Journal of Archaeological Science 147 (2022) 105666

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Journal of Archaeological Science

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n-Alkanes and their carbon isotopes (δ13C) reveal seasonal foddering and
long-term corralling of pastoralist livestock in eastern Mongolia
Natalia Égüez a, b, c, *, Carolina Mallol b, Cheryl A. Makarewicz c
a
Graduate School Human Development in Landscapes. Institute for Prehistoric and Protohistoric Archaeology. Kiel University, 24118, Kiel, Germany
b
Archaeological Micromorphology and Biomarkers -AMBI Lab, Instituto Universitario de Bio-Orgánica Antonio González. Universidad de La Laguna, 38206, Tenerife,
Spain
c
Institute for Prehistoric and Protohistoric Archaeology. Kiel University, 24118, Kiel, Germany

A R T I C L E I N F O A B S T R A C T

Keywords: The compound-specific δ13C values of n-alkanes from pastoralist winter campsites in Mongolia have great po­
Pastoralism tential to reconstruct dietary seasonality of livestock and local vegetation. We analysed leaf wax n-alkanes and
Ethnoarchaeology their carbon isotope ratios in common fodder plants (Artemisia mongolica, Chenopodium album, Elymus dahuricus,
n-Alkanes
Elymus sibiricus and Stipa sibirica) fed to domesticates during the winter months, and also in topsoils (0–8 cm)
Carbon isotope analysis
Plant biomarkers
from four livestock corrals in northern and southern Mongolia. n-Alkanes identified in fresh plant parts reflect
Mongolia signatures with mean δ13C stable isotope values of individual n-alkanes typical for C3 plants (− 34.1‰). Plants
from forest- and desert-steppe exhibit similar mean n-alkane δ13C values but significantly wider isotopic varia­
tion in the desert steppe flora. The non-anthropogenic control samples exhibit δ13C mean values significantly
higher from that of dung deposits and fresh plants for the same regions. Our data support previous research that
n-alkanes profiles and their carbon isotopic values vary depending on the local vegetation and environment,
while simultaneously demonstrating how anthropogenic activity, such as corralling of livestock, is recorded in
the n-alkane of domesticates dung validating the use of biomarker studies in pastoralist archaeological contexts
for distinguishing seasonal foddering, local environmental data and to an extent grazing habitats.

1. Introduction
Pastoralism has reliably supplied meat, milk, fats, and fibers to
people since the early Holocene. Today, extensive pastoral production
occurs in 25% of the global land area, mostly in dry rangelands of sa­
vannahs, grasslands, prairies, steppes and shrub lands, supporting
around 200 million subsistence pastoral households around the world
(Nori et al., 2005; Rust 2019). The term “pastoralism” usually refers to
an outsized importance of bovid livestock in subsistence economies,
with tradition of animal husbandry that may or may not include
mobility and some degree of agriculture, comprising different popula­
tion sizes and social organisation (Cribb 1991; Khazanov 1994; Hon­
eychurch and Makarewicz 2016; Hammer and Arbuckle, 2017). In a
broad sense, archaeologists use the term to refer to mobile societies
relying on herd animals for most of their subsistence economy (Chang
2015; Hammer and Arbuckle, 2017). This definition is problematic as
pastoralism is a complex human lifeway that encompasses many shapes
and is affected by diverse factors such as history, culture, politics and
ecology (Salzman 2004).
Establishing the subsistence and settlement activities of ancient
mobile pastoralists in the archaeological record is often difficult due to
their use of portable dwellings that leave scant traces on land surfaces
and high mobility resulting in minimal accumulation of occupation
debris in seasonally occupied habitation sites (David and Kramer 2001;
Salzman 2004; Biagetti 2014). Often identified though systematic
pedestrian surveys of landscape features, ancient pastoralist sites are
usually evidenced by low density artefact scatters (i.e., pottery, lithics,
and faunal remains) and remnants of pastoralist infrastructure struc­
tures including wells, platforms, and fencing-systems. Notably, one
central piece of pastoralist infrastructure-pens and corrals - contain
another archaeologically visible marker of pastoral activity: dung.
Manure is composed primarily of organic plant tissues reflecting the
composition vegetal forage – grasses, leaves, and browse-ingested by
livestock as well as minerals and nutrients including phosphorus,

* Corresponding author. Graduate School Human Development in Landscapes. Institute for Prehistoric and Protohistoric Archaeology. Kiel University, 24118, Kiel,
Germany.
E-mail address: neguezgo@ull.edu.es (N. Égüez).

https://doi.org/10.1016/j.jas.2022.105666
Received 28 October 2020; Received in revised form 11 August 2022; Accepted 6 September 2022
Available online 21 September 2022
0305-4403/© 2022 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-
nc-nd/4.0/).
N. Égüez et al.
nitrogen, and potassium. These organic and inorganic components of
manure can be identified in the archaeological record through concerted
use of a suite of analytical approaches, including micromorphological,
phosphate, phytolith, and spherulite (Shahack-Gross 2011).
Stable isotopic analyses of livestock bones and teeth, now commonly
employed to elucidate animal management practices involving animal
mobility, foddering, and birth seasonality, offer key insights on ancient
pastoralist husbandry practices at lifetime and seasonal scales (Balasse
et al., 2012; Makarewicz et al., 2018; Ventresca Miller et al., 2018;
Miller et al., 2019). However, practices that are key to promoting animal
survivorship in environmental harsh conditions, such as corralling, are
invisible to isotopic analyses of skeletal tissues. By packing animals
tightly together in corrals, herders help animal survive freezing winter
temperatures by consolidating animal body heat and insulating livestock
from the frozen ground, taking advantage of the thick layer of dung
deposited, often year after year, in the corral. In the bitterly cold ex­
tremes of the Eurasian steppe, winter corralling is essential to keeping
animals alive.
Dung deposited in corrals also preserves a biomolecular archive of
livestock dietary intake that offers information on the seasonality of
corralling and the broad composition of forage and fodder. Previously,
corralling and foddering information has been accessed through nitro­
gen and carbon isotope analyses of organic matter present in hypothe­
sized enclosure sediments (Shahack-Gross et al., 2008; Marshall et al.,
2018). Validated through analyses of ethnoarchaeological control
enclosure sediments and further supported by micromorphological and
phytolith analyses, this approach – while well-suited to historical
archaeological deposits, especially those that are further defined by
architectural elements, may be less successful in identifying corralling
and attendant foddering practices in more ancient deposits that are
subjected to N movements and transformations over short-and long time
scales (Hobbie and Ouimette 2009). 15N enrichment of sediments caused
by the introduction of exogenous nitrogen via manure, for example, is
not necessarily maintained over time due to leaching, and it may be
difficult to decouple the effects of manure fertilization with natural
process 15N-enrichment of local soils via N loss during nitrification and
denitrification, particularly in semi-arid and arid environments inhabi­
ted by mobile pastoralists.
Leaf wax n-alkanes are a highly robust biomolecule resistant to
diagenetic alternation in the burial environment over millennial time
scales (insert citation). The chain length and carbon (δ13C) isotopic
composition of n-alkanes offers information on their floristic origins and
generally used to reconstruct local environmental and vegetation con­
ditions in terrestrial environments. Here, we investigate seasonal fod­
dering practices through analysis of carbon isotope ratios measured
from leaf wax n-alkanes present in summer growth forage collected by
herders as winter fodder as well as dung accumulations present in winter
camp corrals. We sampled sites representing two distinct ecosystems
within the Mongolian steppe – the desert-steppe and forest-steppe. In
doing so, we characterise the origin of the ingested plants by the animals
that were corralled during winter and corroborate fodder and season­
ality of occupation. This approach sets the stage for the application of
molecular methods to characterise and interpret the archaeological
sedimentary organic matter deposited in dung-rich pastoral sites
archaeological pastoralist contexts around the globe.
1.1. Ancient Mongolian pastoralism
The Mongolian Bronze Age (2500-700 BCE) illustrates a moment of
great social interaction and transformations (e.g. transition to intense
animal husbandry and development of mobile pastoralism) that most
probably motivated first social inequalities and the appearance of
nomadic empires (Houle, 2016). The late Bronze Age Deer
Stone-Khirigsuur culture (or ‘DSK’) hints at an important link between
the nomadic herding life and the development of stone funerary mon­
uments such as khirigsuurs with their satellite features: horsehead
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Journal of Archaeological Science 147 (2022) 105666
mounds and stone circles (Fitzhugh, 2009). What is known today from
past Mongolian pastoral societies comes primarily from 1) the study of
those features and their associated materials (Makarewicz et al., 2018);
2) artefact technologies (Taylor et al., 2016); and 3) rock art (Jacob­
son-Tepfer, 2019). In addition, the emergence of DSK complexes has
been traditionally explained by climatic-driven changes in grassland
steppe environments, but the suggested environmental fluctuations are
still not clear (Klinge and Sauer, 2019). Although limited palaeoclimate
data exists (Murakami et al., 2010; Lehmkuhl et al., 2011), more robust
data is needed for most Mongolian regions. Moreover, local, regional
and global signals superimpose in the existing data and it is difficult to
distinguish between climatic and anthropogenic impacts, thus creating
an ambiguous palaeoclimate interpretation for the Holocene in
Mongolia (Lehmkuhl et al., 2011). Proxies derived from anthropogenic
contexts such as archaeological sedimentary deposits have received
little attention, and geoarchaeological information retrieved from
households or seasonal campsites -the archives containing data that
informs about how people lived day to day-is still comparatively few in
number (Honeychurch, 2015).
In a recent review, Wright (2021) describes the importance of
studying pastoralist settlements to tackle the nature of Bronze Age
communities but also to assert to what extent nomadic pastoralism was
central to their structure. Now we are sure that today’s domestic animals
and dairy consumption are present by the end of the late Bronze Age
(Wright, 2021; see also references therein), but local herding and
mobility practices are almost unknown. Establishing the seasonality of
pastoralist campsite occupation and the local climatic conditions in
which mobile communities wintered their animals, are main questions
in need of examination. The resulting data therefore, could be used to
explore complex cultural processes: the interactions of pastoral societies
with their surroundings, past resilience to environmental change and the
formation of cultural landscapes, to name a few current longstanding
archaeological debates in Mongolia and Central Asia pastoralism
studies.
1.2. n-Alkanes as a record of animal diet and local environmental
conditions
While pastoralist domestic settlement typically leaves only ephem­
eral traces of occupation, other infrastructure such as wells, enclosures,
and winter campsites, designed to enhance the management of their
mobile livestock leave more robust and visible traces on the landscape
(Hammer 2014; Rosen 2016). Several of these features, in particular
corrals, support burial environments that preserve biomolecular ar­
chives of pastoralist activity, potentially offering a means to document
ancient livestock herding practices. Dung deposits – a ubiquitous feature
of pastoralist campsites around the world, in particular offer a rich re­
cord of past livestock management practices associated with seasonality
of occupation, stocking rate, separation of animals by species, age, or sex
within enclosures, and dietary intake (Chang and Koster 1986; Macphail
et al., 2004; Shahack-Gross et al., 2008; Marshall et al., 2018). To date,
these aspects have been investigated through analysis of sediment
micromorphology, macrobotanical remains and phytoliths, and isotope
analyses of bulk organic fraction recovered from deposits associated
with dung depostion (Macphail and Goldberg 1995; Shahack-Gross
et al., 2003; Shahack-Gross, 2011; Linseele et al., 2013; Gur-Arieh et al.,
2018).
These high organic-matter dung deposits contain carbon-based
compounds including lipids, cellulose, proteins, chitins and lignins
that are undigestable in the rumen. Wax lipid n-alkanes are among the
carbon-based compounds that are well preserved in dung deposits (Carr
et al., 2010; Égüez and Makarewicz, 2018). n-Alkanes are chemically
inert, non-polar saturated hydrocarbon molecules that originate from
epicuticular waxes of vascular higher plants (Eglinton and Hamilton,
1967). These form a barrier primarily against water evaporation and
biodegradation caused by bacteria and fungi. Intracuticular waxes
N. Égüez et al.
prevent water transpiration while epicuticular waxes regulate light
reflection at the cuticle interface and wettability i.e., the ability of a
liquid to maintain contact with a solid surface (Eglinton and Hamilton,
1967; Koch and Ensikat 2008; Ardenghi et al., 2017). n-Alkanes, indi­
gestible to an extent, have provided relevant insights related to the
botanical composition of the diet in nutritional herbivore studies (Dove
and Mayes 2006; Oliveira and Silva 2007). They are also highly resistant
to microbial degradation, are well preserved in soils and lacustrine
sediments (Huang et al., 1995; Otto and Simpson 2005; Smith et al.,
2007; Garcin et al., 2012), and are easily extractable by chemical
methods from plant tissues, excrements and sediments (Mayes et al.,
1986; Malossini et al., 1996; Duncan et al., 1999; Ardenghi et al., 2017).
In addition, due to their non-polar single covalent bonds and the absence
of functional groups, n-alkanes often preserve authentic stable isotope
ratios that are stable at geological time scales (Cranwell, 1981; Schim­
melmann et al., 2006; Vajda, 2017; Seki et al., 2012; Ardenghi et al.,
2017). These properties make n-alkanes one of the most reliable bio­
markers for animal diet estimation (Duncan et al., 1999; Dove and
Mayes 2005; Dungait et al., 2010) and paleoenvironmental reconstruc­
tion (Sachse et al., 2006; Niedermeyer et al., 2010, 2016; Schemmel
et al., 2016).
The composition and abundance on n-alkanes in plants varies
depending on species, tissue type and growth stage (Wiesenberg et al.,
2004; Bush and McInerney, 2013; Diefendorf and Freimuth, 2017).
Plants produce a range of n-alkanes of different chain lengths, typically
exhibiting a strong predominance of odd-over-even numbered chains
due to decarboxylation from even-numbered fatty acid molecules
(Kolattukudy 1980). Generally, odd carbon numbered long-chained
(>C27) alkanes are characteristic components of terrestrial vegetation
(Eglinton and Hamilton, 1967), mid-chain homologues (C21–C25) char­
acterise submerged and floating aquatic macrophytes (Barnes and
Barnes, 1978; Cranwell, 1981; Ficken et al., 2000), and short-chain
homologues (C17–C21 n-alkanes) characterise aquatic algae, mosses
and ferns (Cranwell et al., 1987 and microbial organisms (Eckmeier and
Wiesenberg, 2009). Leaf wax carbon chains lengths are also determined
by environmental conditions, and as such n-alkane profiles derived from
sediments provide specific biota biomarkers (Samuels et al., 2008; Bush
and McInerney, 2015).
2. Materials and methods
2.1. Geographical and ethnographical setting
Mongolia is situated in strongly seasonal continental climate char­
acterized by relatively short warm summers and long, extremely cold
winters. Most precipitation falls during the later summer months with
annual precipitation ranging from less than 50 mm per year in the
warmer Gobi steppe desert to 400 mm in the cooler northern forest
steppe (Vandandorj et al., 2015). Grassland covers more than 80% of
Mongolian landscapes heavily grazed by sheep, goat, cattle, horses and,
in the north and central regions, also yaks. Mongolian herders manage
their livestock for their milk, meat, fat, and fibres using a wide variety of
husbandry strategies that adjust animal mobility, dietary intake, and
birth seasonality.
Winter camps are essential infrastructure maintained and used by
herders year after year that serve to shelter animals from harsh winter
conditions and thus help promote animal survival. Winter camps are
nestled within landscape features that shield both people and animals
from driving wind and, crucially, are also positioned in immediate
proximity to productive pasture reserved for winter grazing. The pre­
vailing wind patterns for Eastern Mongolia are similar whether at lower
elevations or at ridge-crest locations. The maximum wind speed is
concentrated during April and May, with winds decreasing fast after
those months and having the minimum wind speed during July and
August (Elliot et al. 2001). Fodder, harvested from more distant pastures
during the summer months, is also transported to and stored in winter
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Journal of Archaeological Science 147 (2022) 105666
camps (Makarewicz 2014; Égüez and Makarewicz, 2018). Herding
families maintain winter camps and associated pastures though
long-term generational use rights.
Winter camps, including the pastures and its associated buid infra­
structure, are considerable labor and materials investment by herding
families. Camps vary considerably between regions in their configura­
tion, density, and construction of corrals and shelters, depending on the
local availability of materials and herd composition. In the northern
forest-steppe, winter camps are arranged in large, complex layouts that
include wooden corrals and structures that house livestock according to
their species, sex, and/or age. These constructions are further insulated
by cattle dung used to chink spaces between timber slats, wattle-and-
dung fencing that sub-divides the interiors of roofed shelters, and
wind barriers 2–3 m in height constructed of timber and thick cattle
dung ‘bricks’. Further to the south where timber is absent and expensive
to transport, winter camps may consist of only one or two stone built
corrals, used to tightly pack together sheep and goats during the night,
and shelters with stone walls and wooden roofs for horses.
Herding famílies typically return to the same winter camp year after
year, using them for shelter and pasture from early November until late
April. Dung accumulates in both shelters and open corrals at varying
rates and thicknesses depending on the number of animals penned; herd
size is influenced by herder wealth, access to labor, subsistence needs
and econòmic goals (e.g., cashmere or wool production). Dung accu­
mulatations are left in place, often for years, to provide animals with
further insulation resulting in thick deposits that can reach up to 30 cm
in depth (Fig. 1). This compacted dung is then harvested, typically in
large blocks, from corral deposits and used for cooking fuel and con­
struction purposes (Égüez and Makarewicz, 2018).
2.2. Dung-rich penning deposits samples
Four penning deposits from enclosed corrals were sampled from two
winter campsites located in two contrasting geographical natural zones
during July 2015: Khentii (forest steppe) and Dornogovi (desert steppe)
provinces or aimags (Fig. 2). In the forest steppe, mean temperatures
average − 24 ◦ C/-14 ◦ C (min/max) and mean precipitation is 20 mm
during the winter (November–March), while during summer (June­
–August) are 10◦ /22 ◦ C (min/max) and 104 mm respectively. For the
steppe desert, mean winter temperatures average are − 26 ◦ C/-16 ◦ C
(min/max) and precipitation is 10 mm, while during summer are 14 ◦ C/
26 ◦ C (min/max) and 61 mm (Vandandorj et al., 2015).
Campsite 7 is closely located to a riparian forest within the rich
grasslands of Bayan-Ovoo in the forest steppe of Khentii province
(Fig. 2b). Vegetation communities in the lowlands are dominated by
Poa, Stipa and Leymus spp (Gunin et al., 1999). Sheep and goat corrals
are not roofed, and supported thick, well-preserved dung accumulations
reaching ca 30 cm in depth. Campsite 54 is sheltered between the rocky
outcrops that configure the semi-arid landscape of Dornogovi province
(Fig. 2c). Marshes appear in the low areas when precipitation amount
increases exceptionally, but often these are dry with stands of feather
grasses, mostly Cleistogenes, Artemisia and Stipa spp. (Gunin et al., 1999).
Here, corrals were also not roofed and supported visible, albeit poorly
preserved, dung deposits ca. 8 cm in depth..
In addition, two non-anthropic areas i.e., areas with no observable
animal droppings, approximately 1 km distant from each campsite were
sampled for natural organic matter content control (1 and 2) (Table 1).
To avoid contamination, each sediment sample was collected and
wrapped using aluminium foil.
2.3. Plant samples
Here, we take advantage of the continental climatic seasonality that
influences floral growth and plant tissue isotopic composition in order to
detect the potential range of δ13C variation in plant n-alkanes in
different environments and plant parts (Dove et al., 1996; Wiesenberg
N. Égüez et al.
Fig. 1. Example of winter campsites and dung deposits in a and b) the forest-steppe region of Dadal sum (Khentii aimag), and c and d) the steppe-desert region of
Dalanjargalan sum (Dornogovi aimag). Images: N. Égüez.
et al., 2004; Jansen et al., 2007; Huang et al., 2011; Jambrina-Enríquez
et al., 2018). In order to further establish the dietary origins of bovid
dung, we measured n-alkane compositions by analysing different plant
parts (i.e. leaves, stems, flowers and seeds) Here, a total of eight plant
samples were collected in order to establish the range of carbon isotopic
variation in the n-alkanes in floral growth from the steppe-desert and
forest steppe. These included five different species that are commonly
used during winter for animal fodder (see Jigjidsuren and Johnson,
2003), and thus expected to be present in the dung: Artemisia mongolica
(n = 2), Chenopodium album (n = 3), Elymus dahuricus (n = 1), Elymus
sibiricus (n = 1) and Stipa sibirica (n = 1). Plants were selected on the
basis of stage of growth, and only large mature plants were collected as
n-alkane amount production remain stable at this plant life-stage in
contrast to early stages when production is variable (Tipple et al., 2013).
Plants were collected in whole, air dried in the field, and dry-stored in
paper bags until analysis in the laboratory.
2.4. Lipid extraction, analysis and quantification
Plants were first cleaned with Milli-Q® ultra-pure water and then
dried at 60 ◦ C for 48 h. 1 g of each plant parts was separated from the
plant, treated with liquid nitrogen, and then ground and homogenised
with an agate mortar. 1 g of each of dung-rich sediment samples and
control sediment samples were dried at 60 ◦ C for 48 h and then homo­
genised using an agate mortar. Before extractions, all glassware and
glass wool was placed in a muffle oven at 450 ◦ C for 10 h to eliminate
any residual organic compounds on glassware. The total lipid extract
(TLE) was extracted in 10 mL 9:1 dichloromethane/methanol (DCM/
MeOH) following 3 cycles of 30 min in a sonicator and 10 min in a
centrifuge (4700 rpm). The TLE was then evaporated and separated into
six fractions of different polarity using solid phase extraction (SPE) via
silica gel column chromatography. The n-alkane fraction was eluted
with 3/8 of dead volume in n-hexane. After the addition of a 8 mg/L of an
internal standard (IS) 5α-androstane to the resulting fraction, the total
volume was completed with 150 μL of DCM (Jambrina-Enríquez et al.,
2018).
4
Journal of Archaeological Science 147 (2022) 105666
The compounds in the n-alkane fraction were identified and quan­
tified by gas chromatography with a coupled detection and mass-
selective detector (GC-Agilent 7890 B, MSD Agilent 5977 A) equipped
with an HP-5MS capillary column (30 m, ID: 250 μm, film thickness
0.25 μm). The GC was programmed to an initial temperature of 70 ◦ C for
2 min, heated with a heating rate of 12 ◦ C/min to 140 ◦ C and to final
temperature of 320 ◦ C with a heating rate of 3 ◦ C/min and held for 15
min, with helium as the carrier gas (2 mL/min). The total run time was
82.83 min. The multimode injector was held at a split ratio of 5:1 at an
initial temperature of 70 ◦ C during 0.85 min and heated to 300 ◦ C at
72 ◦ C/min. All measurements were repeated two times. The MS was
operated in full scan mode (m/z 40–580) with an electron ionization
energy of 70 eV and with the temperatures of the ion source and
quadrupole set at 230 ◦ C and 150 ◦ C, respectively. Quantification was
carried out taking the four most intense fragment ions (m/z 43, 57, 71
and 85, and m/z 67, 95, 81 and 245 for IS) for n-alkanes and the total ion
chromatogram for the rest of the analytes. Compounds were identified
by comparison of their retention times and mass spectra with those of
reference compounds (mix C8–C40 alkanes and 5α-androstane, Supelco)
and comparison with the NIST mass spectra library. Quantification of n-
alkanes was based on calibration curves obtained by plotting the ratio
Area/AreaIS versus the concentration of each reference compound.
Correlation coefficients were higher than 0.995. The n-alkane concen­
tration is expressed as μg of individual compound per gram of dry
sample (μg gds− 1).
Common indexes that aid in the environmental interpretation of
occurring wax n-alkanes abundances are the average chain length (ACL)
and the carbon preference index (CPI). ACL is the weighted average of
the various carbon chain lengths, whereas CPI measures the relative
abundance of odd over even carbon (Duan and He, 2011; Bush and
McInerney, 2015). The n-alkane distribution was calculated according
to the CPI using the Bray and Evans equation (1961) (Eq. 1):
CPI25-33 = [(ΣC25-33odd/ΣC24-32even) + (ΣC25-C33odd/ΣC26-C34even)] ×
0.5.
ACL was calculated following Freeman and Pancost (2014). For total
n-alkanes we used the C15–C33 interval (Eq. 2):
N. Égüez et al. Journal of Archaeological Science 147 (2022) 105666

Fig. 2. a) Sampled locations in Mongolia (base map: elevation SRTM 3.0 1 arc second), including the five different plant species collected for this study (Artemisia
mongolica, Chenopodium album, Elymus dahuricus, Elymus sibiricus and Stipa sibirica). Brackets indicate the sample number (Table 1), b) Winter campsite 7, and c)
Winter campsite 54 in relation to local topography. Note the organic-rich dark brown dung deposits in Campsite 7 in contrast to the poor-preserved dung deposits in
Campsite 54. (For interpretation of the references to color in this figure legend, the reader is referred to the Web version of this article.)

Table 1
Summary of the studied samples with basic parameters. Coordinates were recorded in the field by the authors with a handheld GPS.
Sample Natural zone Province Lat Long

Dung Campsite 7 Forest steppe Khentii 49.070275 111.627992

Dung Campsite 54 Desert steppe Dornogovi 45.631038 108.651634
Control sediment 1 Forest steppe Khentii 49.074467 111.640175
Control sediment 2 Desert steppe Dornogovi 45.624320 108.658120

Plant family Photosynthetic pathway

Artemisia mongolica 1 Forest steppe Khentii 49.070366 111.627833 Asteraceae C3

Artemisia mongolica 2 Forest steppe Khentii 48.966686 111.491767 Asteraceae C3
Chenopodium album 1 Desert steppe Sukhbaatar 45.372728 114.119124 Chenopodiaceae C3
Chenopodium album 2 Desert steppe Dornogovi 45.582287 108.624598 Chenopodiaceae C3
Chenopodium album 3 Desert steppe Dornogovi 45.582287 108.624598 Chenopodiaceae C3
Elymus dahuricus Desert steppe Sukhbaatar 45.327778 114.121405 Poaceae C3
Elymus sibiricus Forest steppe Khentii 49.070366 111.627833 Poaceae C3
Stipa sibirica Desert steppe Sukhbaatar 45.372728 114.119124 Poaceae C3

ACL15-33total = Σ(Ci × [Ci])/Σ[Ci]; 15 ≤ i ≤ 33.
The value [Ci] is the concentration of the n-alkane with i carbon
atoms.
2.5. Compound-specific carbon isotope analysis
Carbon isotope analysis by GC-IRMS was performed using a Thermo
Scientific Isotope Ratio Mass Spectrometer Delta V Advantaged coupled
to a GC Trace1310 through a Conflo IV interfaced with a temperature
converter GC Isolink II. n-Alkane fractions were injected by means of a
MultiMode injector (MMI) in splitless mode, with the temperature
increasing from 79 ◦ C (held 0.5 min) to 325 ◦ C (held 3min) at a rate
10 ◦ C⋅s− 1 and finally to 350 ◦ C (held 3min) at 14 ◦ C⋅s− 1. The GC was
fitted with a Trace Gold 5-MS (Thermo Scientific) fused silica capillary
column (30 m length x 0.25 mm i.d., 0.25 μm film thickness). Helium
was used as the carrier gas at a flow rate set at 1.5 mL min− 1. The
combustion reactor temperature was maintained at 1000 ◦ C. The tem­
perature programme comprised a 2 min isothermal period at 70 ◦ C

5
N. Égüez et al.
increasing to 140 ◦ C (held 2 min) at a rate of 12 ◦ C⋅min− 1, followed by
an increase period to 320 ◦ C (held 15 min) at 3 ◦ C⋅min− 1. Data acqui­
sition and processing were carried out using the Isodat 3.0 software
(Thermo Scientific). δ13C values were standardised to VPDB (Vienna Pee
Dee Belemnite) using an n-alkane Schiemmelmann type A6 mixture (n-
C16 to n-C30). The standard deviation of carbon isotope measurements
was better than 0.5‰.
3. Results
Distribution and abundance of n-alkanes for each plant sample
recovered from forest and steppe desert zones are reported in Figs. 3–4,
Fig. 3. Histograms indicating the distribution and abundance of n-alkanes in plant parts (flowers, stems and seeds). The concentrations are given in μg per g of dry
sample (μg/g sample).
6
Journal of Archaeological Science 147 (2022) 105666
and plant δ13C values are reported in Fig. 5. Distribution and abundance
of n-alkanes for dung deposits and sediment control samples are shown
in Fig. 6, and their δ13C values are shown in Fig. 7. Supplementary
material reporting statistical analyses for all samples (i.e. distribution,
abundance and δ13C values of n-alkanes) is available in SM1.
13
3.1. n-Alkane distributions and δ C values observed in fresh plants
All plant lipid profiles include mid- and long-chain n-alkanes (>nC21)
and a predominance of odd-numbered carbon chains. The range of
carbon numbers is slightly wider in stems compared to flowers and seeds
(C21–C35 vs C21–C33) (Fig. 3). Almost all plant species demonstrate a
N. Égüez et al.
Fig. 4. Barplots showing the distributions and abundance of n-alkanes in fresh plants parts and dung deposits of the two campsites with the associated error value (α
= 0.05).
Fig. 5. Biplot showing the n-alkane carbon isotope values for all samples.
Shapes are for type of sample and colors for plant species. The isotopic
composition for all samples fall into the range of C3 plants. (For interpretation
of the references to color in this figure legend, the reader is referred to the Web
version of this article.)
Cmax at C29 in stem parts, except for Artemisia mongolica where maximum
values are visible at C31. Flowers from all species exhibit Cmax at C29
while Cmax values in seeds are variable among species: with maximum
values observed at C23 for Elymus dahuricus, C27 for Stipa sibirica and C29
for Elymus sibiricus.
n-Alkanes are most abundant in flower parts, followed by seeds and
stems (x = 30.4 μg/g, x = 22.1 μg/g and x = 11.9 μg/g respectively).
Statistical observation for plant part pair-wise comparisons, resulted in
7
Journal of Archaeological Science 147 (2022) 105666
significance (p < 0.05) for all the possible pair groups, except for the
stem-flower grouping (ANOVA and Tukey’s test). When all plant parts
are grouped together, n-alkanes abundances between forest and desert
steppe show similar ratios (x = 21.1 μg/g and x = 15.7 μg/g respec­
tively) (Fig. 4). No statistical significance is observed for geographical
variation when the Mann-Whitney U Test was perfomed for not-
normally distributed n-alkane concentrations (w-statistic = 62, p-
value = 0.07022).
Overall, δ 13C values in plant tissues show significant variation be­
tween plant parts as well as between geographical regions. n-Alkane
carbon isotope values measured plants from dung from the forest steppe
and desert steppe exhibit different means averaging − 34.4‰ and
− 31.2‰ respectively, with plants from the desert steppe region, which
include C. album, E. dahuricus and S. sibirica, demonstrating wider range
in carbon isotope values from − 26.4 to − 40.4‰ compared to those from
the forest region, which include A. mongolica and E. Sibirucus, ranging
from − 30.7 to − 37.3‰.
Different plant parts exhibit different carbon isotopes ratios in n-al­
kanes. Mean δ13C vàlues for diferent plant parts, aggregated for forest
steppe and desert steppe specimens are for flowers − 36‰, seeds
− 32.6‰, and stems − 34.9‰ (Fig. 5). After separation by geographical
area, mean carbon isotopic values for plant parts are similar (δ13C forest-
steppe: flowers, − 35.7‰; − 33.6‰ seeds; − 35.7‰, stems; δ13C desert-
steppe; flowers − 36‰, seeds − 32.1‰ and stems − 34.6‰. The approx­
imately 1‰ enrichment in 13C observed in seeds and stem parts from
desert-steppe plants likley reflects lower wàter availability in the Gobi
steppe desert both in terms of lower precipitation levels and [poor
moisture retention in sandy soils] during the period of formation of stem
and seed tissues. Interestingly, δ13C values exhibited by the sediment
control samples show a higher value (2–4‰ difference).
13
3.2. n-Alkane distributions and δ C values in dung deposits
Penning deposits averaged 14.5 μg/g in n-alkanes and, for the forest-
steppe, yielded a n-alkane distribution that included odd-carbon chains
C29, C31 and C33 and for the desert-steppe region C27, C29, C31 (Fig. 6).
The lipid profile obtained from penning deposits from both campsites
includes medium and long-chain n-alkanes (>nC23) with a predomi­
nance of odd-over-even carbon number chains, thus compatible with
profiles of terrestrial vascular plants. Dry dung-rich sediments from the
forest steppe campsite 7, yielded the highest concentrations for C31
(33.1 μg/g), while the desert steppe campsite 54 yielded the highest
concentrations in n-alkane C29 (89.3 μg/g).
N. Égüez et al. Journal of Archaeological Science 147 (2022) 105666

Fig. 6. Histograms indicating the distribution and abundance of n-alkanes in dung corral from winter campsites 7 and 54, and forest-steppe and desert-steppe
sediment control samples. The concentrations are given in μg per g of dry sample (μg/g sample).

2–4‰ relative to nearby campsite 7. δ13C values from the control desert-
steppe sediment sample ranges from − 26.4‰ to − 31.1‰ and is signif­
icantly enriched in 13C ~4–6‰ relative to nearby campsite 54 (Fig. 5).

3.3. CPI and ACL values

CPI values of long-chained n-alkanes extracted from all samples

Fig. 7. Bloxplots showing the n-alkane carbon isotope values for dung deposits
from campsite 7 and 54. The isotopic composition for all samples fall into the
range of C3 plants.
The lipid profiles of the control sediments averaged 1.1 μg/g in n-
alkanes and includes short, medium and long-chain n-alkanes (>nC10).
The carbon chain length distribution from the plant wax-derived n-al­
kanes show a range of C10–C38 with higher n-alkane concentrations
present in the odd-numbered carbon chains. C21 is the most abundant
carbon number in steppe-desert control, while C29 is the most abundant
for the forest-steppe (Fig. 6).
Forest steppe dung n-alkanes exhibited δ13C vàlues of − 34.5‰ on
average, while n-alkanes from dung samples from the desert steppe were
significantly enriched in 13C on average 3.3‰ compared to forest-steppe
dung. δ13C values show similar intra-site range variation between cor­
rals A and B (ca. 2‰) for both campsites (Fig. 7). δ13C values for control
forest-steppe ranges from − 30.6 to − 32.6‰ and are enriched in 13C ~
ranged from 2.18 to 61.28 with values increasing according to deposi­
tional environment and plant part following: control < stem < grain <
flower < dung. CPI values are higher in forest samples when compared
to those of desert samples (mean of 23 vs. 8 respectively). The average
chain length (ACL) mean values range is higher in forest plants than in
desert plants (ca. 29 vs. 27) (Table 2 and Fig. 8).
Differences between dung deposits from campsites in forest and
desert are observable, both for CPI and ACL (Fig. 9). CPI values obtained
from campsite 7 displays a wide range of values (19–61), while campsite
54 values show a narrow range (6–18). In contrast, the reverse pattern
occurs for ACL values, showing a narrow range in campsite 7 and a large
range in campsite 54 with more intra-site variation between corrals 1
and 2 (29.61 and 27.97 respectively).
4. Discussion
4.1. ACL and CPI indexes indicate geographical variation and organic
matter maturation
The considerable differences between dung deposits in forest steppe
and desert steppe ACL values are likely due to differences in the plant
species and parts incorporated into dung deposits. Forest plants yield on
average shorter chain lengths than plants from grasslands or deserts,
because woody plants produce shorter ACL values due to the plant
growth under long periods of lower temperatures Cranwell, 1973). On
the contrary, non-woody plants that would have grown in environments

Table 2
CPI and ACL mean and median values for plant, dung and sediment samples and natural zones (forest and desert).
Grouping CPI ACL

x x Min. Max. x x Min. Max.

Type
Flower (n = 5) 14.76 12.59 8.23 24.10 28.82 28.57 28.23 29.63
Seed (n = 3) 11.21 13.77 6.05 14.97 26.63 26.11 25.10 28.81
Stem (n = 8) 8.89 9.63 2.18 25.75 27.14 28.51 21.23 30.16
Dung (n = 4) 21.20 16.95 7.02 61.28 29.31 29.61 27.97 30.35
Control (n = 2) 3.81 3.81 3.81 3.81 24.40 24.40 20.54 28.27
Natural Zone
Forest 23.78 19.65 12.66 61.28 29.26 29.42 28.27 30.35
Desert 8.38 7.02 2.18 16.95 25.93 27.97 20.54 29.61

8
N. Égüez et al.
Fig. 8. Set of boxplots showing ACL and CPI values for all samples, grouped by natural zones (forest-steppe and desert-steppe).
Fig. 9. Violin plot showing ACL and CPI indexes for dung deposits from campsite 7 and campsite 54.
with periods of higher temperatures would show higher ACL values due
to hydrological stress (Jiang et al., 2020). Moreover, it is suggested that
the modal carbon number is associated to both the temperature and
hydrological characteristics (Poynter et al., 1989; Poynter and Eglinton,
1990; Sachse et al., 2006), with lower latitudes resulting in higher modal
carbon numbers. However, contrary to previous studies, our samples
show higher modal carbon chain numbers in higher latitudes (forest
steppe) and lower in low latitudes (desert steppe) (Fig. 8) suggesting that
in extreme continental climates characterized by wide shifts in tem­
perature both daily and on a seasonal basis, low annual precipitation
levels, and low relative humidity, the modal carbon chains reflect pre­
cipitation records more than temperature variations (Bendle et al.,
2007). The small variation in ACL ranges for the forest steppe suggests
lower evapotranspiration, while the ACL great variation in the desert
steppe dung and stems n-alkanes is reflecting large rates of evapo­
transpiration due to more climatic variations (temperature and rainfall)
within this specific ecosystem (Rao et al., 2009; Sakari et al., 2008), a
pattern that is also observed in the alkanes from dung deposits (Fig. 9).
CPI values in all our samples, except the non-anthropogenic sedi­
ment control sample from desert, exhibit high odd-to-even predomi­
nance, resulting in alkanes from vascular land plants. In this study, CPI
index is used as a reference indicator for organic matter maturity. Our
results show that the n-alkane CPI data extracted from fresh plants in
Mongolia exhibits gradual increment with the increasing latitude
9
Journal of Archaeological Science 147 (2022) 105666
(Fig. 8), and again, dung samples from the corrals of winter campsites
also follow this criterion (Fig. 9), also likely reflecting lower intra-herd
variation in animal diets.
Therefore, our results indicate major contribution of organic matter
in forest deposits than in desert steppe contexts, since the latter are
strongly affected by erosional processes, producing immature soils.
Mature n-alkanes in our desert control sample could be derived from the
pre-existing sedimentary rock and/or the presence of algae, microalgae
or mosses that could grow when marshes are present (Lichtfouse et al.,
1997). Forest soils have more soil organic matter than grasslands or
croplands (Tsui et al., 2004; Wang et al., 2008) due to the condition of
litterfall and the slow decay of it in forest ecosystems. High long-chain
n-alkane carbon preference index values (>1) have been commonly
accepted as an indicator for terrestrial sources of sedimentary organic
matter and as a palaeoenvironmental indicator (Peters et al., 2005), as
higher CPI values usually correlate with cold and dry environments.
Moreover, Luo et al., 2011 demonstrated that CPI values decrease
(≈1/<1) during the transformation of organic matter, being this
occurrence caused by “dilution of n-alkanes without the odd-over-even
predominance from thermal cracking of hydrocarbons”. In addition,
CPI values that are close to one relate to recycled organic matter and
bacterial and marine microorganisms input (Kennicutt et al., 1987; El
Nemr et al., 2016).
N. Égüez et al.
4.2. n-Alkane distributions and δ13C values for dietary insights
The significant differences in δ13C values among plant parts (except
for the stem-flower group that showed no significant differences) as well
as between plants from forest steppe and desert steppe is likely caused
by plant growth stage and diverse environmental variables that are
known to impact floral δ13C values such as temperature and precipita­
tion. The composition and abundance on n-alkanes vary between plant
species, tissues and growth stage (Wiesenberg et al., 2004; Bush and
McInerney, 2013; Diefendorf and Freimuth, 2017). This heterogeneity
in n-alkane composition and concentration is largely controlled by stage
of tissue development, level of humidity and wax production and
regeneration stage (Richardson et al., 2005; Sachse et al., 2012; Kahmen
et al., 2011; Gao et al., 2014; Ardenghi et al., 2017). Altogether, varia­
tion in δ13C values in leaf wax may depend on plant type or water
availability which influences the δ13C values of plant tissues caused by
shifts in CO2 uptake and fixation (Δleaf). The dispersion of CO2 from the
atmosphere into the air-filled spaces within the leaf together with the
plant enzymatic activity produces a13C variation that can range between
− 6‰ and − 37‰ (Chikaraishi et al., 2004; Sachse et al., 2009). There­
fore, leaf water availability derived isotopic signals can be used for
determining the geographic provenance of plant materials and its
derived products (Dawson et al., 2002; West et al., 2006).
The carbon number distribution from n-alkanes from Mongolian
penning deposits ranges from C23 to C33, with maximum values for C31 at
Campsite 7 located the forest steppe, and in C29 for campsite 54, located
in the desert steppe, reflecting ingestion by sheep and goats of herba­
ceous plants and forest shrubby plants respectively (Fig. 6). Previous
research has shown that n-alkanes extracted from sediments discrimi­
nate between different vegetation forms with grassland n-alkanes pri­
marily dominated by C31, and deciduous trees/shrubs n-alkanes
characterized by high abundances of C29 (Cranwell 1973; Cranwell
et al., 1987; Bliedtner et al., 2018). Moreover, previous research in
Mongolian pastoral winter campsites reported penning deposits rich in
C29 and C31 n-alkanes reflecting deposition of dung containing the re­
mains of undigested stem-rich plant tissues (Égüez and Makarewicz,
2018).
4.3. Seasonal foddering detected in the n-alkane composition
Variation in modal carbon numbers observed in dung for both
campsites, emulate the distribution observed in fresh, summer season
plants collected for fodder, stored in camps, and then fed to animals
throughout the winter season. Thus, herders are providing fodder diets
that originate from the same ecosystem where the winter campsite is
located.
The n-alkane composition of sampled dung indicating vascular plants
and high n-alkane concentrations indicating the presence of flowers and
seeds, strongly suggests that dung was formed via ingestion and excre­
tion of summer season flora. The lack of difference between n-alkane
concentrations observed in stems of fresh summer season plants and
dung deposited during the winter months, for both forest- and desert-
steppe regions, shows that sheep and goats fed on plants that had
reached maturity during the summer months. With most precipitation
falling between June and July, floral growth comes into rapid fluores­
cence to produce seeds before the onset of cold temperatures in early
September (Shinoda et al., 2007). Moreover, n-alkanes accumulate in
spring and early summer during plant maturation, with amount pro­
duction (to some extent) regular through the rest of the growing season
(Eglinton and Hamilton, 1967; Avato et al., 1984; Bush and McInerney
2013; Tipple et al., 2013). Tall grasses, chenopods, and some herbaceous
growth are traditionally harvested by herders from late summer pastures
dried and then transported to winter camps and later fed to livestock
during the winter season (Makarewicz and Tuross 2006; Makarewicz
2014).
The similarity in the δ13C values between seed and dung n-alkanes is
10
Journal of Archaeological Science 147 (2022) 105666
unexpected but as herders provide their animals with mature plants
collected in late summer, it is possible that seeds and grains attached to
the stems of grasses and herbs were ingested by animals. Although not
the dominant n-alkane overall, the C27 n-alkane is the Cmax value in
seeds, and also present in the stems and dung deposits, especially on
campsite 54 (desert steppe). Thus, this signal may also be reflecting
seasonal seed input into the diet together with the stems, for example
from Stipa sibirica and Chenopodium album, which are abundant in the
desert-steppe. As reported in Spengler (2019), seeds of chenopodium spp.
were the most abundant assemblage recovered from dung burning ex­
periments, pointing to a high rate of survival from digestion process in
ruminants. Therefore, it is possible that the dung is signalling the pres­
ence of these seeds. It also has been long stablished that carbon number
maxima at C27 in sediments is indicative of forests related to cold cli­
mates and woody species presence (Cranwell 1973; Zech et al., 2009).
Therefore, the higher amount of C27 in our desert-steppe deposits sup­
ports instead the provision and ingestion of grass fodder rich in seeds
and stems collected in summer.
Carbon isotope values of n-alkanes also describe the effects of sea­
sonal foddering in the dung record deposited in Mongolian livestock
corrals. Low δ13C values resemble those of equatorial woods where
organic matter derived from plant tissues and litter are depleted in 13C
due to recycling effects associated with continuous litter deposition over
active soils (Benner et al., 1987; Cotrufo et al., 2013). Notably, these low
values (ca. − 34‰ to − 31‰ δ13C) are sharply different from reported
δ13C values range for the Chinese Inner Mongolian and northern China
grassland natural soils, which range between ca. − 20 and − 28‰
reflecting high local aridity levels (Chen et al., 2007; Yao et al., 2016).
Two mechanisms could explain the low values for dung deposits when
compared to natural soils. First, low δ13C values in dung-rich sediments
could result to high amounts of undigested plant remains rich in lignin
(i.e. stems). Lignin-C is heavily depleted up to 2.5‰ in 13C relative to
bulk plant material for legumes and up to 4.7‰ for grasses (Schweizer
et al., 1999). The carbon isotope composition of lignin detritus gradually
changes during biogeochemical processing in the decomposition phase
due to isotopic discrimination associated with the amino acids phenyl­
alanine and tyrosine, leaving a material that is relatively depleted in 13C
(Benner et al., 1987).
It appears that regular and extensive accumulation of lignin excreted
by livestock and deposited in penning structures follows similar degra­
dation pathways that invoke similar isotopic fractionation responses
during its decomposition. δ13C signatures from the newly formed soil
organic matter together with the products of plant decomposition can
alter or modify those from the original plant material (Schweizer et al.,
1999). Importantly, the massive accumulation of dung would produce a
rapid loss of more isotopically enriched 13C soluble constituents result­
ing in more depleted litter in 13C over time (Schweizer et al., 1999;
Asada et al., 2005). Therefore, continuous litter deposition in corral
dung deposits, which accumulates over the course of several years
providing a incubator for degradation action and organic matter matu­
ration, likely also promotes 13C depletion in plant n-alkanes.
5. Conclusions
Our results showed that preservation of n-alkanes in modern dung
deposits is very good and that, n-alkane homologue distributions,
coupled with the n-alkane δ13C values are a good proxy to identify
penning deposits in ecologically distinct sites in Mongolia and that those
are isotopically distinct from the natural soils. Our ACL and CPI values
directly reflect an open-range diet in desert environments, and a more
restricted diet in forest environments with environmental differences
between forest and desert steppes detected. However, the variability in
the alkane composition could be a result of genetic or life-phase changes,
external environmental elements and changes in the organic matter soil
input (Jansen and Wiesenberg 2017), so we agree with other authors
that caution is needed when using ACL as proxy for past changes in
N. Égüez et al.
vegetation (Wang et al., 2015) as many factors could be influencing the
presence of longer chain lengths.
This study is an example of how current advances in lipid and stable
isotope analytical techniques can set the stage to advance the archae­
ology of pastoralist societies. We have shown that sedimentary dung
deposits preserve digested plant lipids and that these can be character­
ized using molecular markers for distinguishing seasonal foddering,
local environmental data and to an extent grazing habitats. Further
biomarker research on this topic should: 1) involve larger data sets with
different Mongolian plant species to corroborate the environmental
differences observed here and 2) incorporate other proxies such as n-
alkane specific δD analysis to provide further insights into palae­
oenvironmental conditions, as δD variation is related to evapotranspi­
ration and soil evaporation-induced fractionation (Kahmen et al., 2011).
Research in this direction will furnish compound-specific stable isotope
datasets in Mongolia and allow us to explore important aspects of world
pastoralist societies such as husbandry, mobility and pasture land use in
the past.
Credit author statement
Natalia Égüez: Conceptualization, Methodology, Formal analysis,
Investigation, Writing - original draft, Writing - review & editing,
Visualization, Project administration, Funding acquisition. Carolina
Mallol: Methodology, Investigation, Resources, Writing - review &
editing, Supervision. Cheryl A. Makarewicz: Conceptualization, Inves­
tigation, Writing - original draft, Writing - review & editing, Supervi­
sion, Funding acquisition.
Declaration of competing interest
None.
Acknowledgements
The authors would like to thank all members of the Archaeological
Micromorphology and Biomarkers (AMBI Lab, Universidad de La
Laguna, Spain) and all members of the Archaeology Stable Isotopes Lab
(ASIL, Christian-Albrechts-Universität Kiel), including Adiyasuren
Molor, Sarah Martini, Margarita Jambrina-Enríquez, Sarah Pleuger,
Sarah Pederzani, Benjamin Ostwald and Johannes Haenel. NE wishes to
thank Francesc C. Conesa (Catalan Institute of Classical Archaeology) for
his valuable feedback on the analyses performed in R environment and
his comments on the original manuscrit while being a MSCA Fellow at
the University of Cambridge (MarginScapes, n. 794711). Special
acknowledgement is for the all the kind and generous Mongolian fam­
ilies that made this study possible. Funding: work was supported by a
PhD scholarship (NE) from the Graduate School “Human Development
in Landscapes” (Christian-Albrechts-Universität Kiel) in the frame of the
German Universities Excellence Initiative (grant number GSC 208).
Export permit number of collected samples is MN DE 6 1193.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.jas.2022.105666.
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