105–25 in Price, T. D. (ed.), The Chemistry of

Prehistoric Human Bone.. Cambridge: Cambridge University Press.

van der Merwe, N. J. and Medina, E. 1991. The canopy effect, carbon

isotope ratios and foodwebs in Amazonia. Journal of


Vigne, J.-D., Geigl, E.-M., Pruvost, M., Bollongino, R. and Tresset, A.,

2007. Paleogenetique et domestication des Bovides. Ethnozootechnie

79, 7–13.



Stable isotope insights ( δ 18 O, δ 13 C) into cattle and sheep husbandry at Bercy (Paris, France,...

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