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Journal of Anthropological Archaeology
journal homepage: www.elsevier.com/locate/jaa
Integration of Linearbandkeramik cattle husbandry in the forested landscape
of the mid-Holocene climate optimum: Seasonal-scale investigations in
Bohemia
Rémi Berthona,⁎
, Lenka Kovačikováb
, Anne Tresseta
, Marie Balassea
a
UMR 7209 « Archéozoologie, archéobotanique: sociétés, pratiques, environnements » Sorbonne Universités, CNRS-MNHN, 55 rue Buffon, 75005 Paris, France
b
Laboratory of Archaeobotany and Palaeoecology, Faculty of Science, University of South Bohemia, Na Zlaté stoce 3, 370 05 České Budějovice, Czech Republic
A R T I C L E I N F O
Keywords:
Linearbandkeramik
Cattle
Birth seasonality
Forest
Enamel
Carbon isotopes
Oxygen isotopes
Chotěbudice
Černý Vůl
Czech Republic
A B S T R A C T
Domestic animals and plants were introduced to Europe from the Near East and subsequently spread across
Europe, entailing adaptations to different environments with consequences for the biology of organisms, agropastoral
technical systems and socio-economic organisation. Agriculture was introduced to Central Europe by
Linearbandkeramik (LBK) societies between 5600 and 4900 cal. BC, in predominantly forested environments.
LBK farming systems involved intensive permanent field cultivation in natural openings. Milking was practiced
as evidenced from cattle mortality profiles and lipid residues in ceramics. Questions arise as to what extent LBK
cattle husbandry relied on woodland, and as to whether the seasonal scarcity of fodder conditioned cattle reproduction
cycles, with consequences on milk availability. Results from the δ13
C and δ18
O analysis of cattle tooth
enamel at Chotěbudice and Černý Vůl (Bohemia, Czech Republic) suggest a limited use of dense forest for cattle
herding, even on a seasonal scale: cattle were kept in the open component of the forest/steppe mosaic landscape.
Winter forest browsing/provision of leafy fodder was evidenced in one specimen. At Chotěbudice, cattle births
mainly occurred over a two to three-month period, suggesting environmental constraints on cattle fertility cycles,
and possibly seasonal fodder scarcity. A direct consequence of this would be a shorter period of milk
availability throughout the year.
1. Introduction
The primary centres of domestication of European cattle, caprines
and pigs are located in the Near East and date to the 9th millennium cal
BC. These animals were then introduced to Europe at the turn of the 7th
millennium with domestic plants and husbandry practices. They spread
westward along the Mediterranean margins, on one hand, and following
the continental Danubian route, on the other hand, to reach
western Iberian coasts during the second half of the 6th millennium and
European northwestern coasts at the turn of the 5th millennium. The
British Isles were eventually colonized several centuries later (Tresset
and Vigne, 2011). During this expansion across Europe, animals and
husbandry practices had to adapt to diverse environmental conditions,
which differed from those in the Near East (Vigne, 2008). New selection
pressures linked to the climate, landscape, dietary resources and pathogens
led to the development of new physiological abilities/technical
practices regarding reproduction, feeding behaviour, metabolism, animal
products and resistance to diseases (Balasse and Tresset, 2007,
2009, 2017; Flori and Gautier, 2013). In this sense, the spread of
domesticates across Europe represents some of the earliest deliberate
acclimatisation of domestic animals by human societies. More specifically,
environmental factors determine livestock feeding resources and
reproductive behaviours, which in turn greatly impact the rhythms of
pastoral systems on a seasonal scale. Investigating the extent to which
environmental constraints affected agropastoral systems, and the solutions
adopted to cope with these constraints, is paramount for a better
understanding of how Neolithic economies successfully spread across
the diverse climatic zones of Europe.
In Central Europe, agriculture was first introduced by the
Linearbandkeramik (LBK) societies. The LBK developed between 5600
and 4900 cal. BC (Price, 2000; Banffy, 2004; Dolukhanov et al., 2005;
Pavlů, 2005; Jakucs et al., 2016) over a period of winter warming,
summer cooling and increased precipitation (Sanchez Goni et al.,
2016). The mid-Holocene climatic optimum led to the retreat of the
Pleistocene steppe in Central Europe in favour of the spread of deciduous
forests. The degree of openness of these forests has been debated,
and a steppe component has also been brought to light in the landscape,
the extent of which may have varied considerably in accordance with
https://doi.org/10.1016/j.jaa.2018.05.002
Received 23 November 2017; Received in revised form 2 May 2018
⁎
Corresponding author.
E-mail addresses: remi.berthon@mnhn.fr (R. Berthon), lenka.kovacikova@gmail.com (L. Kovačiková), anne.tresset@mnhn.fr (A. Tresset), marie.balasse@mnhn.fr (M. Balasse).
Journal of Anthropological Archaeology 51 (2018) 16–27
Available online 22 May 2018
0278-4165/ © 2018 Elsevier Inc. All rights reserved.
T
regional macroclimatic differences (Kreuz, 2008; Bogucki et al., 2012;
Marinova et al., 2012/2013; Salavert et al., 2014; Pokorný et al., 2015).
LBK communities developed farming systems with the following main
features. Natural openings were exploited for the intensive permanent
field cultivation (Bogaard, 2005; Saqalli et al., 2014) of crops dominated
by hulled wheat (einkorn and emmer; Colledge et al., 2005;
Jacomet, 2007; Dreslerová and Kočár, 2013). Soil fertility may have
been maintained using manure from livestock (Bogaard, 2005; Bogaard
et al., 2013). Cattle (Bos taurus) were raised as an important component
of the animal economy (Bogucki, 1982; Kovačiková et al., 2012;
Arbogast and Jeunesse, 2013), even though regional variability has
been observed in the relative proportion of cattle among domestic
stock, as well as in the role of hunting (Tresset and Vigne, 2001). The
analysis of 19 cattle mortality profiles from LBK contexts concluded
that milk exploitation was a widespread, although non-intensive,
practice across Europe (Gillis et al., 2017). Milk exploitation and
transformation were presumed to have been practised in the LBK of
Central Europe due to the occurrence of perforated vessels similar to
sieves in typology (Bogucki, 1984). Their actual use for cheese making
was confirmed by the identification of dairy fat residues in ceramic
sieves from the classic to late phases of the LBK (5200–4900 cal. BC) in
the region of Kuyavia, Poland (Salque et al., 2012) and at Brodau (LBK
classic phase) in Germany (Salque et al., 2013). Dairy fat residues were
also retrieved in some instances from non-perforated bowls in the
Kuyavia LBK (Salque et al., 2012). Meanwhile, no milk residues were
detected in ceramic assemblages from other LBK sites in Saxony and
Bavaria in Germany (Salque et al., 2013), or in Bohemia (Mátlová et al.,
2017), although the absence of evidence of the use of ceramic vessels
for milk collection or processing does not preclude milk exploitation.
These combined elements raise questions concerning the extent to
which LBK cattle husbandry developed in the woodland component of
the landscape. How was the availability of open grasslands managed
throughout the year to feed these large domestic, predominantly
grazing herbivores, especially if open lands were cultivated?
Palynological evidence indicates pasturing in the floodplains of the
river valleys (Kreuz, 2008). The use of forest resources, including
woodland grazing and the production of fodder from coppicing and
pollarding is often assumed (Kreuz, 2008; Saqalli et al., 2014), building
on direct evidence in more recent contexts (lakeside and wetland settlements
from the last third of the 5th millennium cal BC onwards;
summary in Kühn et al., 2013). It has also been argued that the reconfiguration
of caprine husbandry by early Neolithic farmers in
southeastern Europe, into a cattle pastoral system in the LBK was a
consequence of adaptation to a forested landscape, based on the assumption
that cattle cope relatively well with forest browsing (RowleyConwy
and Legge, 2016). However, beyond the environmental perspective,
social factors including forms of property have been put forward
as the main driving factor for the high representation of cattle in
LBK sites (Shennan, 2011; Manning et al., 2013). The use of the forest
to feed cattle has actually only been directly tested to a small extent in
LBK contexts (see below). The question of subsistence strategies is also
paramount, firstly, for a better understanding of how crop cultivation,
animal husbandry and forest exploitation were functionally interconnected
in LBK economic systems, and secondly, because the management
of the cattle diet would have directly influenced animal production.
Seasonally-reduced forage availability, especially in
wintertime, when meadows are covered with snow and forests do not
produce much fodder, could have conditioned cattle reproduction cycles,
for which food resources are the main restricting factor (Santos
et al., 2006; Burthe et al., 2011). In turn, cattle birth distribution would
have partly determined the organization of pastoral tasks throughout
the year, but also the availability of milk on a seasonal scale: both
elements are key parameters for approaching the socioeconomic modalities
of these agropastoral systems.
These aspects of LBK cattle husbandry can be investigated using
stable isotope analyses of skeletal remains. Previous stable isotope
studies of LBK faunal assemblages consisted in the analysis of bone
collagen stable carbon (δ13
C) and nitrogen (δ15
N) isotope compositions
in order to define the environmental settings. In particular, δ13
C values
can be used to evaluate the degree of habitat closure: in closed forests,
the reduction of light intensity influences photosynthesis efficiency,
leading to lower δ13
C values in plants (the canopy effect; van der
Merwe and Medina, 1991). These values are then passed along the food
chain (Drucker et al., 2008). They may therefore be used to test for
cattle feeding on forest resources. In this article, we first present a
summary of previously published δ13
C values in cattle bone collagen in
LBK contexts; we show that even though the evidence points to cattle
feeding on forest resources in a few sites in the western margin of the
LBK extension area, the question remains open in other regions of
Central Europe. We then provide two additional δ13
C datasets for cattle
remains from the LBK sites of Chotěbudice and Černý Vůl in northwestern
Czech Republic. Feeding on forest resources was additionally
investigated on a seasonal scale, through the sequential analysis of
stable carbon and oxygen isotope ratios in tooth enamel. Our hypothesis
is that stable isotope values measured in bone collagen are less
suitable for detecting the seasonal contribution of forest resources to
cattle diet than the enamel record. Meanwhile, stable oxygen isotope
ratios provide a tool for investigating cattle birth seasonality. Although
this parameter has been investigated in other European prehistoric
contexts, no data are currently available for the LBK. Studies conducted
in the Romanian early Neolithic and Chalcolithic (early 6th and 5th
millennium BC; Balasse et al., 2013, 2014), and in the Middle Neolithic
in France (early 4th millennium BC; Balasse et al., 2012a), point to
restricted birth seasons, compared to year-round breeding in presentday
European husbandry where the nutritional status of cattle is
maintained to a high level throughout the annual cycle. Although a
restricted calving period is also expected for the LBK, this parameter
must be directly determined on faunal remains before it can be confidently
introduced into models aiming to describing the sustainability
of LBK farming systems (for example, Saqalli et al., 2014).
2. Previously published δ13
C datasets from LBK bovine bone
collagen
Available datasets of bone collagen stable isotope composition
measured on LBK assemblages are summarized in Fig. 1A. Values specifically
referred to as “aurochs” in the given publications are not included.
However, it must be noted that the “cattle” datasets from the
studies in Bickle and Whittle (2013) often potentially include aurochs
as a minor component. This is important given that the Holocene aurochs
is commonly referred to as a forest animal. The mean δ13
C values
within each site range from −22.9 to −19.8‰ but variability within
each dataset may be important. In modern ecosystems, the great majority
of δ13
C values in C3 plants (which dominated Neolithic Europe)
range from −29 to −25‰ (−26.5‰ on average) when growing in
open areas (Kohn, 2010). These would lead to δ13
C values of −27.5 to
−23.5‰ (−25‰ on average) in plants from pre-industrial ecosystems
(once corrected by +1.5‰ for the fossil fuel effect: Freyer and Belacy,
1983), and δ13
C values ranging from −22.5 to 18.5‰ (mean −20‰
on average) in collagen (when applying a 5‰ spacing between diet and
collagen δ13
C; Ambrose and Norr, 1993). In contrast, late Boreal/early
Atlantic aurochs feeding in the dense Atlantic forest in Scandinavia
yielded δ13
C values ranging from −24.2 to −22.4‰ (Noe-Nygaard
et al., 2005a, 2005b). Considering this, we use the −22.5‰ value in
collagen as a threshold for a major forest component in the diet.
Among the collected datasets for Central Europe, the δ13
C values
appear to follow a geographical pattern, along a longitudinal gradient
(Fig. 1B). The highest δ13
C values were measured in the eastern margin
of Hungary (settlements of Füzesabony-Gubakút, Polgár-Ferenci-hát
and Balatonszárszó; Whittle et al., 2013a), indicating cattle grazing in
open areas. The lowest δ13
C values were measured in the western
margin of southwestern Germany (Vaihingen; Fraser et al., 2013) and in
R. Berthon et al. Journal of Anthropological Archaeology 51 (2018) 16–27
17
eastern France (Bischoffsheim; Bickle et al., 2013a), clearly demonstrating
cattle feeding on forest resources. This pattern of variability
across Europe is mostly explained by differences in landscape structuration
on a macro-regional scale. Indeed, the pollen profiles obtained
in southern and western Germany suggest the existence of very thick
forests covering most of the territory in these areas (Kalis and
Zimmermann, 1988; Kalis et al., 2003). In contrast, a forest-steppe
mosaic landscape was reconstructed further east, for example in the
Czech Republic, where palynological data, supported by molecular data
on the disjunctive floristic and faunistic elements of continental steppes
(Kajtoch et al., 2016, Kuneš and Abraham, 2017), the mollusc fauna
dwelling exclusively in open habitats (Ložek, 2007, Juřičková et al.,
2013, Kuneš and Abraham, 2017), and the presence of chernozem soils
(Antoine et al., 2013, Kuneš and Abraham, 2017) all attest to a nonforest
component.
Cattle bone collagen from Austria (Gnadendorf and Rutzing; Bickle
et al., 2013b), the Czech Republic (Vedrovice, Těšetice-Kyjovice and
Brno-Starý Lískovec; Whittle et al., 2013b), Slovakia (Blatné; Whittle
et al., 2013b), Poland (Kopydłowo; Marciniak et al., 2017), central
Germany (Karsdorf; Oelze et al., 2011) and southeastern Germany
(Aiterhofen-Ödmühle, Straubing-Lerchenhaid and Heilbronn-Neckargartach;
Dürrwächter et al., 2006; Bentley et al., 2013; Hofmann
et al., 2013) showed intermediate mean δ13
C values. These datasets also
include in some instances a few cattle remains yielding lower δ13
C
values (≤−23‰), clearly reflecting forest feeding. The carbon in bone
collagen is incorporated from the diet over timescales ranging from
months to years, depending on the species and the individual metabolic
state (Libby et al., 1964; Hobson and Clark, 1992). Consequently, bone
collagen stable isotope compositions integrate average dietary contributions
on a (pluri)annual scale, and intermediate δ13
C values could
reflect seasonal woodland grazing or leaf foddering. In contrast, tooth
enamel retains a chronological record of the stable isotope composition
of diet over the duration of tooth formation, which is not consecutively
remodelled once enamel mineralization is completed. A seasonal
dietary contribution can therefore be demonstrated through the sequential
analysis of stable carbon and oxygen isotope ratios in enamel
(Balasse et al., 2012a). Here, we tested the hypothesis of the seasonal
contribution of forest resources to cattle diet in two additional sites
from the Czech Republic.
3. Materials and methods
3.1. Description of the archaeological settlements
In the territory of the present-day Czech Republic, the exploitation
of animal resources by LBK communities focused predominantly on
husbandry. Bovinae (mostly domestic cattle) represent over 40% and
up to 85% of the identified specimens (NISP) in these LBK contexts,
while small stock (sheep, goats and pigs) play a secondary role in the
subsistence economy, with varying respective proportions, possibly due
to specific local environmental conditions (Kovačiková et al., 2012).
The LBK settlement of Chotěbudice is located in the northwestern part
3
4
5
6
7
8
9
10
-24 -23 -22 -21 -20 -19 -18
δ15N(‰)
δ13C (‰)
Bovinebone collagen
, , ,
Czech ,
,
A
B
3
4
5
6
7
8
9
10
-24 -23 -22 -21 -20 -19 -18
δ15N(‰)
δ13C (‰)
Bovinebone collagen - -
-
-
-
-
-
Fig.
1. A – Mean and standard deviation of
δ13
C and δ15
N values in LBK bovine bone
collagen from Central Europe (AT: Austria;
CZ: Czech Republic; DE: Germany; FR:
France; HU: Hungary; PL: Poland; SK:
Slovakia) (ref 1: Bickle and Whittle, 2013;
ref 2: Marciniak et al., 2017; ref 3: Oelze
et al., 2011; ref 4: Fraser et al., 2013). Legend
from East to West. B – Mean and
standard deviation of δ13
C and δ15
N values
in bovine bone collagen from Chotěbudice
and Černý Vůl, compared to macroregional
groups (the convex hulls use the mean va-
lues).
R. Berthon et al. Journal of Anthropological Archaeology 51 (2018) 16–27
18
of the Czech Republic (Fig. 2). The site lies at an altitude of 290 m
above sea level on a gentle southeast-facing slope close to the Dubá
stream. Archaeological excavations revealed nineteen ground plans of
long houses and 800 LBK structures spanning a long-term occupation
ranging from LBK stages IIa to LBK IIIb, i.e., 5400 to 5100 BCE, as well
as a number of features from the Stichbandkeramik (SBK) occupation
(Rada, 1981; Šumberová, 1991). Domestic cattle occur as the main
species in the archaeozoological assemblage throughout the LBK occupation
(Total LBK NISP = 4788. Cattle represent from 63 to 85% of
the NISP – depending on the occupation phase, cattle
NISP = 314–2017), followed by sheep, goat and pig. Cattle kill-off
patterns for the three most representative LBK phases show similar
trends over a period of roughly 200 years and suggest a dual exploitation
of milk and meat (Kovačiková et al., 2012).
Černý Vůl is located in the Prague-West district (Fig. 2). The site lies
at an altitude of 270 m above sea level in the basin of the Únětický
stream. Rescue excavations from 1975 to 1977 revealed the ground
plans of nine LBK post-hole houses dated to the LBK IIc-IIIb (5200–5000
BCE; Řídký et al., 2008). After a hiatus, the settlement was reoccupied
during the latter stage of the SBK IV (4800–4500 BCE; Pavlů and
Zápotocká, 2007). Cattle dominate the LBK faunal assemblage, although
in lower proportions (43% of NISP; Total LBK NISP = 389) than
in Chotěbudice, suggesting variability within the LBK on a regional
scale. At this site, the cattle kill-off pattern resembles the pattern established
at Chotěbudice, suggesting exploitation for milk and meat
(Kovačiková et al., 2012).
3.2. Selected bovine remains and attribution to wild or domestic Bos
The sample under study included ten cattle lower third molars from
the LBK occupation levels at Chotěbudice, five of which (CHO Bos1 to
Bos5) had been previously analysed (Kovačiková et al., 2012), and four
lower third molars from the LBK occupation levels at Černý Vůl. In
addition, thirty bovine (cattle/aurochs) long bones were sampled; 18 of
which come from the LBK occupation at Chotěbudice and five from the
LBK occupation at Černý Vůl. Seven bovine bones were also selected
from the SBK occupation at Černý Vůl. These were used to check for any
change in the degree of openness of the landscape over the centuries.
We paid particular attention to the assessment of the domestic status
of the selected remains to avoid potential misidentification with its wild
counterpart, the aurochs, which is very similar in morphology to the
domestic cattle. Aurochs were hunted at Chotěbudice and Černý Vůl
although their presence is rare in both assemblages (respectively 1.1
and 0.3% of the LBK faunal assemblages; Kovačiková et al., 2012). The
assessment of the domestic/wild status of the selected specimens was
based on osteometric criteria for molars and long bones. For the molars,
the length and breadth of the neck (following Ducos, 1968) were
compared to reference measurements of aurochs’ molars (Bos primigenius)
from Denmark (Degerbøl and Fredskild, 1970) and southeastern
France (Helmer and Monchot, 2006). Aurochs from Central Europe are,
on average, larger than those from Denmark and southwestern Europe
(Bökönyi, 1962; Jarman, 1969; Lasota-Moskalewska and Kobryń, 1990;
Guintard, 1999; Wright and Viner-Daniels, 2015). We are aware that
size criteria may sometimes be misleading for the distinction between
wild and domestic bovines (Vigne et al., 2007; Tresset et al., 2009).
However, the selected teeth from Chotěbudice and Černý Vůl are
smaller than the rather small aurochs from southeastern France and
therefore probably belong to domestic individuals (Supplementary
material 1). Only CHO Bos1M3 (from the previous dataset; Kovačiková
et al., 2012) measurements overlap with the aurochs dataset. Its status
is therefore uncertain.
The attribution of individual long bones to domestic or wild forms is
also challenging, partly because the largest domestic individuals mostly
overlap in size with small wild females (Degerbøl and Fredskild, 1970;
Edwards et al., 2007; Scheu et al., 2008). For this reason, as a precaution,
only specimens with measurements falling within the range of
variation of the male aurochs from Denmark (Degerbøl and Fredskild,
1970) were attributed to Bos primigenius. Specimens smaller than the
wild females from Denmark were attributed to Bos taurus. All remaining
specimens were attributed to Bos sp. (Supplementary material 2).
3.3. Bone collagen extraction and stable isotope measurements
Collagen was extracted from 320 to 350 mg of bone powder, following
the steps described in Bocherens et al. (1991). The analysis was
conducted on 400–600 µg of collagen on a Thermo Flash 2000 EA
Fig. 2. Map of the Czech Republic with the location of Chotěbudice, Černý Vůl, and Vedrovice, Těšetice-Kyjovice and Brno-Starý Lískovec, the stable isotope data of
which are included in the discussion for comparison.
R. Berthon et al. Journal of Anthropological Archaeology 51 (2018) 16–27
19
interfaced with a Thermo DeltaVAdvantage IRMS. The analytical precision,
determined from 7 to 11 analyses of an alanine standard within
each run, was ≤0.06‰ for δ13
C and ≤0.19‰ for δ15
N, and ≤0.5% for
C content, and ≤0.3% for N content.
3.4. Enamel sequential sampling and stable isotope measurements
Sampling was preferentially performed on the lower M3 hypoconid
lobe (for dental nomenclature see Bärmann and Rössner, 2011) in order
to be consistent with previous analyses at Chotěbudice (Kovačiková
et al., 2012), or, failing this, on the protoconid lobe (CerV Bos1M3 and
CerV Bos2M3). Both the protoconid and hypoconid lobes were sampled
in CHO Bos7M3 and CHO Bos8M3 (Supplementary material 3), in order
to check the consistency of the isotopic sequences between the two
lobes of a tooth. Sequential sampling was performed by drilling on the
buccal side of the teeth. The number of samples taken from one tooth
varies from 15 to 30 for a total number of 359 enamel samples. Enamel
samples weighing 5–12 mg were treated in 0.1 M acetic acid (0.1 ml
solution/1 mg sample) for 4 h (Tornero et al., 2013). Pre-treated enamel
samples weighing ∼ 600 μg were analysed on a Kiel IV device interfaced
with a Delta V Advantage IRMS. The accuracy of the data was
checked through the analysis of our internal laboratory carbonate
standard (Marbre LM normalized to the international standard NBS 19).
Over the period of analysis of the samples, the analytical precision
within each run, calculated from 6 to 8 measurements of Marbre LM,
varies from 0.02‰ to 0.06‰ for δ18
O and from 0.01‰ to 0.03‰ for
δ13
C. Results are expressed in V-PDB.
3.5. Interpretation of δ18
O sequences and modelling
In temperate Europe, δ18
O values in skeleton bioapatite from large
mammal skeletons are strongly related to δ18
O values in local precipitation
(Land et al., 1980; D’Angela and Longinelli, 1990). The δ18
O
values in precipitation vary seasonally with ambient temperature, with
higher values in the summer and lower values in the winter (Rozanski
et al., 1993). This annual cycle pattern is recorded in tooth enamel
during mineralization and may be retrieved through sequential δ18
O
analysis (Bryant et al., 1996, Fricke and O’Neil, 1996). Because the
timing of tooth growth is fixed for a species, inter-individual variability
in the positioning of the δ18
O cycle within a given tooth is an expression
of inter-individual variability in the birth season (Bryant et al., 1996,
Fricke and O’Neil, 1996; Balasse et al., 2003). This variability may be
described using the position (distance from the enamel-root junction
ERJ), where the highest δ18
O value is measured in the tooth crown
(Balasse et al., 2003). This position (thereafter referred to as x0) may be
objectively described through the modelling of the δ18
O sequence following
a cosine-function (Balasse et al., 2012b and Supplementary
material 4). Modelling also eliminates the effect of variability in tooth
size through the normalization of the distance (x0) using the period of
the cycle (X, or the length formed over a year). Inter-individual variability
in the period of birth within the annual cycle is then expressed
using the ratio x0/X, or the distance of the highest tooth crown values,
normalized to the period of the cycle which is an expression of the tooth
size (Balasse et al., 2012b).
4. Results
4.1. Collagen preservation and stable carbon and nitrogen isotope values
The results are reported in Fig. 3 and Supplementary material 5. All
the bone samples yielded well-preserved, good-quality collagen. Most
collagen extraction yields vary from 20 to 134 mg/g (mean of
77 ± 32 mg/g). The carbon contents of these extracts vary from 36 to
41% and the nitrogen contents vary from 14 to 16%. Collagen C:N
ratios range between 3.0 and 3.2. Two collagen extracts (CHO Bos17
and CHO Bos22) do not fulfil the preservation criteria of a collagen
yield of ≥20 mg/g (following recommendations by van Klinken, 1999).
However, their %C and %N are within the accepted range (%C > 30
and %N > 11) and their C:N ratios were also consistent with expectations
(3.2). Their δ13
C and δ15
N values are included in the final
dataset.
The δ13
C and δ15
N values are consistent with the expected range for
Neolithic temperate Europe. The δ13
C values range between −22.9 and
−19.5‰ (mean −20.6 ± 0.7‰ at CHO and −20.6 ± 0.4‰ at CerV)
and the δ15
N values vary between 5.5 and 8.4‰ (mean 7.0 ± 1.2‰ at
CHO and 7.3 ± 0.8‰ at CerV) (Fig. 3). The metacarpal CHO Bos25,
identified as an aurochs, stands out with notably low δ13
C (−22.9‰)
and δ15
N (3.7‰) values, which will be discussed below. The values
measured in the aurochs bones overlap with those measured in Bos
taurus and Bos sp. At Černý Vůl, the bovine specimens from the LBK and
SBK occupations yielded similar δ13
C values (respectively
−20.6 ± 0.4‰ and −20.7 ± 0.4‰ on average), suggesting no real
change in the degree of openness of the bovines’ habitat between the
two time periods.
4.2. Tooth enamel stable carbon and oxygen isotope values
Results from δ18
O and δ13
C measurements in enamel bioapatite are
reported in Figs. 4 and 5 and Supplementary material 6. In the sequences
of δ18
O values, the sinusoidal variations in time along the tooth
crown, observed on most individuals, reflect the seasonal ambient
temperature cycle. The δ18
O values vary between −9.7 and −2.2‰,
with an intra-tooth variation amplitude of 2.3 to 5.2‰. Results from
the modelling of δ18
O sequences are reported in Supplementary material
4. Among the parameters used to describe the δ18
O cycles, the X
parameter, i.e., the length of the tooth formed over a year or the cycle
period, varies from 29 to 44 mm with a mean value of 36 mm. CHO
Bos1M3 stands apart with a significantly longer cycle period (55 mm),
in keeping with its larger size (Supplementary material 1). CerV
Bos1M3, CerV Bos2M3 and CerV Bos4M3 could not be modelled due to
the absence of two consecutive maximum and minimal values which
allow for a secure definition of the length of the cycle. The δ13
C values
in enamel vary between −13.6 and −9.8‰. The amplitude of intratooth
variation is moderate (0.4–1.4‰). CHO Bos9M3 is an exception
to this pattern with a higher variation in amplitude (2.4‰), due to
lower δ13
C values in wintertime (Fig. 4).
4.3. Difference between the protoconid and hypoconid lobes
The δ18
O and δ13
C sequences measured in the protoconid and hypoconid
lobes of CHO Bos7M3 and CHO Bos8M3 are compared in
Fig. 6, highlighting a significant shift in both teeth. For each molar, the
modelled δ18
O sequences are similar in both lobes (X = 31.1 vs
31.8 mm and 43.4 vs 43.9 mm respectively in CHO Bos7M3 and CHOBos8M3),
reflecting similar enamel deposition and mineralization rates
(Supplementary material 4). However, the position of the optimum
δ18
O value (x0) shifts by +8.5 mm and +7.9 mm (in CHO Bos7M3 and
CHO Bos8M3 respectively) in the hypoconid lobe, compared to the
protoconid lobe (Supplementary material 4). The locations of the
highest and lowest δ18
O values, as determined from the model, are
reported on the tooth crown in Supplementary material 3, where it can
be seen that the shift is partly explained by an advanced development of
the protoconid compared to the hypoconid lobe. This difference is
slightly exacerbated in the highest part of the crown, where the difference
in the position of maximum δ18
O values is higher than that
observed between the minimum values recorded at a later stage of
crown growth (Supplementary material 3). More importantly, most of
the shift in the distance can be explained by the lower location of the
enamel-root junction (ERJ) in the hypoconid lobe compared to the
protoconid lobe, both leading to higher distances for samples in the
hypoconid lobe.
As a consequence of this shift, the x0/X value calculated in both
R. Berthon et al. Journal of Anthropological Archaeology 51 (2018) 16–27
20
lobes differs by 0.245 (or 24.5% of an annual cycle) in CHO Bos7M3
and 0.17 in CHO Bos8M3 (Supplementary material 4), or approximately
three and two months respectively. This precludes a direct
comparison of x0/X values obtained from the M3 protoconid and hypoconid
lobes when investigating birth seasonality. We applied an
average correction of −0.21 to the x0/X values obtained from hypoconid
lobes in order to compare them with previously published protoconid
(anterior) lobe values.
4.4. Length of the calving period
Fig. 7 is a representation of inter-individual variability in the tooth
crown location of the maximum δ18
O value (x0, normalised to the
length of the tooth formed over a year X in order to eliminate the effects
of tooth size). This variability is an expression of the duration of the
annual cycle, over which all individuals were born. At Chotěbudice,
most individuals were born over a two to three month period (0.21 or
21% of a yearly cycle). Considering the temporal breadth of the sample
(potentially a couple of centuries for the most representative LBK
phases at Chotěbudice) and its small size (ten specimens) due to the
inherent nature of most archaeological assemblages, caution is required
when interpreting the births distribution among what should be considered
a palimpsest herd. At Chotěbudice, the cattle kill-off patterns
for the different LBK phases show very similar trends over two centuries,
suggesting stability in cattle management strategies through
time (Kovačiková et al., 2012). Similarly, reduced variability in the
calving period may suggest a restricted calving period even over a time
scale of centuries. The status of CHO Bos1M3 (domestic vs wild) is
uncertain due to the large size of the molar, but it clusters with the main
group of individuals, meaning that it was born over the same period of
the annual cycle as domestic cattle. The only specimen from Černý Vůl
that could be modelled clusters with the main group at Chotěbudice.
5. Discussion
5.1. Management of cattle diet
The range of δ15
N values and the amplitude of inter-individual
variability are similar to those measured in other LBK bovine populations
from Central Europe (Fraser et al., 2013; Bickle and Whittle, 2013
and Fig. 1). Further interpretation of this δ15
N dataset would require
comparisons with values from other animals from local wild and anthropic
ecosystems. Most of the δ13
C values measured in bovine bone
collagen from Chotěbudice and Černý Vůl indicate feeding in open
areas (> −21‰). In this regard, they resemble those measured at
Vedrovice, Těšetice-Kyjovice and Brno-Starý Lískovec in the southeastern
Czech Republic, averaging −20.2‰, −20.2‰ and −20.3‰
respectively (Whittle et al., 2013b and Fig. 1). Analyses of pollen and
molluscs from sedimentary sequences in the dry lowlands of northern
Bohemia provided evidence for a mosaic landscape with pine-birch
forests and a steppe component (Pokorný et al., 2015). At Chotěbudice,
anthracological analyses highlighted the presence of oak wood, including
an important pine tree component (Kočár et al., 2008) and the
existence of a steppe was also suggested from the mollusc fauna
(Kovačiková et al., 2012). Indeed, our results could suggest either cattle
herding in open-canopy forests or forest boundaries, or preferential
grazing on persistent patches of steppe grassland. Nevertheless, a
denser forest component in the landscape is also evidenced in the
lowest δ13
C value of the dataset (−22.9‰), measured in CHO Bos25
(classified among the aurochs based on osteometric criteria). This value
is comparable to δ13
C values measured in late Boreal/early Atlantic
aurochs feeding in the dense Atlantic forest in Scandinavia (−24.2 to
−22.4‰; Noe-Nygaard et al., 2005a, 2005b). Three specimens, including
one aurochs (CHO Bos23), and one domestic bovine (CHO
Bos12) at Chotěbudice, and one Bos sp. at Černý Vůl (CerVBos12),
yielded intermediary δ13
C values (< −21‰), which could also indicate
a forest component.
The sequential δ13
C values measured in tooth enamel help to clarify
the contribution of forest resources to the cattle diet on a seasonal scale.
In all individuals but one (CHO Bos9M3), the δ13
C values are typical of
feeding in open areas all year round. Limited seasonal changes are
visible in the δ13
C series, although these would be partly attenuated by
the enamel mineralization delay (Balasse, 2002). When seasonal trends
are observed, the highest δ13
C values occur at the end of summer and
the lowest at the end of winter, in accordance with the moderate natural
variation in the δ13
C values of C3 plants on a seasonal scale
(Smedley et al., 1991). Winter provisioning with dried fodder harvested
from meadows, steppe grassland or even cultivated fields has been
discussed elsewhere (Saqalli et al., 2014). The harvesting of fodder
during the growing season and its postponed use until winter would
Fig. 3. Results from stable carbon (δ13
C) and nitrogen (δ15
N) isotope analysis of bone collagen from Chotěbudice and Černý Vůl Bos specimens.
R. Berthon et al. Journal of Anthropological Archaeology 51 (2018) 16–27
21
interfere with the seasonal cycle of variation in plant δ13
C values, by
increasing winter δ13
C values. The moderate inter-seasonal variation in
plant δ13
C values could in our case restrict the possibility of highlighting
this practice, unless the absence of inter-seasonal variations in
some δ13
C sequences (in CHO Bos10 M3 and CerV Bos2 M3, as opposed
to what is observed in CHO Bos2, Bos4, Bos7 and Bos8 for instance;
Figs. 4 and 5) might be an indication of this. However, it is not possible
to deduce clear evidence of winter foddering with summer grass from
this dataset.
In contrast to all the other specimens from Chotěbudice, CHO
Bos9M3 exhibits a higher range of variation in δ13
C values (−13.6 to
−11.2‰), and a marked seasonal variation pattern with lower δ13
C
values in winter, when δ18
O values are lowest. Considering a 14.1‰
13
C-enrichment factor between diet and enamel bioapatite (Cerling and
Harris, 1999), the lowest δ13
C value measured in enamel would correspond
to a diet δ13
C value of −27.3‰, reflecting feeding on plants
from forest understory (Drucker et al., 2008). Forest exploitation by
Chotěbudice LBK settlers (i.e. harvesting of logs, dried twigs and shoots)
was otherwise evidenced by botanical macro-remains (Kočár et al.,
2008). Cattle could have been brought to the forest or fed leaf reserves
Fig. 4. Results from stable carbon (δ13
C; black symbols) and oxygen (δ18
O; open symbols) isotope analysis (V-PDB) of cattle tooth enamel bioapatite from
Chotěbudice (CHO) (erj = enamel–root junction). *
CHO Bos1M3 to CHO Bos5M3 data from Kovačiková et al. (2012). Sampling on the hypoconid (hypo) or protoconid
(proto) lobe.
R. Berthon et al. Journal of Anthropological Archaeology 51 (2018) 16–27
22
collected for this purpose. Winter leaf foddering of cattle has been
suggested from similarly low δ13
C values in tooth enamel at MăguraBoldul
lui Mos Ivănus (Starčevo-Criş I phase, early 6th millennium cal
BC) in Romania (Balasse et al., 2013). At Măgura, this low winter δ13
C
signature was only observed in one individual (among five), suggesting
a similar system to that described at Chotěbudice. At both sites, winter
leaf foddering did not seem to be a prevailing practice but could rather
have been a response to an unusually difficult winter, unlike what was
shown for instance at the Middle Neolithic site of Bercy (4th millennium
BC) in northern France, where this pattern occurred more systematically
(Balasse et al., 2012a). Nevertheless, although the low δ13
C
values measured in CHO Bos9M3 actually indicate a dense forest
component in the surrounding landscape, the absence of such signals in
all other cattle teeth does not totally preclude the gathering of tree
branches from forest margins, in areas where the canopy effect on plant
δ13
C values would not apply.
Seasonal feeding on forest resources would not imprint bone collagen
as heavily as a year-round contribution. We tried to evaluate how
such seasonal contributions would be reflected in bone collagen values
on a pluri-annual scale, using the sequences measured in CHO Bos9M3
with winter feeding on forest resources. In order to compare enamel
sequential δ13
C values and bone collagen δ13
C values: (1) a mean annual
value was calculated from the sequential δ13
C values measured in
enamel over an annual cycle (using the period X), starting from the
apex; (2) the latter value was transposed to diet using a 14.1‰ 13
Cenrichment
factor between diet and enamel bioapatite (Cerling and
Harris, 1999); (3) diet values were transposed to collagen applying 5‰
spacing between diet and collagen δ13
C (Ambrose and Norr, 1993). The
Fig. 5. Results from stable carbon (δ13
C; black
symbols) and oxygen (δ18
O; open symbols) isotope
analysis (V-PDB) of tooth enamel bioapatite of
cattle from Černý Vůl (CerV) (erj = enamel–root
junction). Sampling on the hypoconid (hypo) or
protoconid (proto) lobe.
Fig. 6. Results from stable carbon (δ13
C; black symbols) and oxygen (δ18
O; open symbols) isotope analysis (V-PDB) of tooth enamel bioapatite of cattle from
Chotěbudice. Comparison between the hypoconid lobe (continuous line) and the protoconid lobe (dashed line). In CHO Bos7M3, the higher δ18
O values in the
hypoconid lobe are due to the accidental use of a more concentrated acetic acid during the pre-treatment (Supplementary material 3).
R. Berthon et al. Journal of Anthropological Archaeology 51 (2018) 16–27
23
enamel sequential δ13
C values measured in CHO Bos9M3 would correspond
to a bone collagen δ13
C value of −21.4‰. This value falls
within the intermediate δ13
C values measured in LBK settlements in
Europe as a whole (Fig. 1), which would not necessarily demonstrate
cattle feeding on forest resources. It is therefore possible that this
practice could have been overlooked in bone collagen datasets.
Fig. 8 compares the estimated diet δ13
C value of CHO Bos9 M3 (as a
reference for seasonal feeding on dense forest resources) and other
cattle M3s (as references for grazing in open areas all year round) to the
bone collagen values obtained from bovine bone collagen at Chotěbudice
and Černý Vůl. The undetermined zone, between the lowest value
estimated for feeding in open areas (δ13
C collagen −21.03‰) and the
value estimated for CHO Bos9M3 is very narrow, highlighting once
again the lack of resolution of the bone collagen dataset to investigate
diet on a seasonal scale. One Bos primigenius (CHO Bos23) and one Bos
taurus (CHO Bos12) fall within this undetermined zone. Apart from the
aurochs CHO Bos25, with an outstandingly low bone collagen δ13
C
value (−22.9‰), one Bos sp. from Černý Vůl (CerV Bos12, from the
SBK occupation) falls within the range of values possibly reflecting
seasonal feeding on forest resources. These occurrences are in a minority
compared to all the remaining samples with bone collagen δ13
C
values higher than −21‰, reflecting feeding in open areas all year
round. In conclusion, our data from Chotěbudice and Černý Vůl suggest
either a limited occurrence of dense forests in the area, or a limited use
of forests for cattle herding by LBK farmers.
5.2. Cattle birth seasonality
Domestic cattle are physiologically capable of breeding throughout
the year, with no interruption in fertility. However, food resources are a
major restricting factor in reproductive performances and in the survival
probability of offspring (Burthe et al., 2011). The present-day
pattern of year-round calving is only observed in herds provided with
shelter and artificial food all year long. In contrast, cattle populations
living in unmanaged conditions exhibit seasonal reproduction in synchrony
with vegetation dynamics (Lecomte and Le Neveu, 1986).
Aseasonal breeding has been observed in Chillingham cattle feral populations,
in north-eastern England (Hall and Hall, 1988) but this
characteristic is predicted to reflect selection as a result of their previous
history of domestication (Burthe et al., 2011). At Chotěbudice,
the restricted birth period (two to three months) observed in bovines
resembles that described for modern free ranging cattle of the hardy
Highland cattle breed living in northern France, with calving occurring
mostly between May and July (Lecomte and Le Neveu, 1986), and in
Germany with calving occurring in March and April (Reinhardt et al.,
1986). The Grey cattle bred today in the Thrace region of Turkey, extensively
raised with no feeding supplements, or only a little in very
cold winters, also give birth over a two-month period in March and
April (Soysal and Kök, 2008). This strongly suggests environmental
constraints on the reproductive cycle of livestock at Chotěbudice. In
spite of moderate climate seasonality, and mild and wet winters during
the time period considered here (Kalis et al., 2003; Sánchez Goñi et al.,
2016), seasonal scarcity in grazing resources may have been a limiting
factor for the female fertility rate. Indeed, our stable isotope dataset
suggests that food supplements in winter were far from systematic in
the studied assemblage. Nevertheless, this constraint was not major, as
evidenced by the presence of two outliers (CHO Bos7M3 and CHO
Bos8M3), demonstrating the survival of calves born out of the main
calving season to adulthood (Fig. 7). Out of season births are reported
in free ranging cattle herds with seasonal calving: in the semi-wild
Scottish Highland breed, cattle kept with minimal management in the
Rhein-Taunus Naturpark in Germany, only 9% of the births were reported
to occur out of the main calving season (Reinhardt et al., 1986).
Interestingly, the presence of these outliers might also reflect the absence
of strong control on cattle reproduction by LBK herders.
Similar datasets from the LBK are not available for comparison.
Cattle birth seasonality at the earlier Neolithic site of Măgura-Boldul lui
Mos Ivănus (early 6th mill BC) was investigated on a limited number of
specimens (four) and on the upper third molar, precluding direct
comparison (Balasse et al., 2013). At Cheia in Romania, where the
Hamangia occupation (5200–4950 cal BC) spanned the latter part of the
occupation at Chotěbudice (LBK IIc to LBK IIIb), cattle birth distribution
occurred over a similar timescale (3 months) to the LBK site and over
the same period of the year, although maybe with a slightly earlier start
(Balasse et al., 2014). At the Middle Neolithic site of Bercy in Paris,
France (beginning of the 4th mill BC), cattle births extended over two
additional months. This pattern could have been favoured by a recurrent
winter contribution of forest resources to the cattle diet (Balasse
et al., 2012a).
At Chotěbudice, a restricted calving period would have directly
impacted the period of milk availability throughout the year. Firstly,
0.00
0.08
0.17
0.25
0.33
0.42
0.50
0.58
0.67
0.75
0.83
0.92
1.00
x0/X
Fig. 7. Cattle birth distribution, expressed by inter-individual variability in the
position of the maximum δ18
O values (x0) normalized by the period (X) of the
δ18
O in the lower M3 from Chotěbudice (CHO) and Černý Vůl (CerV). Values
obtained on the protoconid lobe. *
Values calculated from those obtained on the
hypoconid lobe with a correction of −0.21.
Fig. 8. Stable carbon isotope ratios
(δ13
C) of the diet of cattle from
Chotěbudice and Černý Vůl, estimated
from values measured in tooth enamel
(circles) and in bone collagen (squares).
R. Berthon et al. Journal of Anthropological Archaeology 51 (2018) 16–27
24
this is because shorter lactation may be assumed for Neolithic hardy
cows compared to present-day cattle: figures of 6 to 8 months are reported
in the literature for modern European unimproved cattle breeds
(Peške, 1994; Tresset, 1996; Bignal et al., 1999). Secondly, it is rather
unlikely that each cow gave birth to a calf every year: in cattle the
280 day gestation period is followed by an interval between calving and
the next conception, which may greatly vary according to the nutritional
state (Ball and Peters, 2004). For example a median figure of
174 days is given for the Chillingham unmanaged herd of white cattle
in northern England, meaning a calving interval of 454 days (Hall and
Hall, 1988). In the free-ranging Highland cattle of the Marais Vernier,
showing a strongly seasonal breeding pattern, food scarcity may also
lead to females simply skipping the mating period until the following
year (Lecomte and Le Neveu, 1986). In such conditions, maintaining a
level of milk production throughout the year would have been possible
in the case of aseasonal breeding, but would have been more critical if
breeding was restricted to a two to three-month period. The reduced
availability of fresh milk could have been overcome by storing dairy
products in the form of cheese. As mentioned above, ceramic sieves
dated to the classic and late phases of the LBK have been identified in
Poland and Germany, demonstrating that these communities mastered
the technology for cheese making (Bogucki, 1984; Salque et al., 2012,
2013). However, no such evidence was found in Bohemia, where no
perforated potsherd has yet been reported for the LBK, and where a
recent study on a ceramic assemblage from the late LBK phase at Bylany
(N = 163) did not reveal any milk lipid residues (Mátlová et al., 2017).
Even though the absence of milk residues in ceramic potsherds does not
preclude milk processing in perishable containers, for now there is no
archaeological evidence for cheese making in the area under study.
Consequently, the use of storable dairy products by LBK farmers in
Bohemia is still uncertain.
6. Conclusions
Previous stable isotope analyses in LBK bovine remains from the
western area of the LBK (southern and western Germany, eastern
France) have shown the reliance of cattle husbandry on resources from
the thick forests occupying most of the territory in these zones. In
contrast, bone collagen δ13
C data from other parts of central Europe
(Hungary, western Slovakia, the Czech Republic, Poland and Austria)
suggest that cattle herding predominantly took place in open areas,
although with low resolution on a seasonal scale. The sequential analysis
of δ13
C values in enamel conducted in the present study introduced
the seasonal dimension of diet management/landscape exploitation by
LBK herders, with important outcomes. The majority of cattle from
Chotěbudice and Černý Vůl grazed in open areas all year round. This
suggests that cattle were mainly kept in the steppe component of the
forest/steppe mosaic landscape prevailing at this time in Bohemia.
These results do not preclude the gathering of tree branches from forest
margins, in areas where the canopy effect on plant δ13
C values would
not apply: the δ13
C signal may reach its limit in this regard. Meanwhile,
the presence of a dense forest component was confirmed in the low δ13
C
value measured in one aurochs bone, and the seasonal exploitation of
this dense forest was also evidenced in one domestic bovine as winter
browsing/provision of fodder. The estimation of a bone collagen value
based on the integration of this seasonal signal on a (pluri)annual basis
also led to the conclusion that such a contribution may have been largely
overlooked when bone collagen δ13
C values alone were taken into
consideration. Future research should adopt this analytical strategy as a
complement to bone collagen analysis, in order to refine our understanding
of the modalities of forest exploitation by LBK herders on a
seasonal scale, which is the required scale for the analysis of agropastoral
systems.
The definition of the key parameter of cattle birth distribution is
also essential in order to obtain a clearer picture of work organisation
and animal product availability. This study provides the first dataset on
LBK cattle birth seasonality. At Chotěbudice, cattle births were shown
to occur mainly over a restricted two to three-month period, suggesting
environmental constraints on the cattle fertility cycle, and possible
seasonal fodder scarcity in a context where our stable isotope dataset
did not evidence recurrent food supplements in wintertime. A direct
consequence of a short birth period would be a relatively short period of
milk availability. In the absence of archaeological evidence for cheese
making in the LBK of Bohemia, uncertainty persists as to whether dairy
resources may have been secured by transforming them into storable
products. The development of similar works on other LBK assemblages
will help to define whether the traits identified at Chotěbudice are typical
of LBK husbandries, or whether regional diversity can be brought
to light. We thus call for the multiplication of such studies in order to
provide models with direct evidence of husbandry practices and zootechnical
parameters.
Acknowledgements
This work was supported by the ERC Starting Grant “SIANHE” [GA
202881, 2008–2014] for stable isotope analysis and a post-doctoral
fellowship to Rémi Berthon. Stable isotope analyses were performed at
the SSMIM (Paris) with the technical support of Joël Ughetto. English
editing by Louise Byrne.
Appendix A. Supplementary material
Supplementary data associated with this article can be found, in the
online version, at https://doi.org/10.1016/j.jaa.2018.05.002.
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