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Located at the crossroads between Africa, Europe and Asia, the Southern Caucasus is a prime location to study occupations by H. heidelbergensis, H. neanderthalensis and anatomically modern humans. Azokh Cave is an important site for the understanding of human evolution in its archaeological, palaeontological, environmental and ecological context. The main objective of this work is to use rodents to infer the climatic and environmental conditions that prevailed during the formation of the site. The small-mammal remains come from the archaeological excavation campaigns carried out in Azokh 1 in 2003, 2005, 2014, 2015 and 2018; they are from Unit V, Units III–IV and Unit II. The small-mammal assemblage is composed of at least 13 taxa: seven arvicoline, two cricetine, two gerbilline, one dipodid and one murine species. Units III–IV do not yield enough material to draw palaeoclimatic inferences. The palaeoclimatic conditions for Units V and II, ascertained by means of the bioclimatic model, suggest temperatures and precipitation similar to nowadays; the climate seems to be relatively warm-temperate in both units. The palaeoenvironmental reconstruction by means of habitat weighting points to an environment mainly composed of desert and steppe habitats, as well as portions of grassland and forest. This interpretation differs from that inferred from the large-mammal and archaeobotanical data, which indicate a woodland environment. These differences could be explained by the origin of the accumulation. There was no evidence of a major palaeoenvironmental or palaeoclimatic change between the Middle and Late Pleistocene layers, indicating favourable conditions throughout the study period.
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Archaeological and Anthropological Sciences (2022) 14: 96
https://doi.org/10.1007/s12520-022-01555-w
ORIGINAL PAPER
Palaeoecological reconstructions oftheMiddle toLate Pleistocene
occupations intheSouthern Caucasus using rodent assemblages
IvánRey‑Rodríguez1,2,3,4 · JuanManuelLópez‑García1,2 · EmmanuelleStoetzel3 · ChristianeDenys5 ·
JulieArnaud3,4 · SimonPartt6 · YolandaFernández‑Jalvo7 · TaniaKing8
Received: 31 January 2022 / Accepted: 1 April 2022 / Published online: 2 May 2022
© The Author(s) 2022, corrected publication 2022
Abstract
Located at the crossroads between Africa, Europe and Asia, the Southern Caucasus is a prime location to study occupa-
tions by H. heidelbergensis, H. neanderthalensis and anatomically modern humans. Azokh Cave is an important site for the
understanding of human evolution in its archaeological, palaeontological, environmental and ecological context. The main
objective of this work is to use rodents to infer the climatic and environmental conditions that prevailed during the formation
of the site. The small-mammal remains come from the archaeological excavation campaigns carried out in Azokh 1 in 2003,
2005, 2014, 2015 and 2018; they are from Unit V, Units III–IV and Unit II. The small-mammal assemblage is composed of
at least 13 taxa: seven arvicoline, two cricetine, two gerbilline, one dipodid and one murine species. Units III–IV do not yield
enough material to draw palaeoclimatic inferences. The palaeoclimatic conditions for Units V and II, ascertained by means of
the bioclimatic model, suggest temperatures and precipitation similar to nowadays; the climate seems to be relatively warm-
temperate in both units. The palaeoenvironmental reconstruction by means of habitat weighting points to an environment
mainly composed of desert and steppe habitats, as well as portions of grassland and forest. This interpretation differs from
that inferred from the large-mammal and archaeobotanical data, which indicate a woodland environment. These differences
could be explained by the origin of the accumulation. There was no evidence of a major palaeoenvironmental or palaeocli-
matic change between the Middle and Late Pleistocene layers, indicating favourable conditions throughout the study period.
Keywords Rodentia· Taxonomy· Taphonomy· Palaeoenvironment· Palaeoclimate
This article is part of the Topical Collection on Microvertebrate Studies in
Archaeological Contexts: Middle Paleolithic to early Holocene past environments
* Iván Rey-Rodríguez
ivanreyrguez@gmail.com; ivan.rey@urv.cat
Juan Manuel López-García
jmlopez@iphes.cat
Emmanuelle Stoetzel
emmanuelle.stoetzel@mnhn.fr
Christiane Denys
christiane.denys@mnhn.fr
Julie Arnaud
rndjmr@unife.it
Simon Parfitt
s.parfitt@ucl.ac.uk
Yolanda Fernández-Jalvo
y@mncn.csic.es
Tania King
taniacking@gmail.com
1 Departament d’Història i Història de l’Art, Universitat Rovira
i Virgili, Avinguda de Catalunya 35, 43002Tarragona, Spain
2 Institut Català de Paleoecologia Humana i Evolució Social
(IPHES), Zona Educacional 4, Campus Sescelades URV
(Edifici W3), 43007Tarragona, Spain
3 HNHP UMR 7194, CNRS/Muséum National d’Histoire
Naturelle, UPVD/Sorbonne Universités, Musée de lHomme,
Palais de Chaillot, 17 place du Trocadéro, 75016Paris, France
4 Sezione Di Scienze Preistoriche e Antropologiche,
Dipartimento Di Studi Umanistici, Università Degli Studi Di
Ferrara, C.so Ercole I d’Este, 32-44121Ferrara, Italy
5 ISYEB UMR 7205, CNRS/Muséum National d’Histoire
Naturelle/UPMC, EPHE/Sorbonne Universités, CP51, 57
Rue Cuvier, 75005Paris, France
6 Institute ofArchaeology, University College London, 31–34
Gordon Square, LondonWC1H0PY, UK
7 Museo Nacional de Ciencias Naturales (CSIC), José
Gutiérrez Abascal 2, 28006Madrid, Spain
8 Blandford Town Museum, Bere’s Yard,
Blandford,DorsetDT117AZ, UK
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96 Page 2 of 25
Introduction
The area of Western Asia plays an important role in efforts
to understand the biological history of human lineages and
their techno-cultural complexes during the Pleistocene, as
well as large- and small-mammal migrations (Agustí and
Lordkipanidze 2019; Bermúdez de Castro and Martinón-
Torres 2013; Belmaker etal. 2016). Western Asia was a
natural corridor and refuge between Africa and Eurasia
during the Pleistocene. The work of Abbate and Sagri
(2012) indicates that hominin dispersals occurred during
favourable climatic periods, and this area was very attrac-
tive for humans. This is evidenced by the high number of
sites located in this region, such as Dmanisi (Lordkipan-
idze etal. 2013; Coil etal. 2020), Qesem Cave (Maul etal.
2016), Misliya Cave (Hershkovitz etal. 2018; Weissbrod
and Weinstein-Evron 2020), Nesher Ramla (Hershkovitz
etal. 2021), Agithu-3 (Kandel etal. 2017) and Dzudzuana
Cave (Belmaker etal. 2016).
Studies of small vertebrates from archaeological sites
in this region have been on the increase in the last decade
(Belmaker and Hovers 2011; Demirel etal. 2011; Weiss-
brod and Zaidner 2014; Maul etal. 2015, 2016, 2020;
Smith etal. 2015; Belmaker etal. 2016; Parfitt 2016;
Kandel etal. 2017; Weissbrod and Weinstein-Evron 2020;
Rey-Rodríguez etal. 2020; Tilby etal. 2022). Here we
analyse the Azokh 1 site, which is one of the most com-
plete and, to date, one of the oldest archaeological sites
found in the Southern Caucasus.
The main objective of our work is to use rodents to infer
the climatic and environmental conditions that prevailed
during the formation of the site. We compare our results
with previous studies carried out in Azokh 1 Cave, as well
as other sites in the region where rodents have been ana-
lysed, in order to obtain a framework in which to discuss
the palaeoecological conditions where humans pursued
their activities in the Southern Caucasus.
Azokh Cave system
Azokh Cave takes its name from the village situated
nearby. The site is also known as Azykh Cave. It is located
in the Ishkhanaget river valley, Southern Caucasus (39°
37 15 N; 46° 59 32 E) (Asryan etal. 2020). It is an
important site for the understanding of human evolution
in its archaeological, palaeontological, environmental and
ecological context (Fernández-Jalvo etal. 2016b).
The cave is developed in thickly bedded Meso-
zoic carbonates. This system comprises a succession
of sub-rounded chambers oriented NNW to SSE and
interconnected for almost 130m. Several entrance pas-
sageways connect the inner parts of the cave to the exte-
rior, but geo-archaeological sediments have only been
found in Azokh 1, Azokh 2 and Azokh 5 (Fig.1). Our
research focused on Azokh 1, because this is the only
entrance found to date filled with Pleistocene-Holocene
sediments (Murray etal. 2016).
Azokh 1
Azokh 1 is a broadly linear chamber 40m long and 11.5m
high, with a WSW-ENE alignment. The cave has provided
evidence of occupation by hominins from the Middle Pleis-
tocene to the Holocene, and is the only well-stratified and
dated sequence from this period in the region (Asryan 2015).
It was discovered by M. Huseinov (also named Guseinov
by other authors) at the end of the 1950s, and the first exca-
vation started in the 1960s. Excavations of Azokh 1 were
carried out from the 1960s to the 1980s, yielding abundant
archaeological remains (Guseinov 2010). During this period,
the volume of the sediment excavated was about 70% of
the cave. Unfortunately, the information and descriptions of
the excavation procedures and finds before 1975 were too
schematic, making it difficult to interpret the area (Asryan
2015; Fernández-Jalvo etal. 2016b). The current excavations
began in 2002, after an initial survey of the site in 1999 and
2001. The excavation project at Azokh Cave was restarted
through the collaboration of an interdisciplinary research
team. The work focused primarily on the undisturbed, com-
plete sequence of deposits in the upper levels (I–V) at the
back of the cave. The systematic recovery and the detailed
recording of the material, with new methodologies applied,
provided high-quality information on the formation of the
site and on human behaviour and evolution (Asryan 2015).
In Azokh 1, nine stratigraphic units are identified, cur-
rently separated into two sediment sequences, which can
no longer be readily correlated due to the removal of all the
intervening stratigraphy (Fig.2) (Murray etal. 2010, 2016):
1. Sediment Sequence 1: contains Units IX to VI. The
test-trench excavation carried out has so far shown that
this sequence does not contain archaeological material.
The exact age of Sediment Sequence 1 remains unclear,
although previous palaeomagnetic results suggest that
it is probably older than the upper units and may be of
Early Pleistocene age.
2. Sediment Sequence 2: contains Units V to I. The new
excavations focus on these units; all of them present an
archaeological record, with an age range from the Mid-
dle (MIS 9–8) to the Late Pleistocene (MIS 5) (Units V
to II), with some Holocene sediments at the top (Unit I)
(Fernández-Jalvo etal. 2010, 2016b; Murray etal. 2010;
Asryan 2015; Asryan etal. 2017, 2020).
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Unit V
Unit V is located at the base of the sequence and is the larg-
est unit, being approximately 4.5m thick. It is divided into
two subunits: Vb (located at the base) and Va (above Vb).
Uranium-series dating indicated an age of ca. 200ka; race-
mization analysis (D/LAsp) indicated an age closer to ca.
300ka; and ESR dating suggested an age of 293 ± 23ka
(Murray etal. 2016). For the present study, we decided to
refer to it simply as Unit V without considering the subdivi-
sions, notably because the latter may be subject to further
revision (Murray etal. 2016).
According to Asryan etal. (2020), the lithic artifacts from
this unit include a relatively high presence of retouched
flakes and flake fragments, as well as a few cores. There
are no unknapped cobbles/pebbles or large tools (bifaces,
choppers, chopping tools). The characteristics of the Unit V
lithic assemblage indicate that most artifacts made from all
the raw materials were introduced as ready-made tools; how-
ever, the presence of a refit set may point to some isolated
insitu knapping. The techno-typological characteristics and
chronology (~ 300ka) of Unit V share similarities with the
Acheulo-Yabrudian techno-culture of the Caucasus. From
a broader perspective, this assemblage is late Acheulean or
pre-Mousterian, without large cutting tools.
In 1968, a fragment of hominin mandible was discovered
in Unit V by M. Huseinov (King etal. 2016). It is a right
mandible consisting of the posterior portion of the body
and the inferior part of the ramus. The study carried out by
Kasimova (2001) suggests that the mandible belonged to a
female aged 20–25years. This specimen has been assigned
to Homo heidelbergensis, based on the primitive features
(relief of the mylohyoid line) that it displays (King etal.
2016).
According to Van der Made etal. (2016), Unit V is
mainly composed of the following large mammals: carni-
vores (Canis aureus, Crocuta crocuta, Lynx sp., Felis chaus,
Panthera pardus, Ursus spelaeus), artiodactyls (Cervus ela-
phus, Capra aegagrus) and perissodactyls (Stephanorhi-
nus hemitoechus, Stephanorhinus kirchbergensis, Equus
Fig. 1 Azokh Cave system: a view of exterior of cave system, showing three main entrance passages (Azokh 1, 2 and 5); b plan of Azokh Cave
showing the main entrances and internal galleries (Murray etal. 2010); c location of Azokh Cave
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Archaeological and Anthropological Sciences (2022) 14: 96
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hydruntinus, Equus ferus). The faunal spectrum is mainly
composed of “interglacial” species; however, during glacial
times, the altitude of Azokh Cave (926m above sea level)
would have resulted in a harsh environment around the cave
(Van der Made etal. 2016).
According to the herpetofauna of Azokh 1 published by
Blain (2016), Unit V is composed of amphibians (Pseude-
pidalea viridis sensu lato, Ranidae/Hylidae indet. and
Pelobates cf. syriacus) and squamates (Pseudopus apo-
dus, Lacerta sp., Eryx jaculus, cf. Coronella austriaca, cf.
Elaphe sp., cf. “Coluber” sp. and “Colubrinae” indet.). The
herpetofauna seems to be more consistent with a meadow-
steppe environment, indicated by species such as Pelobates
syriacus.
The study of bats was carried out by Sevilla (2016) and
shows an assemblage in Unit V characteristic of an open-
ground landscape with steppe vegetation in Unit V, mainly
by the presence of Myotis blythii and the genus Rhinolophus.
The rodent remains were studied by Parfitt (2016), and these
results will be compared with the new study in “Results and
discussion” (see “Palaeoecological reconstruction of Azokh
1 Cave Sequence 2” for more details).
Allué (2016) identified a low number of charcoal remains
in Unit V, attributed to three taxa only: Prunus, Maloideae
and deciduous Quercus sp. These taxa reflect mild, humid
environmental conditions.
Unit IV
The contact between the top of Unit V and the overlying Unit
IV is 100–130cm thick but diffuse and irregular. No system-
atic excavation was carried out in this unit, although bone
and charcoal were recovered from the test-trench. ESR dat-
ing indicates an age of 205 ± 16ka for the contact between
Units IV and V (Fernández-Jalvo etal. 2016b).
Unit III
The transition between Unit IV and III is marked by a change
in the colour of the matrix. This unit is approximately 60cm
thick and contains charcoal, fossil bones and a few stone
tools. No date is available for Unit III.
The faunal list identified by Van der Made etal. (2016)
for Unit III is composed of carnivores (Panthera pardus,
Ursus spelaeus), artiodactyls (Cervus elaphus, Capra
aegagrus and Sus scrofa) and perissodactyls (Stephanorhi-
nus hemitoechus and Equus hydruntinus). Unit III can be
considered practically sterile in fossil bat remains (Sevilla
2016). Blain (2016) recognized several squamates: Lacerta
sp., Eryx jaculus, “Coluber” sp. and Vipera sp. The rodent
remains were studied by Parfitt (2016), and his results will
be commented on in the “Results anddiscussion” (see “Pal-
aeoecological reconstruction of Azokh 1 Cave Sequence 2”).
The material from this unit is not abundant enough to
draw climatic inferences.
Unit II
Unit II is 100–200cm thick and includes fossil fauna and
stone tools. Sediment diagenesis most likely caused by bat
guano has strongly affected the preservation of fossil bones
and some stone artifacts in this unit. ESR dating indicates an
age of 184 ± 13ka for the bottom of Unit II and 100 ± 7ka
for the contact between Units I and II (Fernández-Jalvo etal.
2016b).
Lithic artifacts were studied by Asryan etal. (2015,
2020), showing that the operative chain of different raw
materials is based primarily on knapping products, includ-
ing flakes, flake fragments and broken flakes. Levallois tech-
nology clearly dominates the core assemblage and knap-
ping products. The small size of most cores and the further
retouch of some indicate maximum exploitation and use of
good-quality, non-local raw materials. These characteristics
imply that almost all artifacts entered this area of Unit II as
ready-made tools, with most presenting a clearly predeter-
mined character. There may have been some insitu knapping
and retouching activities, as suggested by the presence of
knapping waste.
During the excavations in August 2010, an isolated homi-
nin tooth was discovered in this unit and studied by King
etal. (2016). It was identified as an upper left first permanent
molar that showed typical features of Homo neandertha-
lensis, on the basis of its morphology (swollen hypocone
and skewed shape) and taurodontism, as well as the crown
dimensions and root robusticity.
The list of large mammals (Van der Made etal. 2016) is
composed of carnivores (Panthera pardus, Ursus spelaeus,
Ursus sp., Vulpes vulpes and Canis lupus), artiodactyls (Cer-
vus elaphus, Capra aegagrus, Saiga tatarica, Dama sp. and
Sus scrofa) and perissodactyls (Stephanorhinus kirchbergen-
sis). The faunal spectrum suggests that this unit was char-
acterized by a typical “interglacial” temperate environment.
Unit II yielded slightly more bat remains than Unit III
(Sevilla 2016). The bat representation in Unit II shows a
low diversity and hints at a change towards colder conditions
Fig. 2 Stratigraphy of Azokh 1: a cross section through the entrance
passage (facing NW) showing the extent of the cave sediments
remaining in the chamber. These are physically separated and are
labelled Sediment Sequences 1 and 2. The undated Sequence 1
includes Units VI to IX. The archaeological Sequence 2 (inside the
rectangle) includes Units I to V; b cross section (orthogonal to the
section shown in a) of Azokh 1 showing the keyhole shape of the pas-
sage; c stratigraphy of Sediment Sequence 1; d stratigraphy of Sedi-
ment Sequence 2 with dating results of archaeological units (photos
modified from Murray etal. (2016))
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Archaeological and Anthropological Sciences (2022) 14: 96
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96 Page 6 of 25
compared to Unit V, indicated notably by the presence of
Myotis mystacinus and the absence of Rhinolophus mehelyi
(Sevilla 2016).
According to Blain (2016), the fossil herpetofauna from
Unit II is composed of amphibians (Pseudepidalea viridis
sensu lato, Pelobates cf. syriacus) and squamates (Pseudo-
pus apodus, Lacerta sp., Ophisops elegans, Eryx jaculus, cf.
Coronella austriaca, cf. Elaphe sp., cf. “Coluber” sp., the
Vipera berus complex and Vipera sp.). This assemblage rep-
resents a drier period, with the presence of a representative
of the V. berus complex (probably V. ursinii) and the small
colubrine Coronella austriaca. A rodent study was carried
out by Parfitt (2016); see “Results and discussion” (“Palaeo-
ecological reconstruction of Azokh 1 Cave Sequence 2”).
According to the study carried out by Allué (2016), the
charcoal remains from Unit II show a wide diversity of taxa,
with Prunus, Acer, Quercus sp., Maloideae, Lonicera sp.,
Paliurus/Ziziphus sp., Celtis/Zelkova sp., Euonymus sp. and
Ulmaceae. This assemblage composition indicates mild,
humid environmental conditions.
Unit I
Unit I is 80–135cm thick. This unit presents consider-
able disturbance and reworking of the sediments by recent
mammal burrowing activity. Palaeolithic faunal and lithic
remains were recovered from these burrows together with
modern artifacts. Unit I evidences the presence of domestic
animals (Sus scrofa, Equus asinus, Equus caballus, Capra
hircus) and is considered to be of Holocene age (Van der
Made etal. 2016). The amphibians (Pseudepidalea viridis
sensu lato) and squamates (Agamidae indet., Pseudopus
apodus, Lacerta sp., Ophisops elegans, Lacertidae indet.,
Eryx jaculus, cf. Coronella austriaca, cf. Elaphe sp., cf.
Coluber” sp., Vipera sp.) present in this unit indicate an
environment corresponding to an arid mountain steppe
(Blain 2016).
The large-scale bioturbation has greatly interfered with
the internal stratigraphic details. Charcoal from the fumier
in Unit I provided a radiocarbon age of 157 ± 26years BP
(OxAC1419424). A Soviet coin from around the mid-1960s
was also found in 2006, arguing in favour of high biotur-
bation in this level (Fernández-Jalvo etal. 2016b). Conse-
quently, palaeoenvironmental interpretations must be treated
with caution.
In summary, Azokh Cave is an important site for sev-
eral reasons: (1) it is located on the migration route through
the Caucasus used by hominins and fauna at the crossroads
between Africa, Europe and Asia; (2) the site was occupied
by three hominin species (documented by human remains
and lithic artifacts): H. heidelbergensis (Azokh 1 Unit V)
(Fernández-Jalvo etal. 2010), H. neanderthalensis (Azokh
1 Unit II) and H. sapiens (Azokh 2 and 5) (Fernández-Jalvo
etal. 2016b); (3) it is a site that has been well studied using
a multidisciplinary approach (Fernández-Jalvo etal. 2016c)
including the study of lithic assemblages (Asryan 2015;
Asryan etal. 2017, 2020) as well as of geology, geomorphol-
ogy (Murray etal. 2016), taphonomy (Andrews etal. 2016;
Marin-Monfort etal. 2016), zooarchaeology, palaeontology
(Blain 2016; Parfitt 2016; Van der Made etal. 2016) and
anthracology (Allué 2016).
The aim of this study is to investigate the palaeoenviron-
mental and palaeoclimatic conditions that prevailed during
the formation of Azokh 1 Cave using these new data and to
compare our results with other sites in the study area and
with previous works carried out in Azokh 1 Cave. Before
undertaking our palaeoecological study, we had to identify
the origin of the accumulation and the potential influence of
this on the palaeoecological results.
Material andmethods
The rodent remains used in this study come from the archae-
ological excavation campaigns carried out in Azokh 1 in
2003, 2005, 2014, 2015 and 2018. They are from Sediment
Sequence 2: Unit V, Units III–IV and Unit II.
The samples comprise disarticulated bones and isolated
teeth that were collected in the field by water screening using
superimposed 5 and 0.5mm mesh screens. Most of the sam-
ples were sieved in the field by the excavation team, and
the resulting residues were air-dried and sorted in the site
laboratory. In subsequent years, the small-vertebrate bones
were picked out by hand and under a microscope. The mate-
rial was identified with a binocular microscope, and photos
were taken with a Dino-Lite (model AM7915MZTL). In
total, 855 samples were analysed: 251 from Unit V, 10 from
Units III–IV and 594 from Unit II. The material is housed
in the village of Azokh.
Taxonomy
We focused on rodents because these can be considered
one of the most useful tools for palaeoenvironmental and
palaeoclimatic reconstructions in archaeological sites (e.g.
López-García etal. 2010; Maul etal. 2015). The taxonomic
identification of the rodent remains is based mainly on molar
morphology and measurements (Rey-Rodríguez etal. 2020).
We focused especially on the lower m1 (the most discrimi-
nant) for Arvicolinae; mandibles and maxillae were used
for the taxonomic identification of Cricetidae, Gerbillinae
and Murinae.
The remains were identified to species level whenever
possible. We used specimens from the modern reference
collections of the Natural History Museum of London, the
Field Museum of Chicago and the American Museum of
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Archaeological and Anthropological Sciences (2022) 14: 96
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Natural History of New York. We also used comparative
morphological and biometric data from the literature, nota-
bly for Microtus (Coşkun 2016; Kryštufek & Shenbrot 2016;
Kryštufek and Vohralík 2009; López-García 2008; Rusin
2017; Shenbrot, Kryštufek & Molur 2016; Tesakov 2016),
Clethrionomys (Kryštufek etal. 2009, 2020), Chionomys
(Krystufek 2017; Kryštufek and Vohralík 2009; López-
García 2011), Cricetulus (Bogicevic etal. 2011; Kryštufek
etal. 2017; Sándor 2018), Mesocricetus (Kryštufek and
Vohralík 2009), Meriones (Coşkun 1999; Kryštufek and
Vohralík 2009; Darvish 2011; Dianat etal. 2017; Stoetzel
etal. 2017), Allactaga (Karami etal. 2008; Shenbrot 2009)
and Apodemus (López-García 2008; Kryštufek and Vohralík
2009; Bogicevic etal. 2011; Amori etal. 2016; Knitlová and
Horáček 2017).
For the genus Ellobius, we based our identifications on
data from the literature (Moradi Gharkheloo 2003; Kryštufek
and Vohralík 2009; Tesakov 2016), and we also applied geo-
metric morphometric methods (GMM) in order to detect
morphological differences and variations in shape and size,
following Rey-Rodríguez etal. (2021). The Ellobius first
lower molars were all photographed in occlusal view under
constant conditions with a digital camera. To investigate
the first lower molar size and shape, we combined 2D land-
marks (LM) and semi-landmarks (SLM) on the photographs
using the TPSdig2 v.2.32 software package (Rohlf 2016) for
two-dimensional geometric morphometric analyses. All the
analyses were performed with R (R Core Team 2020) using
the geomorph (Adams etal. 2020) and Morpho (Schlager
2017) packages.
For the genus Arvicola, we used the molar morphology
(Maul etal. 2020) as well as the enamel differentiation index
developed by Heinrich (1987), also named SDQ (Schmelz-
band-Differenzierungs-Quotient). The enamel differentiation
index was calculated in accordance with the formula:
SDQ = [Σ(teet × 100/leet)]/N.
teet: posterior part of the triangle.
leet: distal part of the triangle.
N: number of triangles.
where N refers to the number of dentine fields of the
studied tooth, teet (trailing edge enamel thickness) refers to
the maximum thickness of the posterior enamel band and
leet (leading edge enamel thickness) refers to the maximum
thickness of the anterior enamel band of each dentine field
(Heinrich 1987; Lozano-Fernández etal. 2013).
We also used the standard measurements of the total
length (L) and total width (W) of the molars in order to
identify some species, particularly among the genera Crice-
tulus and Mesocricetus.
The quantification of taxonomic frequencies was based on
standard indices used in zooarchaeological analyses, includ-
ing the number of identified specimens (NISP) and mini-
mum number of individuals (MNI) (Weissbrod and Zaidner
2014). The latter was estimated using the most abundant
skeletal element present in the assemblage (molars in our
case).
Taphonomy
A preliminary study was performed on the rodent remains
from Azokh 1 Cave. This was based on the systematic
descriptive method that examines the modifications of prey
bones induced by predation, focusing on the degree of diges-
tion observed on teeth during identification (Andrews 1990;
Fernández-Jalvo etal. 2016a).
Predation is closely related to ecology due to two main
factors: the density dependence of prey according to popu-
lation fluctuations and the trophic preferences of the preda-
tor. Accordingly, identifying the predator(s) makes it easier
to identify the nature and origin of the assemblage. It is
necessary to take these factors into account in undertaking
palaeoecological interpretations based on small-mammal
accumulations (Fernandez-Jalvo and Andrews 2016).
We performed a preliminary study focusing on a subsam-
ple composed of 100 incisors and 90 molars per unit (for
Unit V and Unit II), looking at the digestion intensity and
broadly defining the category of modification produced by
the predator. A taphonomic study was previously performed
by Andrews in Azokh 1 Cave (Andrews etal. 2016); in this
work, we include an analysis of the new material.
Palaeoenvironmental andpalaeoclimatic
reconstructions
Habitat weighting method
Palaeoecological interpretations derived from faunal data are
based on analyses of community composition (Belmaker and
Hovers 2011). The method used for the palaeoenvironmental
reconstruction is the habitat weighting method (Evans etal.
1981; Andrews 2006; modified by Blain etal. 2008; López-
García etal. 2011), which is based on the current distribu-
tion of each taxon in the habitat(s) where it can be found
nowadays, according to the values proposed by the IUCN
red list. We adapt the method to our studied area (Rey-Rod-
ríguez etal. 2020) in assuming that the Azokh 1 Cave spe-
cies had equivalent ecological requirements to their present-
day relatives. We consider the following types of habitats:
forest (Fo), a large area covered with trees; shrubland (Sh),
vegetation dominated by shrubs; grassland (Gr), an open
area covered with grass; desert (De), an area with little pre-
cipitation and no vegetation cover; wetland (We), an area
where water covers the soil; steppe (St), a dry grassy plain;
and rocky (Ro), a rocky or stony substrate. Each species has
a score of 1.00, which is divided between the habitats where
the species can be found at present (Table1).
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Bioclimatic model
In order to reconstruct the past climate at Azokh 1 Cave,
we applied the bioclimatic model (BM), which was devel-
oped by Hernández-Fernández (2001) and updated by
Royer etal. (2020). This method is based on the faunal
spectrum, assuming that small- and large-mammal spe-
cies can in general be ascribed to ten different climates
(Hernández-Fernández 2001; Hernández-Fernández
etal. 2007). It was first necessary to calculate the climatic
restriction index (CRIi = 1/n, where i is the climatic zone
where the species appear and n is the number of zones
where the species is present) and the bioclimatic compo-
nent (BCi = (Σ CRIi) 100/S, where i is the climatic zone
and S is the number of species). All statistical analyses
were performed using the R software package (R Core
Team 2020) using the script PalBER developed by Royer
etal. (2020) in order to calculate the bioclimatic spectra
and infer the climatic zone.
The different climatic groups defined by Hernández-
Fernández (2001) and Hernández-Fernández etal. (2007)
present in this work are as follows: II/III, transition tropi-
cal semi-arid; III, subtropical arid; IV, subtropical with
winter rains and summer droughts; VI, typical temperate
with winters that are cold but not very long, but summers
that are cool; VII, arid-temperate with large temperature
contrasts between winter and summer; VIII, cold-temperate
with cool summers and long cold winters (boreal) and IX,
arctic (Table2).
By means of the BM, we were able to estimate various
climatic parameters, such as the mean annual temperature
(MAT), the mean temperature of the coldest month (MTC),
the mean temperature of the warmest month (MTW) and the
mean annual precipitation (MAP). This method enabled us
Table 1 Scores attributed to
each rodent species found at
Azokh 1 Cave according to
its ecological requirements,
used for the habitat weighting
method: forest (Fo), shrubland
(Sh), grassland (Gr), desert
(De), wetland (We), steppe (St)
and rocky (Ro)
Fo Sh Gr De We St Ro
Clethrionomys glareolus 0.5 0.5
Ellobius lutescens 0.33 0.33 0.33
Microtus arvalis/socialis 0.33 0.33 0.33
Arvicola persicus 1
Chionomys gud 1
Chionomys nivalis 0.33 0.33 0.33
Cricetulus migratorius 1
Mesocricetus brandti 0.33 0.33 0.33
Meriones dahli 0.33 0.33 0.33
Meriones tristrami 0.5 0.5
Meriones libycus 0.33 0.33 0.33
Meriones persicus 0.5 0.5
Allactaga willamsi 0.5 0.5
Apodemus spp. 0.5 0.5
Table 2 Scores attributed
to each rodent species
found at Azokh 1 Cave for
the bioclimatic model (in
accordance with Hernández-
Fernández (2001); Hernández-
Fernández etal. (2007), updated
by Royer etal. (2020)). See
text for the significance of the
Roman numerals corresponding
to the climatic groups
Species II/III III IV VI VII VIII IX
Clethrionomys glareolus 0 0 0 0.333 0 0.333 0.333
Ellobius lutescens 0 0 0.5 0 0.5 0 0
Arvicola persicus 0 0 0.25 0.25 0.25 0.25 0
Microtus arvalis 0 0 0 0.5 0 0.5 0
Microtus socialis 0 0 0.5 0 0.5 0 0
Chionomys gud 0 0 0.333 0.333 0 0.333 0
Chionomys nivalis 0 0 0.333 0.333 0 0.333 0
Cricetulus migratorius 0 0 0.333 0.333 0.333 0 0
Mesocricetus brandti 0010000
Meriones libycus 0 0.5 0.5 0 0 0 0
Meriones tristrami 0 0 0.333 0.333 0 0.333 0
Meriones persicus 0 0.5 0.5 0 0 0 0
Allactaga williamsi 0.333 0 0.333 0 0.333 0 0
Apodemus spp. 0 0 0.333 0.333 0 0.333 0
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to infer the palaeoclimatic conditions that prevailed during
the Middle and Late Pleistocene at Azokh 1 Cave.
Results anddiscussion
Systematics, current distribution andecology
A total of 434 molars were identified following the identi-
fication keys, representing a minimum number of 237 indi-
viduals (MNI) (Table3). This sample comprises the new
remains from the 2003, 2005, 2014, 2015 and 2018 excava-
tion campaigns, the previous material having been analysed
by Parfitt (2016), as later discussed in “Palaeoecological
reconstruction of Azokh 1 Cave Sequence 2”.
Order Rodentia Bowdich, 1821
Family Cricetidae Fischer, 1817
Genus Cricetulus Milne-Edwards, 1867
Cricetulus migratorius Pallas, 1773
Material: 10 isolated teeth. Unit II: eight isolated teeth,
one left lower m1, two right lower m1, one right upper M1,
one left upper M1, one right upper M2 and two left upper
M2. Unit V: two isolated teeth, one left lower m1 and one
right maxilla with M1 and M2.
Discussion: the first molars (m1 and M1) are brachyo-
dont and cuspidate, with two longitudinal series of cusps.
Each series of cusps consists of three pairs. The m1 and
M1 are the largest, and the m3/M3 the smallest. The lower
m3 only has two pairs of cusps (Kryštufek and Vohralík
2009). We identified Cricetulus migratorius (Fig.3.1) in
Unit II and Unit V of Azokh 1 Cave in accordance with
the measurements (Table4) and the identification keys for
molars based on the morphology and arrangement of the
tubercles and cusps provided by Kryštufek and Vohralík
(2009). We also drew comparisons with the reference col-
lection from Iran, Afghanistan and Azerbaijan housed in the
Natural History Museum of London regarding the morphol-
ogy of the teeth. The grey hamster, or migratory hamster, is
the smallest hamster species (Bogicevic etal. 2011; Sándor
2018). We measured the teeth, and the results indicate that
our specimens are the same size as those from the reference
collection, excluding the small-sized hamster Allocricetus
(Parfitt 2016).
Habitat and distribution: Cricetulus migratorius
extends from eastern Europe through Russia and Central
Asia to Mongolia and western China (Kryštufek etal. 2017;
Kryštufek and Vohralík 2009). The habitats of this spe-
cies are mostly dry grasslands, steppes and semi-deserts.
Arid areas with relatively sparse vegetation are preferred
(Kryštufek etal. 2017; Maul etal. 2015).
Cricetulus sp.
Material: 10 isolated teeth. Unit II: six isolated teeth,
two left lower m1, two right lower m2 and two left lower
m2. Unit V: four isolated teeth, two left lower m1, one right
lower m1 and one left lower m2.
In our record, we recognized as Cricetulus sp. broken
teeth that could not be measured and identified to the spe-
cies level.
Genus Mesocricetus Nehring, 1898
Mesocricetus cf. brandti Nehring,
Material: 11 isolated teeth. Unit II: five isolated teeth,
one left lower mandible with m1 and m2, one left lower
Table 3 Representation of the
Azokh 1 Cave rodent species in
terms of number of identified
specimens (NISP), minimum
number of individuals (MNI)
and percentage of the MNI (%)
Unit V Unit III–IV Unit II
Taxa NISP MNI % NISP MNI % NISP MNI %
Cricetulus sp. 4 4 3.48 6 2 1.74
Cricetulus migratorius 2 1 0.87 8 2 1.74
Mesocricetus brandti 6 3 2.61 5 3 2.61
Clethrionomys glareolus - - - - - - 4 3 2.61
Arvicola ex. gr. persicus - - - 1 1 0.87
Chionomys gud 3 2 1.74
Chionomys nivalis - - - 4 3 2.61
Microtus gr. arvalis-socialis 154 77 66.96 7 5 71.43 135 75 65.22
Microtus (Terricola) spp. 13 7 6.09 1 1 14.29 7 4 3.48
Ellobius cf. lutescens 13 7 6.09 1 1 14.29 22 12 10.43
Meriones spp. 5 3 2.61 4 2 1.74
Meriones gr. dahli-libycus 4 3 2.61 1 1 0.87
Meriones gr. persicus-tristrami 6 3 2.61 2 2 1.74
Apodemus spp. 3 2 1.74 7 4 3.48
Allactaga sp. 2 2 1.74
Allactaga cf. williamsi 2 1 0.87 2 1 0.87
Total 217 115 100 9 7 100 208 115 100
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m1, three left upper M2 and one left upper M3. Unit V: six
isolated teeth, one right lower m1, one left upper M1, three
left upper M2 and one right upper maxilla with M1 and M2.
Discussion: the specimens from Azokh 1 Cave are
attributed to Mesocricetus brandti (Fig.3.2) on the basis
of the morphology of the teeth, the disposition of the cusps
and the size. The molars are significantly larger than Cri-
cetulus migratorius, but smaller than representatives of
the Cricetus genus. The first molars have six tubercles;
the second and third molars only four. The largest molars
are m1 and M1, whereas m2/M2 and m3/M3 are reduced
(Kryštufek and Vohralík 2009). In the region of Azokh,
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Mesocricetus raddei is also found, presenting a similar
morphology of the teeth to Mesocricetus brandti, but
compared with the NHM reference collection of the lat-
ter, Mesocricetus raddei is bigger (Table5).
Habitat and distribution: Mesocricetus brandti has
the largest distributional area of the species belonging to
the genus Mesocricetus, ranging from Anatolia, Transcau-
casia and southeast Dagestan to northwest Iran (Qazvin
in the east, Lorestan in the south; Lay 1967). This species
is found at altitudes ranging from sea level up to 2600m.
However, the primary range is from 1000 to 2200m. Mes-
ocricetus brandti is found in arid and semi-arid steppe
habitats in lowlands and in mountainous areas (Kryštufek,
Yigit & Amori 2015; Kryštufek and Vohralík 2009; Neu-
mann etal. 2017).
Subfamily Arvicolinae Gray, 1821
Genus Clethrionomys Tilesius, 1850
Clethrionomys glareolus Schreber, 1780
Material: four isolated teeth. Unit II: one right lower m1
and three left lower m1.
Description and discussion: In our sample, we identified
Clethrionomys glareolus (Fig.3.3) on the basis of the pres-
ence of roots and the morphology of the first lower molar
(m1) (Kryštufek etal. 2020). In our study region, there are
only two genera of Arvicolinae with rooted molars: Clethri-
onomys and Ellobius, and these can be easily differentiated
by their molar morphology. In Clethrionomys, the m1 dis-
plays two roots, and has five triangles with a highly variable
anteroconid complex; however, T4–T5 do not vary. T5 is
integrated into the anterior cap or more rarely entirely iso-
lated from it; the anterior cap is rarely simple and oval but
mostly broadly confluent with T5. There are three or four
re-entrant angles on the inner side and three on the outer side
(Kryštufek & Vohralík 2005).
Habitat and distribution: The bank vole (Clethriono-
mys glareolus) is widely distributed in the Palaearctic area,
which stretches from the British Isles through continental
Europe and Russia to Lake Baikal. It was present at Azokh
1 Cave in the past; however, this species no longer exists in
the study region today.
Regarding its habitat, it is present in all kinds of wood-
land, preferring densely vegetated clearings, woodland edges
and river and stream banks in forests. It can also be found in
scrub, parkland and hedges (Bergl etal. 2017).
Arvicola ex. gr. persicus de Filippi, 1865
Material: one isolated tooth. Unit II: one left lower m1.
Description and discussion: recent works have shown
that the Iranian phylogroup of water voles can be consid-
ered a valid species, named Arvicola persicus (Fig.3.4)
(Chevret etal. 2020; Mahmoudi etal. 2020). We chose to
follow Maul etal. (2020) in using the name Arvicola ex. gr.
persicus. In our material, only one m1 could be attributed to
the genus Arvicola based on its size and morphology. The
m1 is rootless, with clear Mimomys-type enamel differentia-
tion and crown cementum filling the re-entrant angle. The
m1 consists of an anterior cap (AC), five triangles (T) and a
posterior lobe (PL). Enamel thickness (SDQ) is a parameter
applied to lower molars for the identification of Arvicola
species. In our material, we could only use SDQ5 on the
triangles, because the AC and the PL were digested. We
obtained a value of 124, which assigns our sample to Arvi-
cola ex. gr. persicus according to Maul etal. (2020). The
shape of the anteroconid complex also corresponds to this
species, with a BRA3 that is not as deep as in A. nahalensis.
The Mimomys-type enamel differentiation excludes the pos-
sibility that it could be A. amphibius, the other current repre-
sentative of the genus in the region (Mahmoudi etal. 2020).
The SDQ values show that there is a close relation between
A. persicus from Iran and Turkey, A. italicus from Italy and
A. sapidus from Spain, because all of these display a SDQ
value > 100 (Maul etal. 2020). In our sample, Arvicola ex.
gr. persicus is the only possible assignation, considering the
geographic distribution of the other taxa.
Habitat and distribution: the precise ecology and
distribution of Arvicola persicus remain to be clarified
(Mahmoudi etal. 2020). However, the group of water voles,
within which our species is included, is always rather scarce
in dry areas, most probably because these animals require
suitable aquatic habitats, i.e. the nearby presence of perma-
nent water bodies such as rivers, streams and marshes. We
assume that A. persicus has a semi-aquatic lifestyle, indicat-
ing the presence of water habitats (Harrison and Bates 1991;
Maul etal. 2020; Mahmoudi etal. 2020). Nowadays, there
is a river called the Ishkhanaget near the site at the bottom
of the mountain, which may very well have existed in the
past and favoured the presence of Arvicola around the site.
Genus Chionomys Miller, 1908
Fig. 3 Some rodents identified at Azokh 1 Cave. 3.1 Cricetulus
migratorius, Azokh 1 Cave, 2003, Unit V, D45, Z: 224, right M1
and M2, number 221. 3.2 Mesocricetus brandti, Azokh 1 Cave,
2014, Unit II, E49, Z: 320–330, left m1, number 74. 3.3 Clethriono-
mys glareolus, Azokh 1 Cave, 2018, Unit II, G56, Z: 46.48, left m1
occlusal and lingual view, number 703. 3.4 Arvicola ex. gr. persicus,
Azokh 1 Cave, 2014, Unit II, G51, Z: 330–340, left m1, number 536.
3.5 Chionomys gud, Azokh 1 Cave, 2003, Unit V, D46, Z: 111, left
m1, number 349. 3.6 Chionomys nivalis, Azokh 1 Cave, 2014, Unit
II, F48, Z: 330–340, left m1, number 529. 3.7 Microtus gr. arvalis-
socialis, Azokh 1 Cave, 2014, Unit II, G53, Z: 330–340, right m1,
number 5. 3.8 Microtus (Terricola) sp., Azokh 1 Cave, 2005, Unit V,
E40, Z: 136, left m1, number 43. 3.9 Microtus (Terricola) sp., Azokh
1 Cave, 2005, Unit V, D45, right m1, number 170. 3.10 Ellobius cf.
lutescens, Azokh 1 Cave, 2015, Unit II, F51, Z: 320–330, left m1
occlusal and lingual view, number 226. 3.11 Meriones gr. persicus-
tristrami, Azokh 1 Cave, 2003, Unit V, D45, right m1 occlusal and
lingual view, number 169. 3.12 Meriones gr. dahli-libycus, Azokh 1
Cave, 2003, Unit V, D45, Z: 224, left M1, number 222. 3.13 Apode-
mus (Sylvaemus) sp., Azokh 1 Cave, 2014, F52, Z: 330–340, left m1,
number 80. 3.14 Allactaga cf. williamsi, Azokh 1 Cave, 2018, H49,
G50, right m1, number 635. Scale 1mm
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In our sample, eight isolated teeth show the typical traits
of the genus Chionomys. The enamel is of the Microtus
type with cement in the re-entrant angles (Kryštufek 2017;
Kryštufek and Vohralík 2005; López-García 2011). These
specimens differ from Microtus in that they only have five
triangles, and an AC with a typical rounded morphology.
Chionomys gudSatunin, 1909
Material: three isolated teeth. Unit V: one isolated right
lower m1 and two isolated left lower m1.
Description and discussion: in Unit V we observed three
m1, which mainly show four closed triangles, with the wear
facets of triangles T5 and T6 confluent with the anterior cap;
they are typical of the Caucasian snow vole, Chionomys gud
(Fig.3.5) (Kryštufek and Vohralík 2005; Sözen etal. 2009).
These specimens were also compared to the Chionomys rob-
erti specimens from the NHM of London and were found to
be quite different in appearance, notably in the configuration
of the triangles and the morphology of the AC.
Habitat and distribution: Chionomys gud is closely
associated with rocky habitats. It occurs in alpine mead-
ows, in sparse fir and spruce forests and in the valleys of
brooks and small rivers. It prefers more mesic habitats than
Chionomys nivalis (Kryštufek and Vohralík 2005).
Chionomys gud is endemic to the Caucasus and the east-
ernmost part of the Pontic Mountains of Turkey, and is not
represented today in Armenia (Kryštufek 1999).
Chionomys nivalis Martins, 1842
Material: four isolated teeth. Unit II: one right lower m1
and three left lower m1.
Description and discussion: in Unit II, four first lower
molars display five closed triangles and an anteroconid
complex (AC) with a morphology characteristic of the
nivalis morphotype (Fig.3.6), where triangles T6 and T7
are reduced and tightened at their base, and the anterior cap
is of an arrowhead or oval shape, inclined towards the labial
part (Nadachowski 1991; Kryštufek and Vohralík 2005).
Habitat and distribution: Chionomys nivalis has a
global distribution extending from southwestern Europe
through southeastern Europe to the Caucasus, Turkey, Israel,
Lebanon, Syria and Iran (Kryštufek 1999).
Regarding the habitat of Chionomys nivalis, the species
is present in open rocky areas, typically above the tree line
and with scarce vegetation cover (Amori 1999).
Chionomys sp.
Material: one isolated tooth. Unit II: one isolated left
lower m1.
We opted for the classification Chionomys sp. because
of the lack of discriminant characters in the broken tooth.
Genus Microtus Schrank, 1798
The molars identified in the genus Microtus are hyp-
sodont and arhizodont, with crown cementum in the re-
entrant angles and Microtus-like enamel differentiation.
Eight species of Microtus are currently recognized in the
southern Caucasus: Microtus arvalis, Microtus daghestani-
cus, Microtus levis, Microtus majori, Microtus nasarovi,
Microtus schelkovnikovi, Microtus schidlovskii and Microtus
socialis (Aşan Baydemir and Duman 2009; Golenishchev
etal. 2019; Firouz 2005). Microtus m1s are characterized
by four outer and five inner re-entrant angles, with a pos-
terior lobe (PL), seven closed triangles (T) and an anterior
cap (AC). In most of our material, triangles T4 and T5 are
closed, ruling out the subgenus Terricola.
Microtus is the most abundant taxon in all the sequence,
but identifying the species was difficult because we lack
good reference collections and comparative data from the lit-
erature. However, we were able to differentiate two groups,
based on the morphology of the lower m1.
Microtus gr. arvalis-socialis
Material: 296 isolated teeth. Unit II: 135 isolated teeth;
75 right lower m1, 60 left lower m1. Units IIIIV: seven
isolated teeth; five left lower m1 and two right lower m1.
Unit V: 154 isolated teeth; 77 right lower m1 and 77 left
lower m1.
Description and discussion: the species (Fig.3.7)
included in this group are Microtus arvalis, Microtus socia-
lis and Microtus guentheri. The triangles from T1 to T5 are
Table 4 Measurements of the Cricetulus migratorius first lower
molar, m1, (in mm): L, total length; W, width. NHM Natural History
Museum of London. NISP number of identified specimens
Cricetulus migratorius
Azokh 1, NISP: 4 NHM, NISP: 16
L Min–max 1.41.81 1.41176
Mean 1.56 1.57
W Min–max 0.580.77 0.740.95
Mean 0.68 0.85
Table 5 Measurements of
Mesocricetus brandti and
Mesocricetus raddei first lower
molar, m1, (in mm): L, total
length; W, width. NHMNatural
History Museum of London.
NISPnumber of identified
specimens
Mesocricetus brandti Mesocricetus raddei
Azokh 1, NISP: 3 NHM, NISP: 7 NHM, NISP: 4
L Min–max 2.14–2.19 2–2.28 2.39–2.58
Mean 2.17 2.14 2.52
W Min–max 1–1.07 1.02–1.2 1.3–1.11
Mean 1.03 1.1 1.23
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Archaeological and Anthropological Sciences (2022) 14: 96
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closed and opposite one another. T6–T7 are confluent and
parallel to BRA4 and LRA5 (Kryštufek and Vohralík 2005).
Habitat and distribution: Microtus species belonging
to the group arvalis-socialis mostly occur in steppe, shrub-
land or semi-desert habitats (Fernandez-Jalvo etal. 2016).
According to the IUCN Red List, all of them are currently
present in the southern Caucasus.
Microtus (Terricola) spp.
Material: 21 isolated teeth. Unit II: seven isolated teeth;
three right lower m1 and four left lower m1. Units IIIIV:
one isolated tooth; one left lower m1. Unit V: 13 isolated
teeth; seven right lower m1 and six left lower m1.
Description and discussion: the second group is Micro-
tus (Terricola) spp., (Fig.3.8 and 3.9) characterized by
broadly confluent T4 and T5, forming the pitymyan rhombus
(Kryštufek and Vohralík 2005). In our sample, we identified
at least two species, based on the size and the morphology
of the AC. There are several Terricola species in the region:
Microtus daghestanicus, Microtus nasarovi, Microtus majori
and Microtus schidlovskii. All of them are very close to one
another in their molar morphology, and without a good refer-
ence collection, it is not possible to identify them to species
level.
Habitat and distribution: Caucasian voles, belonging
to the Microtus (Terricola) spp. group, are found in a quite
wide range of habitats: Microtus daghestanicus and Micro-
tus nasarovi prefer pastures, alpine meadows and steppe;
Microtus majori favours clearings in forests and shrubland,
as well as alpine pastures; and Microtus schidlovskii is more
closely associated with xerophytic steppes and meadow
steppes. All of them are currently present in the southern
Caucasus, according to the IUCN Red List.
Genus Ellobius Fischer, 1814
Ellobius cf. lutescens Thomas,
Material: 36 isolated teeth. Unit II: 22 isolated teeth;
10 right lower m1 and 12 left lower m1. Units IIIIV: one
isolated tooth; one left lower m1. Unit V: 13 isolated teeth;
seven right lower m1 and six left lower m1.
Description and discussion: 36 isolated teeth show the
typical traits of the genus Ellobius (Miller 1896; Hinton
1962; Kretzoi 1969; Coşkun 2016; Kryštufek & Shenbrot
2016; Kryštufek and Vohralík 2005; Rusin 2017; Kryštufek
& Molur 2016; Tesakov 2016).
The Ellobius lower m1 is composed of the anterior cap
(AC), five triangles (T) with three buccal and four labial re-
entrant angles, and one posterior lobe (PL). Ellobius molars
are notably characterized by the presence of roots that are
clearly visible in adults and old individuals. Moreover, Ello-
bius molars lack cement in the re-entrant angles (Coşkun
2016).
For modern representatives, most of the discriminant
characters are based on skull morphology (Kaya etal.
2018) and external characters (Kryštufek and Vohralík
2005), whereas in an archaeological context we mostly
rely on isolated molars or broken maxillae and mandi-
bles. The lower m1 is quite similar among the different
current Ellobius species (Ellobius fuscocapillus, Ellobius
lutescens and Ellobius talpinus), but some specific mor-
phological characters have been pointed out in previous
studies: the AC is broad in Ellobius lutescens, narrow in
Ellobius talpinus and elongated in Ellobius fuscocapillus
(Maul etal. 2015); the distance between T4 and T5 (W)
and the total length (L) differ among the species, Ello-
bius fuscocapillus showing the largest teeth and Ellobius
talpinus the smallest (Rey-Rodríguez etal. 2020). How-
ever, these morphological and biometric characters are not
always clear or reliable, and the use of more powerful
methods is necessary to obtain reliable identifications. To
this end, we applied geometric morphometric analyses to
m1 molars following the methodology established by Rey-
Rodríguez etal. (2021) for Ellobius. We used specimens
from the modern reference collections of the Natural His-
tory Museum of London, the Field Museum of Chicago
and the American Museum of Natural History of New
York, and the archaeological sample is from Azokh 1 Cave
(Unit II and Unit V).
The PCA performed on the normalized landmarks and
sliding semi-landmarks of the right lower molar reveals
significant differences among the analysed species, the first
two principal components (PCs) accounting for 57.3% of the
total variance (Fig.4a).
The main variation along PC1 (41.4%) relates to the mor-
phology of the anterior cap, which is more flattened for the
positive values and more rounded for the negatives ones.
Ellobius talpinus occurs in the positive part of the PC1 axis
whereas Ellobius fuscocapillus and Ellobius lutescens are
located in the negative part, reflecting a broader and more
rounded AC. Along the PC2 (15.9%) scores, the positive
values reflect a rounded and less developed AC turned to the
labial side, whereas the negative values show an elongated
AC turned to the buccal side. In PC2, there is no clear dif-
ferentiation between the three species. However, it is worth
noting that the Ellobius lutescens specimens are located
principally in the upper half of the graphs, showing negative
values in PC2. We can thus confirm the species identifica-
tions and corroborate that Ellobius talpinus is not present in
the archaeological samples.
To look at possible allometric effects on the samples, we
performed a linear regression of the PCs onto the log of the
size of the centroid (Mitteroecker etal. 2015). Including size
allows us to discriminate E. lutescens from E. fuscocapillus,
the latter showing greater dimensions (Fig.4b). E. talpinus
presents a wide size range overlapping the two latter species
and is only distinguished by the molar conformation (PC1).
The archaeological remains from Azokh 1 Cave are distrib-
uted among E. lutescens and E. fuscocapillus (Fig.4b).
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Other criteria can allow species to be identified, such as
the configuration of the anterior cap (Maul etal. 2015), with
the transition between T4 and T5 being narrower in E. lutes-
cens than in E. fuscocapillus, leading to a smaller and more
closed AC in E. lutescens. By combining GMM and PCA
analyses with morphological criteria (AC and distribution
of the triangles), we were able to attribute the material from
Azokh 1 to the species Ellobius cf. lutescens (Fig.3.10).
Habitat and distribution: Ellobius species frequent
steppes, grasslands and semi-deserts in eastern Europe
and central Asia; these fossorial species are specialized in
subterranean life (Coşkun 2016; Kryštufek and Vohralík
2005). Ellobius lutescens (western mole vole) is distributed
in northwestern Iran, Iraq, Azerbaijan, Armenia and eastern
Anatolia (Thomas 1905; Ellerman and Morrison-Scott 1951;
Darlington 1957; Osborn 1962; Walker 1964; Lay 1967;
Hassinger 1973; Roberts 1977; Corbet 1978; Corbet and
Hill 1991; Coşkun 1997; Nowak 1999; Wilson and Reeder
2005, 2017; Kryštufek and Shenbrot 2016).
Subfamily Gerbillinae Gray, 1825
Genus Meriones Illiger, 1811
The genus Meriones is one of the most diverse among
the tribe Gerbillini in the Palaearctic region, particularly
in arid regions of Asia (Darvish 2011; Denys 2017). The
Meriones species currently reported in the southern Cauca-
sus are Meriones dahli, Meriones libycus, Meriones persi-
cus, Meriones tristrami and Meriones vinogradovi (Dianat
etal. 2017; Fernández-Jalvo etal. 2016b; Kryštufek and
Vohralík 2009). In our archaeological material from Azokh 1
unit V, III–IV and II we have identified at least two different
groups of Meriones species based on their molar size and
morphology, as well as the number of roots on the molars.
Meriones gr. persicus- tristrami
Material: nine isolated teeth. Unit II: two isolated teeth;
two right lower m1. Units III-IV: one isolated tooth; one
left upper M1. Unit V: six isolated teeth; three right lower
m1 and three left lower m1.
Description and discussion: the part of the mate-
rial from Azokh 1 Cave attributed to the genus Meriones
(Fig.3.11) displays the typical morphology of this group,
including semi-hypsodont molars with prismatic enamel
triangles linked by a longitudinal crest and with no trace of
cusps. In our sample, we identify first upper molars (M1)
with three roots, which is characteristic of Meriones per-
sicus and Meriones tristrami. Unfortunately, the dental
morphology of these two species is very similar; their m1
and M1 display three roots, the second molars (m2 and M2)
have two transverse plates and two roots, whereas the third
Fig. 4 aPrincipal component analysis of the normalized landmarks and sliding semi-landmarks and shape configuration at the extreme ends of
the PCs. b First principal component of size and shape including the reference collection and Azokh 1 Cave material
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molars (m3 and M3) are simple and rounded with a single
root (Coşkun 2016; Kryštufek and Vohralík 2009).
Habitat and distribution: the genus Meriones is distrib-
uted across North Africa, Central Asia, Turkey and Pakistan
(Darvish etal. 2014; Wilson etal. 2017). It lives mostly in
dry steppes of short or tall grass, on open hillside, among
rocky outcrops in steppes, or in open dry meadows.
The distribution of Meriones persicus ranges from the
Caucasus (including the southeastern foothills of the Lesser
Caucasus and the Talysh Plateau in Azerbaijan) in the
west, through northeastern Iraq and Iran to Turkmenistan,
Afghanistan (Habibi 2004) and Pakistan, where it is widely
distributed. The species generally occurs in arid, rocky or
mountainous regions (Kryštufek and Vohralík 2009; Molur
& Sozen 2016).
The habitat of Meriones tristrami is limited to areas with
100mm of rainfall annually; the species needs well-drained
soil although it avoids rocky conditions. It has been found in
Armenia, Azerbaijan, Georgia, Iran, the Islamic Republic of
Iraq, Israel, Jordan, Lebanon, the Syrian Arab Republic and
Turkey. Its habitat is mainly shrubland and desert (Kryštufek
and Vohralík 2009; Sozen etal. 2016).
Meriones gr. dahli-libycus
Material: five isolated teeth. Unit II: one isolated tooth;
one right upper M1. Unit V: four isolated teeth; one right
lower m1 and three left upper M1.
Description and discussion: in our sample, we also
identified upper first molars with two roots, which are char-
acteristic of Meriones dahli and Meriones libycus. We pro-
visionally attribute our specimens to this group (Fig.3.12)
in the light of the number of roots and the morphology of
M1, pending a revision of the Middle Eastern species of
the genus.
Habitat and distribution: Meriones dahli is known in
the border area of Armenia and Turkey, and possibly Azer-
baijan. It has been found only in local desert habitats (Kefeli-
oglu etal. 2008).
Meriones libycus has a wide global range, occurring
in North Africa (from Western Sahara and Mauritania to
Egypt) and in Asia (from the Eastern Arabian Peninsula to
China). M. libycus occupies semi-desert and desert habitats;
it is most abundant in unflooded river plains and is often
found close to wadis and dayas (Granjon 2016).
Meriones sp.
Material: nine isolated teeth. Unit II: four isolated teeth;
one right lower m1, two right upper M1 and one left lower
m1. Unit V: five isolated teeth; three right lower m1 and
two left lower m1.
Description and discussion: we identify as Meriones
sp. broken or digested molars without clear discriminating
characters.
Family Muridae Illiger, 1811
Genus Apodemus Kaup, 1829
Seven Apodemus species are currently recognized in the
southern Caucasus: A. hyrcanicus, A. flavicollis, A. with-
erbyi, A. ponticus, A. agrarius, A. mystacinus and A. uralen-
sis, all belonging to the Sylvaemus subgenus (Jangjoo etal.
2011).
Apodemus (Sylvaemus) sp.
Material: 10 isolated teeth. Unit II: seven isolated teeth;
two right lower m1, one left mandible with an m1, m2 and
m3, one left upper M1 and three left lower m1. Unit V: three
isolated teeth; one right upper M1, one right maxilla with an
M1 and M2 and one right lower m1.
Description and discussion: the first lower molars (m1)
from our material show the traits characteristic of the genus
Apodemus (Fig.3.13): the occlusal surface is low, with
six main cusps and a small anterior mesial tubercle; the
anterolabial and posterolabial cusps of m1 converge in an
X-shape; the posterior cusp of m1 is low, rounded and well
developed, with two or three secondary cusps in the labial
part. Identifications to the species level are difficult due to
a lack of documented and genetically typed reference col-
lections. Moreover, comparative morphological data avail-
able in the literature (López-García 2008; Siahsarvie and
Darvish 2008; Kryštufek and Vohralík 2009; Bogicevic etal.
2011; Darvish etal. 2015; Amori etal. 2016; Knitlová and
Horáček 2017) are sometimes unclear and/or display high
variability, and some species are not documented.
Habitat and distribution: the genus Apodemus has a
large distribution range extending from Great Britain across
much of continental Europe to the Urals. It also extends east
through Turkey to western Armenia, the Zagros Mountains
of Iran and south to Syria, Lebanon and Israel. It inhabits a
variety of woodland and bushy habitats (Amori etal. 2016).
Family Dipodidae Fischer, 1817
In the Caucasus, three species are currently represented:
Allactaga williamsi, Allactaga major and Allactaga elater.
Genus Allactaga Cuvier, 1837
Allactaga cf. williamsi
Material: four isolated teeth. Unit II: two isolated teeth;
one right lower m1 and one left upper M1. Unit V: two
isolated teeth; one right upper M3 and one left lower m2.
Discussion: this rodent group is poorly known in the
Middle East (Shenbrot 2009; Naderi etal. 2011; Dianat etal.
2013), and sometimes there is a size overlap between the
species. In our sample, we only found six remains attributed
to Allactaga. The remains are not damaged and were com-
pared with specimens from the reference collection of the
Natural History Museum of London and with data from the
literature. We have an M1 with four roots, a lower m1 with
two roots and an M3 with three roots. All of them share the
same morphological features as Allactaga williamsi (lower
molars narrower than upper ones, third molars less reduced,
first lower molar with a small fold in front and two folds on
either side). In comparison, Allactaga major and Allactaga
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elater have an M1 with only three or two roots (Kryštufek
and Vohralík 2009; Markova 1982). We thus tentatively
attribute the remains from Azokh 1 to Allactaga cf. williamsi
(Fig.3.14), awaiting further studies on the molar morphol-
ogy among the species of this genus.
Habitat and distribution: Allactaga williamsi is distrib-
uted in Anatolia (Turkey), the Caucasus (Armenia, Azerbai-
jan, Iran and Turkey) and northwestern Iran, with an isolated
population in central Afghanistan. As regards its habitat,
this species occurs in steppes with sparse vegetation (Eken
etal. 2016).
Allactaga sp.
Material: two isolated teeth. Unit V: two isolated teeth;
two lower m3.
Description and discussion: we classified as Allactaga
sp. broken teeth whose roots could not be counted or whose
molar morphology could not be seen properly.
Taphonomic study
For the taphonomic study, we focused on a subsample
composed of 380 incisors and molars (100 incisors and 90
molars for each unit). We considered that it was necessary
to identify the origin of the accumulation and the potential
influence of this on any interpretation before a palaeoeco-
logical study could be undertaken.
A considerable percentage of the molars and incisors
show evidence of digestion: 26.83% in Unit V and 22.22%
in Unit II (Fig.3, Table6). According to Andrews (1990)
and Fernández-Jalvo etal. (2016a), the intensities of diges-
tion are mostly light (14.53% in Unit V and 17.50% in Unit
II) or moderate (5.77% in Unit V and 8.37% in Unit II) in
both units (Table6), suggesting that the remains were prob-
ably accumulated by a category 1 predator such as the barn
owl (Andrews 1990; Fernández-Jalvo etal. 2016a). Such
owls have opportunistic hunting habits and are sedentary, so
their prey spectrum is assumed to be a good representation
of the ecosystem in which they live, taking into account their
nocturnal habits and their size.
These results are in agreement with previous tapho-
nomic work published by Andrews etal. (2016), where the
molars and incisors showed similar levels of digestion. The
digestion levels are mainly light or moderate, indicating that
the sample was probably accumulated by Tyto alba, which
produces low degrees of digestion except at its nest site
(Andrews etal. 2016).
Note that Parfitt (2016) found Rattus sp. remains in Unit
V (Table7), which may raise some taphonomic issues. How-
ever, we were unable to verify the identification of these
remains, so we cannot draw any conclusion on this point
for now.
Palaeoecological reconstruction ofAzokh 1 Cave
Sequence 2
The small-mammal sequence of Azokh 1 Cave is quite
diverse, with at least 13 taxa identified in this work (Table3).
Compared to previous studies (Parfitt 2016), the new taxa
we identified include (Table7) Mesocricetus brandti, Arvi-
cola ex. gr. persicus, Ellobius cf. lutescens, Meriones gr.
persicus-tristrami, Meriones gr. dahli-libycus and Allactaga
cf. williamsi. Conversely, some species identified by Parfitt
were not found in our samples: Allocricetus sp., Rattus sp.,
Mus cf. macedonicus and Dryomys nitedula.
There is no significant turnover between the studied units,
which contain similar rodent assemblages (Table3). The
most abundant taxon in all the units is Microtus gr. arvalis-
socialis (77 individuals in Unit V, seven in Units III–IV and
75 in Unit II), followed by Ellobius cf. lutescens (seven indi-
viduals in Unit V and 12 individuals in Unit II) and Microtus
(Terricola) spp. (seven individuals in Unit V and four indi-
viduals in Unit II). Most of the species identified at Azokh
1 Cave are still present in the area today, but there are some
exceptions, such as Chionomys gud, Ellobius cf. lutescens
and Clethrionomys glareolus.
Units III–IV do not yield enough material to draw pal-
aeoecological inferences (MNI < 30). For Units V and II, the
bioclimatic model shows similar results (Table8). The BM
based on the small mammals from Azokh 1 suggests mean
annual temperatures and precipitation similar to the present
(Table8), it being a bit colder and drier in Unit II than in
Unit V. The climate seems to be relatively warm-temperate,
with continental conditions in both units.
Table 6 Representation (N) and percentages of digestion (%) on isolated incisors and isolated molars from Azokh 1 Cave, Unit V and Unit II
Unit V Unit II
Elements Total No digestion Total
digested
Light diges-
tion
Moderate
digestion
Total No digestion Total
digested
Light diges-
tion
Moderate
digestion
N N N %N%N%N N N %N%N%
Isolated incisor 100 73 27 27 21 21 6 6 100 75 25 25 22 70 3 7
Isolated molars 90 66 24 26.67 19 11.11 5 5.56 90 72 18 20 9 10 9 10
Total 190 139 51 26.83 40 21.05 11 5.78 190 147 43 22.63 31 16.31 12 6.31
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The habitat weighting method (Fig.5) shows an environ-
ment mainly composed of steppes and shrublands, notably
indicated by the presence of Meriones spp. and Mesocri-
cetus brandti. In Unit V, grasslands (Allactaga cf. wil-
liamsi), desert (Cricetulus migratorius) and rocky habitats
(Chionomys gud, C. nivalis) are also well represented. Unit
II presents a higher representation of wetland habitat than
Unit V, due to the presence of Arvicola ex. gr. persicus. The
proportion of forest in Unit II is also greater than in Unit V,
as indicated by the presence of Clethrionomys glareolus and
Apodemus (Sylvaemus) sp.
The current vegetation map of Azokh shows the pres-
ence of a broad belt of semi-xerophilous woodland in the
lowlands (Manuk 2010). Further to the east are belts of sage-
brush steppe and sagebrush desert, and both would have
formed part of the habitat ranges of larger mammals and
birds of prey (Andrews etal. 2016).
In summary, the BM does not reveal significant palaeo-
climatic differences between Unit V, Unit II and the current
values. The HW shows some differences, especially in the
proportions of forest and wetlands. However, the environ-
mental data from both Units V and II are consistent with
steppe environments with shrubland, rocky areas and arid
conditions.
Comparisons withother palaeoenvironmental
proxies fromAzokh 1 Cave
The results we obtained with the rodent assemblages were
compared with previous studies carried out at Azokh 1 Cave,
where palaeoenvironmental proxies were estimated using
large mammals, small vertebrates and archaeobotanical data
Table 7 Comparison between
the new studies carried out
in Azokh 1 Cave and Parfitt
(2016), X: indicates presence in
the sample
This study Parfitt (2016)
Taxa Unit V Unit III-IV Unit II Unit V Unit III Unit II/III Unit II Unit I
Cricetulus migratorius X X X X X X
Mesocricetus brandti X X
Mesocricetus sp. X X X X
Allocricetus sp. X X
Clethrionomys glareolus X X X
Arvicola ex. gr. persicus X
Chionomys gud X X X
Chionomys nivalis X X X X
Microtus gr. arvalis-socialis X X X X X X X X
Microtus (Terricola) spp. X X X X X X X
Ellobius sp. X X X X X
Ellobius cf. lutescens X X X
Meriones spp. X X X X X X X
Meriones gr. dahli-libycus X X
Meriones gr. persicus-tristrami X X
Apodemus sp. X X X X X X
Rattus sp. X
Mus cf. macedonicus X X X
Dryomys nitedula X
Allactaga sp. X x
Allactaga cf. williamsi X
Table 8 Temperature and precipitation values for Azokh 1 Unit V
and Unit II, obtained by the bioclimatic model. MAT mean annual
temperature; MTC mean temperature of the coldest month; MTW
mean temperature of the warmest month; MAP mean annual precipi-
tation; Max maximum of values obtained; Min minimum of values
obtained. The current values were obtained from https:// en. clima te-
data. org/
Unit V Unit II Current values
MAT 11.42 10.86 10.1
MAX 18.08 17.5
MIN 4.76 4.24
MTW 22.5 22.44 22.9
MAX 27.81 27.71
MIN 17.2 17.16
MTC 1.43 0.11 − 3
MAX 16.65 15.23
MIN − 13.78 − 15.02
MAP 528.13 500.01 668
MAX 1144 1112.61
MIN − 87.74 − 112.6
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(Table9). We paid special attention to the study undertaken
by Parfitt (2016), who analysed most of the small mammals
collected between 2002 and 2009 in Sediment Sequence 2
from Unit V to Unit I, as well as to the palaeoecological
interpretations proposed by Andrews etal. (2016).
Most of the previous studies of the small mammals
from Azokh have been based on Unit V, which yielded
the most abundant material. Parfitt’s results showed subtle
differences in faunal composition throughout the Middle
(Unit V) to Late Pleistocene (Units III and II) sequence of
Azokh, indicating changes in aridity and temperature, com-
bined with fluctuations in woodland cover and the prox-
imity of trees to the site (Parfitt 2016). Our study made it
possible to add one new species (Arvicola ex. gr. persicus)
to the faunal list established by Parfitt (2016) and to specify
the identifications for Ellobius cf. lutescens, Meriones gr.
persicus-tristrami, Meriones gr. dahli-libycus and Allac-
taga cf. williamsi, which were previously limited to the
genus level. Moreover, our study allowed us to provide
more consistent data for Unit II, which was the least repre-
sentative unit among the samples in Parfitt (2016).
According to Marin-Monfort etal. (2016), the large mam-
mals were accumulated both by carnivores and humans.
Taxonomic identifications have been performed by Van der
Made etal. (2016), with at least 29 species represented in
the sequence (Table9). Some of the species, such as Cervus
elaphus, are present in the whole sequence. Ursus spelaeus
was identified in all the Pleistocene units, and a taphonomic
study revealed that cave bear remains were relatively com-
plete, with some bones in anatomical connection, suggest-
ing that the bears living in the cave were using it as a den
(Marin-Monfort etal. 2016). The remains of other mammals
are, in most units, extremely fragmented, and mainly repre-
sented by teeth, horn/antler and foot bones. According to
Andrews etal. (2016), using the habitat weighting method,
Units V–II have the highest index value for deciduous wood-
land and also for Mediterranean evergreen woodland, as
indicated by Stephanorhinus kirchbergensis, Sus scrofa and
Dama sp. Unit I differs from the others by its more steppic,
arid signal.
The herpetofauna of Azokh 1 is composed exclusively of
extant genera and species (Table9), most of them belonging
to thermophilous and xerophilous forms (e.g. Pelobates syri-
acus, Agamidae, Pseudopus apodus, Ophisops elegans, Eryx
jaculus, Elaphe sauromates, among others). The anuran
Pseudepidalea viridis has a wide ecological tolerance. Most
taxa frequent open wooded or bushy areas, such as Pseu-
dopus apodus or Ophisops elegans (Blain 2016). Accord-
ing to Blain (2016), the faunas from Units II to I reflect a
drier period, including a representative of the Vipera berus
complex (probably V. ursinii) and the small colubrine Cor-
onella austriaca. In Unit V, the environment seems to be
more consistent with a meadow steppe. The results obtained
with amphibians and squamates are consistent with those
obtained with rodents and indicate an environment mainly
composed of desert and steppes, especially in Unit V.
The bat fauna from Azokh 1 is composed of extant genera
(Table9), and the main habitats that it reflects are mountains
steppes, followed by mountain grassland. Unit V is domi-
nated by species with Mediterranean or humid affinities,
indicating woodland conditions. The bat assemblage indi-
cates colder conditions for Unit II, with a higher proportion
of steppe environment. We can thus see some discrepan-
cies with respect to the results obtained from rodents, which
indicate more steppic-arid conditions in Unit V and more
woody-temperate conditions in Unit II.
Archaeobotanical data (Table9) obtained from charcoal
studies (Allué 2016) of Units V and II show that wood could
have been carried into the cave by humans. However, no
hearth or other intentionally constructed feature has been
identified in the site at the rear of the cave where these fossil
assemblages were deposited during the Pleistocene. Wood
could have also been carried in by animals or brought in
by natural surface fires (Andrews etal. 2016). The main
Fig. 5 Results obtained by the
habitat weighting method for
Azokh 1 Cave (Unit V and Unit
II)
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Archaeological and Anthropological Sciences (2022) 14: 96
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taxa identified from the charcoal remains are Prunus, Acer
and Quercus, a combination of large woodland trees and
small trees and shrubs. The results of the phytolith analy-
sis of coprolites (Scott etal. 2016) for Unit II can also be
included, indicating a temperate steppe mosaic with grassy
conditions, but the density of woody components cannot
be determined. This interpretation is consistent with our
rodent analysis, which shows a wide spectrum of species
suggesting an open or semi-open landscape formed mainly
by woody trees and shrubs.
In summary, the large mammals and the charcoal data
reflect woodland environments, whereas the small ver-
tebrates and coprolites mainly depict arid environments,
mostly steppes and shrubland. These differences were
at least partly explained by Andrews etal. (2016), who
argued that they indicate different taphonomic trajectories,
or origins of the accumulations. Both the large mammals
and wood could have been selected in specific habitats and
introduced to the site by humans (Allué 2016). By con-
trast, the small vertebrates and coprolites were accumu-
lated by non-human predators, which could have hunted
over a wider diversity of habitats, especially open areas
(Andrews etal. 2016).
Azokh 1 Cave andtheSouthern Caucasus context
Several other Pleistocene sites have provided small-mammal
studies in the Middle East: Qesem Cave (Smith etal. 2015;
Maul etal. 2016), Hummal (Maul etal. 2015), Aghitu-3
Cave (Kandel etal. 2017; Nishiaki and Akazawa 2018;
Frahm 2019), Nesher Ramla (Weissbrod and Zaidner 2014),
Amud Cave (Belmaker and Hovers 2011), Misliya Cave
(Weissbrod and Weinstein-Evron 2020), Dzudzuana Cave
(Belmaker etal. 2016), Kaldar Cave (Rey-Rodríguez etal.
2020), Shanidar Cave (Tilby etal. 2020), Kudaro Caves
(Baryshnikov 2002) and Karain Cave (Demirel etal. 2011).
Most of these studies have highlighted the problems of
identifying rodent species in this region, especially the Mid-
dle Pleistocene species. We focus on Qesem Cave and Mis-
liya Cave, both of them located in Israel (Table10, Fig.6).
We are aware that Israel is quite a long way from our study
area for a comparative analysis; the sites do not belong to
the same climatic zones (Fig.6), and the faunas may be
very different. However, these sites are present well-studied
rodent assemblages that could be chronologically compared
with Azokh 1 Cave.
Qesem Cave is an archaeological site located near Tel
Aviv (32° 06 36 N, 34° 58 48 E) at 90m a.s.l. It can
be compared with Azokh Unit V (Table9) in terms of its
chronological framework. The palaeoecology of Qesem
Cave was explored using two methods. The first was the
nearest living relative (NRL) approach, and the second was
the coexistence approach, both of which are based on the
assumption that at least since the Neogene taxa have had
climatic requirements similar to their NRLs, allowing infer-
ences to be drawn on past climatic conditions. The modern
climatic requirements were taken from the IUCN (Maul
etal. 2015). The results show a climate of Mediterranean
type during the accumulation of the two microvertebrate
concentrations. A large proportion of the annual precipita-
tion occurred in winter, whereas summers were dry. The
temperatures were lower than today during the deposition of
Concentration 2, with especially harsh winters, and winter
precipitation was lower, resulting in lower precipitation sea-
sonality. The landscape was a mosaic of open habitats with
sparse vegetation, shrubland, Mediterranean forest, rocky
areas and riverbanks (Maul etal. 2015; Smith etal. 2015).
Both concentrations were accumulated by the barn owl Tyto
alba (Smith etal. 2015).
Misliya Cave is located on the western slope of Mount
Carmel (32° 44 29 N, 34° 58 21 E), facing the coastal
plain and situated at an elevation of about 90m a.s.l. It is
chronologically comparable with Azokh Unit II (Table10).
The environmental conditions of Misliya Cave are charac-
terized by a heterogeneous landscape, or habitat mosaics
(Weissbrod and Weinstein-Evron 2020).
Misliya Cave is characterized by a combination of Afri-
can rodent taxa, such as Mastomys and Arvicanthis, and
Euro-Siberian taxa, such as Ellobius. This could suggest
that the Transcaucasia region (like the Central Mountains
in Israel and the Caucasus mountains), the Zagros and Tau-
rus Mountains may have acted more as a barrier than as a
migration corridor for the Azokh rodents. This hypothesis,
combined with the greater Mediterranean influence on the
Israeli sites, could explain the differences in the palaeocli-
matic signal observed between Qesem and Misliya Cave on
the one hand, and Azokh 1 Cave on the other.
Figure6 shows the location of Qesem Cave and Misliya
Cave in relation to the different main climates in present-day
Israel according to Köppen (1936, modified by Beck etal.
2018). The climate ranges mainly from temperate (Misliya)
to an arid limit (Qesem), with a hot summer and steppe envi-
ronment. For Azokh 1 the climate is classified as “snowy”,
with dry winters and warm summers.
The rodent assemblages of Azokh 1 show a strong Asiatic
influence, whereas the large mammals (Van der Made 2016)
are dominated by Western Eurasian species, also with Afri-
can, Indian and central Asian elements. The rodent fauna did
not record any African species, whereas some were found
at the Israeli sites.
The identified rodent species present several biogeo-
graphic affinities but are mainly from Europe and Asia.
Misliya also recorded the presence of African rodent taxa,
such as Mastomys and Arvicanthis (Maul etal. 2016;
Weissbrod and Weinstein-Evron 2020). Such species were
not found at Azokh, indicating that the Caucasus, Zagros
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Archaeological and Anthropological Sciences (2022) 14: 96
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96 Page 20 of 25
and Taurus Mountains probably acted as a barrier for small
mammals coming from Africa. Moreover, most of the spe-
cies found at Azokh display an Asiatic origin, suggesting
the possible importance of the Black and Caspian seas as
barriers to small mammals coming from Europe, as was
previously indicated by Yanina (2014).
Conclusions
We have identified 434 rodent remains, corresponding
to a minimum number of 237 individuals. The rodent
assemblages from Azokh 1 Cave are composed of at
Fig. 6 aLocation of Qesem Cave, Misliya Cave and Azokh 1. b Climatic map of the studied region, obtained from Zittis (2015)
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Archaeological and Anthropological Sciences (2022) 14: 96
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Page 21 of 25 96
Table 9 Species and elemental interpretations carried out previously in Azokh 1 Cave with different environmental proxies
Large mammals (Van der made etal.2016) Small vertebrates other than rodents (Blain2016; Sevilla2016) Archeobotanical data (Allué2016; Scott etal.2016)
Species Environmental conditions Species Environmental conditions Species Environmental conditions
Unit V Carnivores: Canis aureus,
Crocuta crocuta, Lynx sp.
Felis chaus, Panthera pardus,
Ursus spelaeus
Interglaciar species. Central
Asia aspect
Bats: Rhinolophus ferrumequi-
num, Rhinolophus mehelyi,
Rhinolophus euryale, Myotis
blythii, Myotis nattereri/
schaubi, Myotis mystacinus,
Plecotus auritus/macrobul-
laris, Barbastella barbastel-
lus, Barbastella laucomelas,
Pipistrellus nathusii, Pipist-
relus pipistrelus, Miniopterus
schreibersii
Bats indicate a predominance
of Mediterranen species.
Herpetofauna indicates an
environment consistent with
a meadow-steppe, by the
presence of species such as P.
syriacus and small vipers
Charcoal remains indicate the
presence of three taxa: Pru-
nus, Maloideae and deciduous
Quercus sp. Phytoliths were
studied from a coprolite
The data obtained with charcoal
indicate mild and humid envi-
ronmental conditions. However,
phytoliths indicate a temperate
steppe mosaic, with grassy
conditions
Artiodactyla: Cervus elaphus,
Capra aegagrus Amphibians: Pseudepidalea
viridis sensu lato, Ranidae/
Hylidae indet. and Pelobates
cf. syriacus
Perissodactyla: Stephanorhinus
hemitoechus, Stephanorhinus
kirchbergensis, Equus hydrun-
tinus, Equus ferus
Squamates: Pseudopus apodus,
Lacerta sp., Eryx jaculus,
cf. Coronella austriaca, cf.
Elaphe sp., cf.“Coluber” sp.
and “Colubrinae” indet
Unit II Carnivores: Panthera pardus,
Ursus spelaeus and Ursus sp.
and Vulpes vulpes, Canis lupus
Interglaciar temperate environ-
ment
Bats: Rhinolophus ferrumequi-
num, Myotis blythii, Myotis
dasycneme, Pipistrelus pipist-
relus, Miniopterus schreibersii
Bats indicate colder condi-
tion in this unit, because of
the absence of Rhinolophus
mehelyi. Accoding to the
herpetofaunal remains, this
assemblage would represent a
drier period, with the presence
of a representative of the V.
berus complex (probably V.
ursinii) and the small colu-
brine Coronella austriaca
Charcoal remains: Prunus,
Acer, deciduous Quercus sp.,
Maloideae, Lonicera, Paliu-
rus/Ziziphus, Celtis/Zelkova,
Euonymus, and Ulmaceae
This assemblage composition
indicates mild and humid envi-
ronmental conditions
Artiodactyla: Cervus elaphus,
Capra aegagrus, Saiga
tatarica, Dama sp. and Sus
scrofa
Amphibians: Pseudepidalea
viridis sensu lato, Pelobates
cf. syriacus
Perissodactyla: Stephanorhinus
kirchbergensis Squamates: Pseudopus apodus,
Lacerta sp., Ophisops elegans,
Eryx jaculus, cf. Coronella
austriaca, cf. Elaphe sp., cf.
Coluber” sp., Vipera spp.
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Archaeological and Anthropological Sciences (2022) 14: 96
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96 Page 22 of 25
least 13 taxa: seven arvicoline (Clethrionomys glareo-
lus, Microtus gr. arvalis-socialis, Microtus (Terricola)
spp., Arvicola ex. gr. persicus, Chionomys nivalis,
Chionomys gud and Ellobius cf. lutescens), two crice-
tine (Cricetulus migratorius and Mesocricetus brandti),
two gerbilline (Meriones gr. persicus-tristrami, Meri-
ones gr. dahli-libycus), one dipodid (Allactaga cf. wil-
liamsi) and one murine species (Apodemus spp.).
The palaeoclimatic parameters obtained with the biocli-
matic model suggest mean annual temperatures and pre-
cipitation similar to today, although the climate seems to
be relatively warm-temperate in both units.
The palaeoenvironmental reconstruction, based on the habi-
tat weighting method, shows an environment mainly com-
posed of shrubland and steppe, with patches of deciduous
forests and desert, similar to that currently found in the area.
Whereas large-mammal and charcoal studies indicate a
woodland environment, small vertebrates and phytoliths
from coprolites mainly reflect arid environments, such
as steppes and desert. These differences can be partly
explained by the origin of the accumulations.
A comparison between Azokh Cave on one hand and
Qesem Cave and Misliya Cave (Israel) on the other indi-
cates that there is no exact correspondence between the
rodent faunas, both because the sites belong to differ-
ent climatic regions and because the Israeli sites record
African influences, which were not observed in Azokh 1.
Acknowledgements We would like to thank Roberto Portela Miguez,
Senior Curator in Charge of Mammals, for his help with the refer-
ence collection in the Natural History Museum of London; Lawrence
Heaney, Adam Ferguson and Lauren Smith of the Chicago Field
Museum; and Marisa Surovy, Judith Galkin and Carl Mehling of the
American Museum of Natural History of New York. We would like
to thank Rupert Glasgow for reviewing the English language of the
manuscript. We also want to thank the Guest Editor Angel Blanco, as
well as Dr. Alexey S. Tesakov and the anonymous reviewer for their
comments and suggestions, which greatly improved the final version
of the manuscript.
Funding Open access funding provided by Universitat Rovira i Virgili.
I. Rey-Rodriguez is the beneficiary of a Margarita Salas postdoctoral
scholarship (2021URVMS03) at Universitat Rovira i Virgili funded
by the European Union – NextGenerationEU. J.M. López-García was
supported by a Ramón y Cajal contract (RYC-2016–19386) with finan-
cial sponsorship from the Spanish Ministry of Science, Innovation and
Universities.
Declarations
Conflict of interest The authors declare no competing interests.
Open Access This article is licensed under a Creative Commons Attri-
bution 4.0 International License, which permits use, sharing, adapta-
tion, distribution and reproduction in any medium or format, as long
as you give appropriate credit to the original author(s) and the source,
provide a link to the Creative Commons licence, and indicate if changes
were made. The images or other third party material in this article are
included in the article's Creative Commons licence, unless indicated
otherwise in a credit line to the material. If material is not included in
the article's Creative Commons licence and your intended use is not
permitted by statutory regulation or exceeds the permitted use, you will
need to obtain permission directly from the copyright holder. To view a
copy of this licence, visithttp:// creat iveco mmons. org/ licen ses/ by/4. 0/.
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Article
This study addresses the roles of biotic agents in site formation in the B1 strata of Block 2 at Dmanisi, Georgia, using theoretical and analogous frameworks for the interpretation of spatial behaviors of carnivores and hominins. For this study, stone material, faunal remains, and coprolites are analyzed to determine if any spatially distinct behaviors can be identified, located, and attributed to either hominins or carnivores. Faunal, stone, and coprolite assemblages are compared with each other, and lithic, taxonomic, and taphonomic subassemblages are compared with the overall distribution of their parent material. The spatial and taphonomic signatures suggest that hominin activity was only a small part of the contributing factors to site formation, whereas carnivores played a major role in the accumulation of bone.
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In this study, we provide the first taphonomic and taxonomic descriptions of the micromammals from Misliya Cave, where recently a Homo sapiens hemimaxilla has been reported. This finding significantly extends the time frame for the out-of-Africa presence of anatomically modern humans. It also provides an opportunity to reassess variation in early modern human population responses to climate change in the Levantine sequence. Information on species ranking and diversity estimations (Shannon functions) is obtained from quantitative data across 31 Levantine assemblages and investigated in a broad comparative frame using multivariate analyses. Recent models of human-climate interactions in the late Early–Middle Paleolithic of the southern Levant have drawn heavily on on-site associations of human fossils with remains of micromammals. However, there has been little, if any, attempt to examine the long-term picture of how paleocommunities of micromammals responded qualitatively and quantitatively to climatic oscillations of the region by altering their compositional complexity. Consequently, our understanding is vastly limited in regard to the paleoecosystem functions that linked past precipitation shifts to changes in primary producers and consumers or as to the background climatic conditions that allowed for the development of highly nonanalog ancient communities in the region. Although previous studies argued for a correspondence between alternations in H. sapiens and Neanderthal occupations of the Levant and faunal shifts in key biostratigraphic indicator taxa (such as Euro-Siberian Ellobius versus Saharo-Arabian Mastomys and Arvicanthis), our data indicate the likelihood that early H. sapiens populations (Misliya and Qafzeh hominins) persisted through high amplitudes of paleoecological and climatic oscillations. It is unlikely, given these results, that climate functioned as a significant filter of early modern human persistence and genetic interactions with Neanderthals in the Levant.
Chapter
The Caucasus is an important intercontinental passageway for fauna and hominin dispersal from Africa to Eurasia. Numerous Pleistocene sites emphasise the importance of this region for the study of human evolution and hominin ‘Out of Africa’ dispersals. The Azokh 1 site in the Southern Caucasus provides a stratigraphic sequence, the renewed excavations of which have shown the presence of well-contextualised lithic and faunal assemblages dated between 300 and 100 ka associated with hominin remains (Homo heidelbergensis and Homo neanderthalensis) also found in the site.