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Citation: Venditti, F.; McCartin, M.J.;
Ostermann, M.-L.; Conard, N.J.; Wolf,
S. Foxes in Retrospect—Unraveling
Human-Fox Relationships through
Fox Tooth Ornaments in the Swabian
Jura. Quaternary 2023,6, 50. https://
doi.org/10.3390/quat6030050
Academic Editor: Juan Manuel López
García
Received: 16 August 2023
Revised: 2 September 2023
Accepted: 6 September 2023
Published: 21 September 2023
Copyright: © 2023 by the authors.
Licensee MDPI, Basel, Switzerland.
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4.0/).
quaternary
Article
Foxes in Retrospect—Unraveling Human-Fox Relationships
through Fox Tooth Ornaments in the Swabian Jura
Flavia Venditti 1,* , Madison J. McCartin 2, Melanie-Larisa Ostermann 2, Nicholas J. Conard 1,3
and Sibylle Wolf 1,3,*
1Department of Early Prehistory and Quaternary Ecology, University of Tübingen, Schloß Hohentübingen,
Burgsteige 11, 72070 Tübingen, Germany; nicholas.conard@uni-tuebingen.de
2
Institute for Archaeological Sciences, University of Tübingen, Hölderlinstrasse 12, 72074 Tübingen, Germany;
mjmccartin@ucdavis.edu (M.J.M.); laraostermann20@gmail.com (M.-L.O.)
3Senckenberg Centre for Human Evolution and Palaeoenvironment at the University of Tübingen,
Hölderlinstrasse 12, 72074 Tübingen, Germany
*Correspondence: flavia.venditti@uni-tuebingen.de (F.V.); sibylle.wolf@ifu.uni-tuebingen.de (S.W.)
Abstract:
Personal ornaments play an important role in our understanding of human cultural and
behavioral change during the Upper Paleolithic, providing insights into intangible aspects of human
cultural behavior. Some ornament forms are better studied than others, and fox tooth ornaments,
despite their frequent occurrence and broad spatiotemporal span, are relatively under-addressed.
Here we present the first comprehensive study of 40 perforated fox teeth recovered from four cave
sites in southwestern Germany. This region’s rich record of symbolic representations, as well as
evidence of long-standing human–fox relationships, make the Swabian Jura an ideal case study
for investigations of fox tooth ornaments. By applying a holistic approach, including geometric
morphometrics and traceology coupled with experimental archaeology, we show that fox teeth were
mostly perforated by bifacial scraping and grooving and were worn as ornaments. We discuss the role
of foxes within human socio-symbolic and paleoenvironmental systems during the Upper Paleolithic
of the Swabian Jura, and we contextualize our results within the broader context of sites across
Europe during the Upper Paleolithic. The data we provide are in line with general trends observed
across the continent and offer insight into the role of foxes during the Upper Paleolithic, especially
regarding human subsistence, cultural expression, and ornament production.
Keywords:
personal ornamentation; traceology; geometric morphometrics; Upper Paleolithic; Swabian
Jura; foxes (Vulpes vulpes and Vulpes lagopus)
1. Introduction
In the Swabian Jura (southwestern Germany), Upper Paleolithic symbolic culture is
characterized by ivory figurines, musical instruments, and personal ornamentation [
1
–
9
].
Although small, ornaments are particularly informative, as they are numerous, well-
preserved, diverse in form, and occur throughout the entirety of the Upper Paleolithic.
Their hypothesized role as social and symbolic objects also lends insight onto intangible
aspects of human cultural behavior, such as personal identity, group affiliation, language,
movement, trade, and social status [8,10–17].
During the Upper Paleolithic, ornaments were produced from a variety of organic
and inorganic raw materials, including ivory, tooth, bone, shell, fossil, and stone. Those
made from ivory, shells, and cervid teeth are well studied [
8
,
10
,
11
,
18
–
30
], while fox tooth
ornaments, primarily made from red (Vulpes vulpes) and arctic fox (Vulpes lagopus) canines,
are comparatively under-addressed.
Fox tooth ornaments first appear during the Châtelperronian and persist throughout
the entire Upper Paleolithic at sites spanning the Iberian Peninsula to the East European
Plain (see Supplementary Materials) [
13
,
17
,
31
]. Most assemblages are small, usually less
Quaternary 2023,6, 50. https://doi.org/10.3390/quat6030050 https://www.mdpi.com/journal/quaternary
Quaternary 2023,6, 50 2 of 36
than twenty specimens (Table S1), although exceptional cases include sites like Pavlov
I (Czechia) and Sunghir (Russia), each represented by over 200 fox tooth ornaments in
association with Gravettian burials [10,32–34].
Despite their abundance and wide geographic span, detailed information on fox
tooth ornaments is limited (e.g., on species and tooth selection, metric data, production
methods, use-wear, taphonomy). This is especially disappointing, given their potential
to illuminate not only the socio-symbolic aspects of ornamentation, but also the broader
relationship between humans, foxes, and their shared environment. Unlike other ornament
forms, animal teeth come from a living organism and are, therefore, directly related to local
ecology and human subsistence practices. Fox tooth ornaments can thus be linked to the
broader role of foxes on the landscape (e.g., paleoecology, human–animal relationships),
in human diets (e.g., hunting methods, nutrition), and as a raw material (e.g., fur, bone
tools, ornamentation).
This study presents an assemblage of Upper Paleolithic fox tooth ornaments
(n = 40) from four sites in the Swabian Jura: Hohle Fels (n = 12), Geißenklösterle
(n = 12), Brillenhöhle (n = 3), and Hohlenstein-Stadel (n = 13) (Supplementary Materi-
als and Figures 1and 2). This region’s rich record of symbolic representation, as well as
evidence of long-standing human–fox relationships [
35
–
39
], make the Swabian Jura the
ideal location to explore the production and use of these ornaments and their relevance for
broader human–fox relationships in the Upper Paleolithic.
In this study, we explore fox tooth ornaments from a diachronic and synchronic per-
spective by analyzing ornaments from the entire Upper Paleolithic sequence (Aurignacian
to Magdalenian) and by considering intra- and inter-ornament variability and the influence
of culture versus technological preferences. In particular, we aim to:
1.
investigate diachronic changes in technology, production, and function and how these
changes relate to the methods of the perforation and mode of use among the three
cultures (Aurignacian, Gravettian, and Magdalenian) and within the four analyzed
sites in the Swabian Jura,
2.
explore the potential for species preferences between red fox and arctic fox within
the analyzed sites and discuss the results in the framework of fox ecology and
behavior, and
3.
contextualize the results from the Swabian Jura within a broader context, drawing
from an extensive literature review of fox tooth ornaments from sites across Europe
during the Upper Paleolithic.
To do so, we apply three analytical methods to obtain a more holistic view of fox
exploitation and ornament production, use, and discard. Geometric morphometrics facil-
itates the reliable differentiation of red versus arctic fox in isolated teeth, allowing us to
investigate species selection and preference, fox paleoecology, and the interaction between
humans and foxes in the Swabian Jura. This is complemented by traceological analysis cou-
pled with experimental archaeology, in which microscopic observations of the perforations
are used to identify manufacturing techniques and use-wear traces.
Foxes in the Swabian Jura
During the Late Pleistocene, climatic and environmental conditions influenced the dis-
tribution and composition of fox species on the landscape and their subsequent availability
to hunter-gatherer populations [
40
,
41
]. At the start of the Upper Paleolithic, both red and
arctic foxes were sympatric in Central Europe. Subsequent cooling associated with the
Last Glacial Maximum forced red foxes to retreat into glacial refugia, such as the Iberian
Peninsula [
42
,
43
]. Sympatric populations appeared again in Central Europe during the
Late Glacial, but Allerød warming later pushed arctic foxes northward. A final period of
sympatry is recorded during the Younger Dryas, followed by the northern retreat of arctic
foxes in response to Holocene warming, after which both species attained their modern
distributions [42].
Quaternary 2023,6, 50 3 of 36
Quaternary2023,6,xFORPEERREVIEW3of38
Figure1.Perforatedfoxteethanalyzedinthisstudyandtheirgeographiclocation.Itemsfromleft
toright.(a)Upperrow:ID461,ID631.1,ID1278.1,ID1295;middlerow:ID1517.1,ID1137,ID
1228.2,ID964.1;lowerrow:ID1227.1,ID1742,ID2099.1,ID1713.(b)Upperrow:ID1B,ID1A,ID
391,ID89,ID369;middlerow:ID286,ID38F,ID658.2,ID226,ID918;lowerrow:ID487,ID1511.
(c)Upperrow:ID1.0380,ID2.0380,ID3.0380,ID4.0380,ID.5.0380,ID141-1,ID14-1;lowerrow:ID
33-1,ID36-1,ID167-1,ID178-1,ID9000-1,ID9001-1.(d)ID130,ID131,ID132.
Figure 1.
Perforated fox teeth analyzed in this study and their geographic location. Items from left to
right. (
a
) Upper row: ID 461, ID 631.1, ID 1278.1, ID 1295; middle row: ID 1517.1, ID 1137, ID 1228.2,
ID 964.1; lower row: ID 1227.1, ID 1742, ID 2099.1, ID 1713. (
b
) Upper row: ID 1B, ID 1A, ID 391,
ID 89, ID 369; middle row: ID 286, ID 38F, ID 658.2, ID 226, ID 918; lower row: ID 487, ID 1511.
(
c
) Upper row: ID 1.0380, ID 2.0380, ID 3.0380, ID 4.0380, ID. 5.0380, ID 141-1, ID 14-1; lower row:
ID 33-1, ID 36-1, ID 167-1, ID 178-1, ID 9000-1, ID 9001-1. (d) ID 130, ID 131, ID 132.
In the Swabian Jura, both red and arctic foxes are found at numerous archaeological
sites throughout the Paleolithic, generally accounting for 1–5% of NISP. Their abundance
increased over time compared to other carnivores and large herbivores. A similar increase
is recorded in hare, likely due to common acquisition methods, such as traps and snares,
which elevated the catch rate of small game [
44
,
45
]. Juvenile foxes (<1 year) are rare, only ac-
counting for ~2% of fox NISP. Most come from the Middle Paleolithic of Hohlenstein-Stadel
(n = 17) [
46
], reflecting natural deaths from denning animals. The comparative dearth of ju-
veniles from Upper Paleolithic contexts (Vogelherd Aurignacian, n = 1 [
35
]; Geißenklösterle
Gravettian, n = 4 [
39
]; Vogelherd Magdalenian, n = 1 [
35
]; Langmahdalde Magdalenian,
n = 6 [
47
]) may indicate issues with preservation or excavation techniques, that caves were
not used as fox dens, or that humans did not intentionally hunt juveniles [44].
Quaternary 2023,6, 50 4 of 36
Quaternary2023,6,xFORPEERREVIEW4of38
Figure2.MapoftheSwabianJurawiththelocationofimportantPaleolithiccavesitesfromtheAch,
Lone,andLauchertvalleys.FoursitesoutlinedinredprovideUpperPaleolithicfoxtoothpendants
andarethefocusofthisstudy:HohleFels(1)Geißenklösterle(2),Brillenhöhle(3),Hohlenstein-
Stadel(4).(MapprovidedbyChristianSommer/ROCEEH(2019):AmapcollectionofthePaleolithic
oftheSwabianJura(Version1.0.0)(Dataset).(modifiedbyF.V.).
FoxesintheSwabianJura
DuringtheLatePleistocene,climaticandenvironmentalconditionsinfluencedthe
distributionandcompositionoffoxspeciesonthelandscapeandtheirsubsequentavail-
abilitytohunter-gathererpopulations[40,41].AtthestartoftheUpperPaleolithic,both
redandarcticfoxesweresympatricinCentralEurope.Subsequentcoolingassociated
withtheLastGlacialMaximumforcedredfoxestoretreatintoglacialrefugia,suchasthe
IberianPeninsula[42,43].SympatricpopulationsappearedagaininCentralEuropedur-
ingtheLateGlacial,butAllerødwarminglaterpushedarcticfoxesnorthward.Afinal
periodofsympatryisrecordedduringtheYoungerDryas,followedbythenorthernre-
treatofarcticfoxesinresponsetoHolocenewarming,afterwhichbothspeciesattained
theirmoderndistributions[42].
IntheSwabianJura,bothredandarcticfoxesarefoundatnumerousarchaeological
sitesthroughoutthePaleolithic,generallyaccountingfor1–5%ofNISP.Theirabundance
increasedovertimecomparedtoothercarnivoresandlargeherbivores.Asimilarincrease
isrecordedinhare,likelyduetocommonacquisitionmethods,suchastrapsandsnares,
whichelevatedthecatchrateofsmallgame[44,45].Juvenilefoxes(<1year)arerare,only
accountingfor~2%offoxNISP.MostcomefromtheMiddlePaleolithicofHohlenstein-
Stadel(n=17)[46],reflectingnaturaldeathsfromdenninganimals.Thecomparative
dearthofjuvenilesfromUpperPaleolithiccontexts(VogelherdAurignacian,n=1[35];
GeißenklösterleGravettian,n=4[39];VogelherdMagdalenian,n=1[35];Langmahdalde
Magdalenian,n=6[47])mayindicateissueswithpreservationorexcavationtechniques,
thatcaveswerenotusedasfoxdens,orthathumansdidnotintentionallyhuntjuveniles
[44].
EvidenceoffoxbutcheryisrarecomparedtoothertaxaintheSwabianJura[44].
DuringtheMiddlePaleolithic,asinglephalanxfromBocksteinexhibitsacircularcut
markassociatedwithskinning[48].DuringtheAurignacian,cutmarksontwomandibles
atVogelherd[49,50]andafifthmetacarpalatGeißenklösterle[39]alsoindicateskinning,
Figure 2.
Map of the Swabian Jura with the location of important Paleolithic cave sites from the Ach,
Lone, and Lauchert valleys. Four sites outlined in red provide Upper Paleolithic fox tooth pendants
and are the focus of this study: Hohle Fels (1) Geißenklösterle (2), Brillenhöhle (3), Hohlenstein-Stadel
(4). (Map provided by Christian Sommer/ROCEEH (2019): A map collection of the Paleolithic of the
Swabian Jura (Version 1.0.0) (Data set). (modified by F.V.).
Evidence of fox butchery is rare compared to other taxa in the Swabian Jura [
44
].
During the Middle Paleolithic, a single phalanx from Bockstein exhibits a circular cut mark
associated with skinning [
48
]. During the Aurignacian, cut marks on two mandibles at
Vogelherd [
49
,
50
] and a fifth metacarpal at Geißenklösterle [
39
] also indicate skinning, while
cut marks on a rib and ulna at Hohle Fels are from defleshing [
37
]. During the Gravettian,
only two cut marks are present on fox remains at Hohle Fels, one on a pelvis and the
other on a femur, both related to defleshing [
37
]. Lastly, during the Magdalenian, one
cut-marked ulna is recorded from Hohle Fels [
51
] and one cut-marked mandible is present
at Langmahdhalde [
52
], indicating defleshing and skinning, respectively. In addition to
cut marks, human tooth marks are also recorded on three Aurignacian and Gravettian fox
elements from Hohle Fels [
53
]. Combined, these traces clearly establish foxes were used for
both fur and food, as well as a raw material for the 40 fox tooth ornaments discussed in
this study.
Human–fox relationships also extended beyond human predation, as evidenced
by isotopic studies of fox paleoecology, which suggest some foxes were commensal to
humans [
38
,
44
,
54
,
55
]. Prior to the Upper Paleolithic, foxes inhabited expected niches as
either rodent specialists or commensals to large carnivores (for an overview on fox ecology
and behavior, see the Supplementary Materials). However, during the Aurignacian, a new
fox niche emerged, characterized by a diet of primarily reindeer with a smaller propor-
tion of rodents and other large game. The likely explanation is that some foxes became
commensal to humans to take advantage of food waste, perhaps in response to increasing
human population density [
38
,
55
]. By engaging in this synanthropic commensal behavior,
foxes were able to access new, more reliable food resources as well as protection from large
carnivores who were deterred by humans. Humans also benefited, gaining more frequent
access to foxes for food, fur, and raw materials, as well as a means of food disposal, which
indirectly protected humans from large carnivores [
54
]. This relationship lasted throughout
the Aurignacian and Gravettian before dogs and wolves largely took over the niche during
the Magdalenian, although some foxes remained commensal [38,55].
Quaternary 2023,6, 50 5 of 36
2. Materials and Methods
2.1. Archaeological and Comparative Materials
The complete assemblage includes 40 perforated fox tooth ornaments from the Au-
rignacian, Gravettian, and Magdalenian horizons of four sites in the Swabian Jura: Hohle
Fels, Geißenklösterle, Brillenhöhle, and Hohlenstein-Stadel (Table 1). Most are canines
(n = 32), although incisors (n = 4) and premolars (n = 2) are also present as well as some
indeterminate root fragments (n = 2).
Table 1.
List of studied perforated fox tooth ornaments from the Swabian Jura. QU = quadrant,
ID = find ID, GH = geological horizon, AH = archaeological horizon, Morph = morphological species
identification, GM = geometric-morphometric species identification from this study,
1
= Kölbl and
Conard [5].
Culture QU ID GH AH Species ID (Morph) Species ID (GM) Tooth
Hohle Fels:
NA (Upper Paleolithic) 28 964.1 1ka-7 0/I-IV Vulpes sp. Vulpes vulpes Canine
Gravettian 58 1295 3c IIc Vulpes lagopus 1Vulpes lagopus Canine
Gravettian 79 1228.29 3c-3d IIc-IId Vulpes sp. Vulpes lagopus Canine
Gravettian 66d 461 3bl IIb Vulpes sp. 1Vulpes vulpes Canine
Aurignacian-Gravettian 76 1227.1 5 IIe Vulpes sp. ? NA Root fragment
Aurignacian-Gravettian 59 2099.1 5-7 IIe-IV Vulpes lagopus 1NA Canine
Aurignacian 24 1137 7 IV Vulpes sp. Indeterminate Canine
Aurignacian 26 631.1 7a Va Vulpes sp. Vulpes lagopus Canine
Aurignacian 69 1713 7 IV Vulpes lagopus 1Vulpes vulpes? Canine
Aurignacian 89 1517.1 7 IV Vulpes sp. Vulpes lagopus Canine
Aurignacian 24 1278.1 7 IV Vulpes sp. Vulpes lagopus Canine
Aurignacian 65 1742 7 IV Vulpes sp. NA Canine
Geißenklösterle:
NA (likely Gravettian) 22 1A disturbed
disturbed
Vulpes sp. Vulpes vulpes ? Canine
NA (likely Gravettian) 22 1B disturbed
disturbed
Vulpes sp. Vulpes lagopus Canine
Gravettian 67 658.2 7 It Vulpes sp. NA Incisor
Gravettian 98 391 7 It Vulpes sp. Vulpes vulpes Canine
Gravettian 99 918 7 It Vulpes sp. Vulpes lagopus ? Canine
Gravettian 100 89 5b Ir Vulpes sp. 1Vulpes sp. Canine
Gravettian 140 226 7 It Vulpes sp. Vulpes lagopus Canine
Gravettian 98 286 7 It Vulpes sp. NA Canine
Gravettian 98 387 7 It Vulpes lagopus 1NA Canine
Aurignacian 59 369 16 IIIb Vulpes sp. 1Vulpes vulpes Canine
Aurignacian 58 487 15 III Vulpes sp. NA Canine
Aurignacian 67 1511 15 IIIa Vulpes sp. NA Canine
Hohlenstein-Stadel:
Aurignacian NA 1.0380 19./20. Meter,
6. Hieb NA Vulpes sp. Vulpes lagopus Canine
Aurignacian NA 2.0380 19./20. Meter,
6. Hieb NA Vulpes sp. Vulpes vulpes Canine
Aurignacian NA 3.0380 19./20. Meter,
6. Hieb NA Vulpes sp. Vulpes vulpes Canine
Aurignacian NA 4.0380 19./20. Meter,
6. Hieb NA Vulpes sp. Vulpes lagopus Canine
Aurignacian NA 5.0380 19./20. Meter,
6. Hieb NA Vulpes sp. Vulpes lagopus Canine
Aurignacian or Magdalenian NA 36-1 backdirt NA Vulpes sp. NA Incisor
Aurignacian or Magdalenian NA 90001-1 backdirt NA Vulpes sp. NA Incisor
Aurignacian or Magdalenian NA 9000-1 backdirt NA Vulpes sp. NA Incisor
Aurignacian or Magdalenian NA 14-1 backdirt NA Vulpes sp. Vulpes sp. Canine
Aurignacian or Magdalenian NA 141-1 backdirt NA Vulpes sp. Vulpes sp. Canine
Aurignacian or Magdalenian NA 178-1 backdirt NA Vulpes sp. NA Premolar
Aurignacian or Magdalenian NA 33-1 backdirt NA Vulpes sp. NA Premolar
Aurignacian or Magdalenian NA 167-1 backdirt NA Vulpes sp. ? NA Root fragment
Brillenhöhle:
Gravettian NA 132 NA VII Vulpes sp. NA Canine
Gravettian P7 131 NA VII Vulpes sp. NA Canine
Gravettian NA 130 NA VII Vulpes sp. NA Canine
Quaternary 2023,6, 50 6 of 36
The ornaments from Hohle Fels are stored at the University of Tübingen as part of a
larger study of the faunal assemblage. Most of the ornaments from Geißenklösterle are also
stored at the University of Tübingen, while two others are on display at the Landesmuseum
Württemberg Stuttgart alongside the ornaments from Brillenhöhle. The Hohlenstein-Stadel
ornaments are stored and curated at the Museum Ulm, some of which are on display.
The Hohle Fels assemblage includes 12 total ornaments, which come from the Au-
rignacian layers Vb-IId (n = 8), the Gravettian layers IIe-IIb (n = 3), and a profile collapse
spanning multiple Upper Paleolithic horizons (n = 1) [3,5,53,56–59].
Geißenklösterle hosts 12 total ornaments, which come from the Aurignacian layers
IIIb-III (n = 3), the Gravettian layers It-Ir (n = 7), and from disturbed contexts that are likely
Gravettian (n = 2) [
60
,
61
]. Here, we publish one of the Aurignacian ornaments (ID 1511) for
the first time. Two additional Gravettian ornaments are known from Kölbl and Conard [
5
]
(Kat. 75, ID 359) and the Geißenklösterle database (ID 786); however, these two objects
could not be located for this study and are not included in the total count.
For Brillenhöhle, the assemblage includes three Gravettian ornaments from layer VII
(Riek, 1973).
Lastly, the Hohlenstein-Stadel assemblage includes 13 total ornaments. Those exca-
vated in association with the lion man figurine (n = 5) are considered Aurignacian [
61
–
64
],
while those excavated from the backdirt (n = 8) cannot be assigned to a specific cultural
period [
63
,
65
]. The latter likely belong to the Aurignacian, although the Magdalenian
cannot be excluded.
The assemblage we present here is likely incomplete. It is likely that excavators in
the early 20th century overlooked smaller finds due to their coarser excavation methods.
Excavations are also ongoing at Brillenhöhle and Hohle Fels, and we can expect to discover
more animal tooth pendants from these sites in the future.
For the geometric morphometric analysis, our sample includes a total of 24 Aurig-
nacian and Gravettian ornaments from Hohle Fels (n = 10), Geißenklösterle (n = 7), and
Hohlenstein-Stadel (n = 7) (Table S2). To establish a comparative baseline, we also curated
42 modern, unmodified, adult fox canines (Table S3) from the comparative zooarchaeo-
logical collection of the University of Tübingen alongside two canines from the private
collection of Chris Baumann (University of Tübingen). The comparative sample includes
13 upper and 9 lower arctic fox canines (Banks Island, Canada) as well as 9 upper and
11 lower red fox canines (southwestern Germany).
2.2. Geometric Morphometric Methods
Micro-computed tomography (
µ
CT) scanning of the archaeological sample was carried
out with a Nikon XT H 320
µ
CT scanner at isotopic voxel length between 20 and 61
µ
from
the Terrestrial Palaeoclimatology working group at the University of Tübingen. For the
comparative sample, 16 arctic and 16 red fox canines were scanned with the same machine,
while 6 arctic and 4 red fox canines were scanned with a GE Phoenix v|tome|x s240
µ
CT
scanner at an isotopic voxel length between 15 and 55
µ
from the Paleoanthropology High
Resolution CT Laboratory at the University of Tübingen.
Our geometric morphometric analysis follows protocols established by Benazzi et al. [
66
]
and Röding et al. [
67
] for the cervical outline of hominin molars with some modifications for
fox canines. As the cervical outline lacks homologous fixed points and a clear start or end
point, landmark collection does not fall into the usual concept of fixed or semi-landmarks,
thus necessitating additional steps during data collection for the removal of orientation,
location, and absolute size from the landmark coordinates.
After scanning, we post-processed and sectioned the
µ
CT images in Avizo 9.2 (Vi-
sualization Sciences Group), orienting each tooth to establish a best-fit plane along the
cervical outline. We then virtually sectioned the images along this outline, removing the
portion below the cervical plane, so that only the crown remained. Next, in Rhinoceros
6 (Robert McNeel and Associates), we translated the best-fit plane into an x-y coordinate
system for landmark selection. To ensure consistent positioning, each tooth was rotated
Quaternary 2023,6, 50 7 of 36
so that the lingual surface was parallel to the right side of the z-axis. Additionally, the
area centroid of the cervical outline was calculated, upon which the outline was centered
and translated to the Cartesian coordinate position (10,10,0). After positioning, sixteen
landmark positions along the cervical outline were generated through equiangular spaced
radial vectors from the centroid. The coordinates at the intersection of the vector with
the cervical outline served as pseudo-landmarks and were recorded clockwise starting
from the middle-upper landmark. Following established methods [
68
,
69
], all left canines
were mirrored and treated as right canines. However, it should be noted that fluctuating
asymmetry in dentition may increase noise when left and right teeth are combined [67].
Before analyzing the landmark data, we tested for intra-observer error to evaluate
the alignment, reliability of landmark positions, and the relative reproducibility of in-
dividual landmarks. This was calculated as the percent error of the Euclidean distance
between the configuration centroid and repeated landmark measurements. To accomplish
this, we performed each step in Aviso and Rhinoceros five times for a random sample
with a time gap of five hours, one day, one week, and one month after the first anal-
ysis. The randomization was performed by assigning a number to each used sample,
and a random number generator was then used to select a sample for error measure-
ments. Afterward, for each repeated landmark set, the centroid and the 16 Euclidean dis-
tances between the centroid and each landmark were calculated using unscaled landmark
data. We then calculated the percent error for each landmark as well as the average and
standard deviation.
After data collection, we performed multivariate statistical analyses in the program
R (R Development Core Team 2011, version 4.1.2,
©
2021) using multiple packages:
Geomorph [
70
,
71
], ggplot2 [
72
], ggpubr [
73
], and ggrepel [
74
]. Analyses were conducted
separately for upper and lower canines. For some teeth, the cervical outline was absent or
unclear due to surface modifications or poor preservation. We considered five ornaments
too incomplete for further analysis (ID 391, ID 89, ID 1227.1, ID 14-1, ID 141-1). However,
for teeth with less severe damage, we attempted to reconstruct the cervical outline with
the function estimate.missing in geomorph (method = “TPS”). The method was successful
for a CN45 upper canine, ID 1A, ID 226, and ID 2.0380 but unsuccessful in the case of the
CN99 upper canine. The former were included in subsequent analyses, while the latter was
removed from the statistical sample.
We analyzed the landmark sets with two Principal Component Analyses (PCA)—a
Shape PCA (gm.prcomp function) and a Form PCA (prcom function) [
70
]. The Shape PCA
only utilizes the scaled and superimposed landmark coordinates, whereas the Form PCA,
defined as shape considered with size, uses the logarithmized centroid size added as a
variable. To satisfy the assumptions of our PCAs, we estimated Cook’s distance for each
individual to ensure our dataset did not include outliers or influential specimens, and no
canines fell over the recommended cut-off value [
75
]. Shape changes along the PCs were
visualized as landmark configurations at ±2 standard deviations [67].
2.3. Traceological Methods
Traceological analyses were carried out on the complete ornament assemblage in the
Material Culture Laboratory (hereafter MCL) at the University of Tübingen. We performed
the analysis using three microscopes to identify manufacture traces, perforation techniques,
and use-wear traces. While we analyzed the manufacturing traces on broken and complete
perforations, observations at higher magnification in search of use-wear traces were only
carried out on teeth with intact and well-preserved perforations.
Following well-established protocols for microwear analysesof material culture [
76
–
78
], we
combined observations at low and high magnification. Macroscopic traces were observed
with an Olympus SZX7 stereomicroscope with magnification ranging from 8
×
to 56
×
and equipped with an LED ring light source in combination with a Hirox HRX-01, a
3D digital microscope covering a magnification range of 20
×
–2500
×
. Observations at
higher magnification were performed with an Olympus BX53M metallographic microscope
Quaternary 2023,6, 50 8 of 36
in transmitted light with magnification up to 500
×
, alongside the aforementioned 3D
digital microscope.
Before analysis, the perforations of the best-preserved teeth were cleaned with a cotton
swab immersed in demineralized water. For the description of macro and micro traces, we
used established nomenclature [
78
–
81
]. When describing the location of traces, we used
the dial plate system [
82
], in which specific “hours” on the dial plate correspond to a sector,
i.e., sector 1 corresponds to hours 12 to 1, sector 2 to hours 1 to 2, and so on.
We compared the technological and use-wear traces observed on the archeological
specimens with a reference collection of 28 perforated fox teeth, which were produced
for this study (see Supplementary Materials). Additionally, comparisons were made
with previous microwear studies of animal tooth ornaments from other prehistoric and
ethnographic contexts [25,82,83,83–88].
3. Results
3.1. Geometric-Morphometric Results
Our novel approach to differentiate red versus arctic fox individuals from loose canines
was ultimately very successful. Of the total assemblage, we identified 17 (71%) ornaments
as either red (n = 10) or arctic fox (n = 7) individuals. Two ornaments (8%) were classified
as indeterminate, and five (21%) were excluded due to damage affecting the visibility of
the cervical outline (Table 2). Of the arctic foxes, six correspond to the Aurignacian and
four to the Gravettian. Of the red foxes, five correspond to the Aurignacian and two to
the Gravettian.
Table 2.
Species classification of the archaeological ornaments after geometric-morphometric analyses.
Site Red Fox Arctic Fox Indeterminate Unsuitable for Analysis
Geißenklösterle (n = 7) 2 3 0 2
Hohle Fels (n = 10) 3 4 2 1
Hohlenstein-Stadel (n = 7) 2 3 0 2
Total (n = 24) 7 10 2 5
The statistical analyses revealed that, while both the Shape and Form PCAs exhibited
significant distinction and little overlap between species, the Shape PCA is best suited for
this analysis. The Form PCA, which considers the size of the cervical outline, reveals more
variability between the archaeological ornaments than the comparative sample.
Error measurements fall under the acceptable 5% deviation between the five repeated
measurements (Figure S7). Landmark 2 exhibited the greatest error (1.3%), where the
error measurements fell under 0.4%. This may indicate landmark 2 was more susceptible
to variations in outline placement in Avizo or in subsequent orientation in Rhinoceros.
However, the overall low deviation between samples indicates our method is suitable for
fox canines.
3.1.1. Upper Canines
The upper canine assemblage includes the archaeological specimens ID 369, ID 226,
ID 1137, ID 964.1, ID 461, ID 1228, and ID 2.0380 alongside the modern comparative upper
canines (n = 21).
The Shape PCA (Figure 3a) cumulatively explains 84% of variance in the first two PCs,
with PC1 and PC2 explaining 64% and 20% of variance, respectively. Red and arctic fox
canines mainly differ across PC1, where arctic foxes plot more positively and red foxes plot
more negatively, although a small overlap occurs between 0.00 and 0.01. On PC2, red foxes
exhibit a narrower range than arctic foxes and fall mostly within the arctic fox range.
Quaternary 2023,6, 50 9 of 36
Quaternary2023,6,xFORPEERREVIEW10of38
Figure3.GM:Shape(a)andForm(b)PCAofuppercaninesandpendants.
TheFormPCA(Figure3b)cumulativelyexplains9%ofvarianceinthefirsttwoPCs,
withPC1andPC2explaining49%and33%ofvariance,respectively.SimilartotheShape
PCA,adistinctionbetweenredandarcticfoxesisapparent.Whileredfoxpresentsa
broaderrangethanarcticfoxinPC1,theprimaryseparationisvisibleinPC2,wherearctic
Figure 3. GM: Shape (a) and Form (b) PCA of upper canines and pendants.
Although most ornaments plot clearly within the arctic or red fox groups, some require
further consideration. ID 1137 plots in an area of overlap along both PC1 and PC2, so a
clear species assignment was not possible. Three pendants plot outside the distribution
of the recent upper canines for both species: ID 1228, ID 2.0380, and ID 5.0380. Although
clear species classifications were not possible, ID 1228 is likely an arctic fox, given the
more positive PC2 value, and ID 5.0380 is likely also an arctic fox, given the more positive
PC1 value.
Quaternary 2023,6, 50 10 of 36
The Form PCA (Figure 3b) cumulatively explains 9% of variance in the first two PCs,
with PC1 and PC2 explaining 49% and 33% of variance, respectively. Similar to the Shape
PCA, a distinction between red and arctic foxes is apparent. While red fox presents a
broader range than arctic fox in PC1, the primary separation is visible in PC2, where arctic
fox plots more positively and red fox plots more negatively. Areas of overlap occur between
−0.04 and 0.04 in PC1 and between 0.00 and 0.01 in PC2.
Although size is considered in the Form PCA, species classifications do not fundamen-
tally change from the Shape PCA; however, some specimens require further consideration:
Ornament ID 226 plots within the arctic fox range in the Shape PCA but plots slightly out-
side of the arctic fox and within the red fox range on PC2 in the Form PCA. This indicates
that ID 226 can be viewed as a larger arctic fox canine. As in the Shape PCA, ID 1137 plots
in an area of overlap between species, inhibiting a clear species assignment. ID 1228 still
plots outside of both groups, but within the red fox range on PC1 and in an overlapping
area on PC2; however, a clear assignment was not possible. ID 2.380 plots within the red fox
distribution on both PC1 and PC2; considering the Shape PCA, this tooth was tentatively
identified as a red fox canine. Lastly, ID 5.0380 plots outside of the red and arctic fox values
on PC1 and within the arctic fox values on PC2, indicating that the size of this tooth falls
outside of the range of recent arctic foxes.
3.1.2. Lower Canines
The lower canine assemblage includes the archaeological specimens ID 1A, ID 1B,
ID 918, ID 1278.1, ID 631, ID 1295, ID 1713, ID 1517, ID 1.0380, ID 3.0380, and ID 4.0380
alongside the modern comparative lower canines (n = 22).
The Shape PCA (Figure 4a) cumulatively explains 76% of variance in the first two
PCs, with PC1 and PC2 explaining 60% and 16% of variance, respectively. The species are
primarily distinguished on PC1, where arctic foxes plot more positively and red foxes more
negatively with an area of overlap between
−
0.01 and 0.01. PC2 exhibits similar ranges
between groups and is thus a poor indicator of species.
Most ornaments exhibit clear adherence to either red or arctic fox, although some
specimens are more ambiguous. ID 1A and ID 1713 both plot within the red fox group,
although overlap with the arctic fox group along PC1 makes this identification tentative.
ID 1B and ID 4.0380 both plot outside the distribution of recent fox canines, but their more
positive PC1 values indicate these individuals correspond to arctic foxes.
The Form PCA (Figure 4b) mostly confirms the identifications presented in the Shape
PCA and explains 84% of variance in the first two PCs, with PC1 and PC2 explaining 59%
and 25% of variance, respectively. The greatest species separation occurs along PC2 with
arctic fox plotting more positively and red fox more negatively with some overlap between
−0.01 and 0.01.
Several ornaments plot outside the defined species groups, indicating the size of
cervical outlines varied more in the archaeological sample than the modern sample. In the
Shape PCA, ID 1A is tentatively classified as a red fox given its more negative PC2 value,
which is supported by the Form PCA’s PC1 value. Although ID 1713 plots within the red
fox group, the potential for overlap is high, making the classification as red fox tentative.
ID 981 overlaps with the red fox range along PC1; however, the location within the arctic
fox group of the Shape PCA combined with the more positive PC2 values of the Form PCA
suggests a tentative identification as arctic fox. For ID 1B, ID 1295, ID 1.0380, and ID 4.0380,
more positive PC2 values indicate they belong to arctic foxes. Lastly, the more negative
PC2 values of ID 3.0380 indicates this individual is a red fox.
3.2. Traceological Results
We present the traceological results by site, focusing on root modifications to determine
perforation techniques. Additionally, we pay special attention to the identification of
potential macro and microwear traces related to the use of the teeth as pendants or as
ornaments sewn onto clothes or other goods.
Quaternary 2023,6, 50 11 of 36
Quaternary2023,6,xFORPEERREVIEW12of38
Figure4.GM:Shape(a)andForm(b)PCAoflowercaninesandpendants.
Figure 4. GM: Shape (a) and Form (b) PCA of lower canines and pendants.
3.2.1. Hohle Fels
Among the 12 fox canines analyzed at Hohle Fels, seven exhibit complete perforations,
while three are broken at the tip of the root and one fragment only represents the upper
part of the eyelet. The twelfth tooth is incompletely perforated. Overall, 11 teeth were
suitable for the macro and microscopic analysis (Table 3).
Quaternary 2023,6, 50 12 of 36
Table 3. List of archeological fox teeth analyzed in this study from Hohle Fels.
Site Culture Find
Number (ID) Tooth Integrity of
Perforation Technique of Perforation Use Traces
Hohle Fels Aurignacian 1517.1 Canine Complete Scraping + grooving +
drilling Used: rounding/
smoothing
Hohle Fels Aurignacian 1137 Canine Fragmentary Scraping Not diagnostic
Hohle Fels Aurignacian 631.1 Canine Complete Probably scraping +
widening Not diagnostic
Hohle Fels Aurignacian 1713 Canine Complete Scraping + grooving +
widening
Used: round-
ing/smoothing/Polish
and striae
Hohle Fels Aurignacian 1278.1 Canine Complete (broke
during analysis) Scraping + grooving Used: rounding/
smoothing
Hohle Fels Aurignacian 1742 Canine Complete Drilling Used: rounding/
smoothing
Hohle Fels Aurignacian-
Gravettian 2099.1 Canine Complete Scraping + grooving Possibly used
Hohle Fels Gravettian 1295 Canine Complete Scraping + grooving+
possibly widening Used: round-
ing/smoothing/Polished
Hohle Fels Gravettian 1228.29 Canine Perforation
unaccomplished Carving + Scraping +
grooving Not Used
Hohle Fels Aurignacian-
Gravettian 1227.1 Root
fragment Fragmentary NA NA
Hohle Fels Gravettian 461 Canine Complete Scraping + drilling Used: rounding/
smoothing
Hohle Fels NA 964.1 Canine Fragmentary Scraping + grooving Not diagnostic
Under the microscopes, the assemblage shows different degrees of preservation: most
teeth have shiny, damaged surfaces and two exhibit “very poor” preservation. Only two
teeth are considered “well preserved”.
Manufacturing Traces
At Hohle Fels, we identified two manufacturing techniques: (1) scraping/grooving
and (2) drilling.
1.
Scraping/grooving: nine teeth show traces of scraping or grooving, which we define
as isolated groups of close, narrow striations, and/or deep grooves (Figure 5a–c).
Technological deformations in sectors 1 and 6 of five teeth indicate deep gouging
for the purpose of creating an opening in the root (Figure 5a). In three cases, we
observed specular lateral deformations in sectors 3 and 9, which we interpret as the
result of the widening and smoothing of the perforations through a semi-circular
motion with a borer or pointed lithic tool (Figure 5c,d and comparison in the Supple-
mentary Materials and Figure S2g–i). This scraping/grooving technique was always
performed bifacially and resulted in elongated perforations on the vertical axis of the
root (Figure 5a,c). Interestingly, tooth ID 1228 exhibits bilateral whittling/carving
traces complemented by scraping, but it was abandoned during early manufacture
before achieving a complete perforation, providing some insight into the order of
operations (Figure 5e).
Quaternary 2023,6, 50 13 of 36
Quaternary2023,6,xFORPEERREVIEW15of38
Figure5.OverviewoftechnologicaltracesrecordedonfoxteethfromHohleFels.(a)ID964,groov-
ingmarks;(b)ID1137,scrapingandgroovingmarks;(c)ID1713,scrapingandgroovingmarksand
evidenceofinnerwideningofthehole(noticethelateralhalf-moonenlargement);(d)ID1517.1,
circularstriationsbysemi-rotationaldrilling;(e)ID1228.2,whittling/carvingmarksandscraping
traces;(f)ID461,drillingtraces;(g)ID1742,drillingtraces.
Figure 5.
Overview of technological traces recorded on fox teeth from Hohle Fels. (
a
) ID 964, grooving
marks; (
b
) ID 1137, scraping and grooving marks; (
c
) ID 1713, scraping and grooving marks and
evidence of inner widening of the hole (notice the lateral half-moon enlargement); (
d
) ID 1517.1,
circular striations by semi-rotational drilling; (
e
) ID 1228.2, whittling/carving marks and scraping
traces; (f) ID 461, drilling traces; (g) ID 1742, drilling traces.
2.
Drilling: Two teeth (ID 1742 and 461) exhibit clear concentric striations on the inner
walls of their perforations, characteristic of the rotating action of a borer or a pointed
Quaternary 2023,6, 50 14 of 36
lithic tool. According to our reference collection, the perforation of tooth ID 461 was
created by bifacial free hand drilling (Figure 5f). Tooth ID 1742 was probably bifacially
perforated with the use of a bow drill or a hafted hand drill (Figure 5g). The difference
between the two techniques is indicated by the high regularity of the drilling traces
and the asymmetrical shape of the perforation, which indicates bow drilling. These
features may result from the difficulty of controlling a bow drill at high speeds (see
for comparison Figure S1h). Only the root surface of ID 461 appears to be prepared by
scraping prior to drilling.
Use-Wear Related Traces
All seven teeth with complete perforations showed very round and smooth perfo-
rations, specimens ID 1742 and ID 1713 in particular. We found clear microwear traces
associated with use on two specimens. On tooth ID 1295 and ID 1713, we recorded smooth
and polished areas in sectors 1, 2, and 12 with linear traces in the form of long, shallow,
and straight striations running parallel to the longitudinal axis of the root (Figure 6a–d, see
Figure S6e for comparison).
Quaternary2023,6,xFORPEERREVIEW16of38
Figure6.Overviewofuse-relatedtracesrecordedonfoxteethfromHohleFels.(a)ID1742,high
roundingandsmoothing;(b)ID1295,roundingandsmoothing;(c)ID1295,roughpolishdeveloped
insector12;(d)ID1713,smoothpolishwithparallellongandnarrowstriationsdevelopedinsector
3.
3.2.2.Geißenklösterle
AtGeißenklösterle,foxcaninesarerepresentedby12totalspecimensdividedasfol-
lows:fiveteethwithcompleteperforations,fivewithbrokenperforations,onenon-hu-
manmodifiedtooth,andanothermodifiedtoothwithabrokenrootdisplayingnoperfo-
ration(Table4).Thestateofpreservationisvariable:twospecimensexhibitveryshiny
surfaces,oneisaffectedbypatina,andthreehavecalciteencrustationaroundtheholes,
whiletheremainderareinmedium-goodstateofpreservation.
ManufacturingTraces
AsinHohleFels,atGeißenklösterleweidentifiedtwomainproductiontechniques:
(1)scraping/grooving/gougingand(2)drilling.
1. Scraping/grooving/gouging.Scrapingandgroovingappearedineightcases,some-
timescombinedwithgouging.Inallcases,thistechniquewasperformedbifacially.
Scrapingisrepresentedbyabundleofcloseandnarrow,longandshortstriations,
whilegroovingisattestedbydeepisolatedlonggrooves(Figure7a–c).Inthecaseof
ID391,theperforationwasopenedbybifacialgougingandsuperficialcarvingofthe
root,whosetracesareevidentinsectors12and6(Figure7d).Theperforationsasso-
ciatedwiththistechniqueareelongatedorsub-oval.
2. Drilling.DrillingisattestedatGeißenklösterlefromonlytwoteeth,bothshowing
tracesofscrapingtopreparethesurfaceoftheroot.Clearcircularandconcentric
drillingtracesarevisibleatlowandhighmagnification.Theirregularityofthe
Figure 6.
Overview of use-related traces recorded on fox teeth from Hohle Fels. (
a
) ID 1742, high
rounding and smoothing; (
b
) ID 1295, rounding and smoothing; (
c
) ID 1295, rough polish developed
in sector 12; (
d
) ID 1713, smooth polish with parallel long and narrow striations developed in sector 3.
3.2.2. Geißenklösterle
At Geißenklösterle, fox canines are represented by 12 total specimens divided as
follows: five teeth with complete perforations, five with broken perforations, one non-
human modified tooth, and another modified tooth with a broken root displaying no
perforation (Table 4). The state of preservation is variable: two specimens exhibit very
Quaternary 2023,6, 50 15 of 36
shiny surfaces, one is affected by patina, and three have calcite encrustation around the
holes, while the remainder are in medium-good state of preservation.
Table 4. List of archeological fox teeth analyzed in this study from Geißenklösterle.
Site Culture Find
Number (ID) Tooth Integrity of
Perforation Technique of Perforation Use Traces
Geißenklösterle Aurignacian 487 Canine Fragmentary Scraping + drilling/
lateral carving Not eligible
Geißenklösterle Aurignacian 1511 Canine Fragmentary Possibly scraped +
widening Not eligible
Geißenklösterle Aurignacian 369 Canine Fragmentary Grooving Not diagnostic
Geißenklösterle most likely
Gravettian 1A Canine Fragmentary Gouging + widening Not diagnostic
Geißenklösterle most likely
Gravettian 1B Canine Complete Scraping + drilling Not diagnostic
Geißenklösterle Gravettian 391 Canine Complete Grooving + Carving
Used: rounding/polish/striae
(lateral sides)
Geißenklösterle Gravettian 918 Canine Non-human modified /
Geißenklösterle Gravettian 658.2 Incisor (?) Complete Light scraping + possibly
widening Used: rounding/smoothing/
Polish (apex)
Geißenklösterle Gravettian 89 Canine Not
preserved NA NA
Geißenklösterle Gravettian 226 Canine Fragmentary Scraping + grooving Not diagnostic
Geißenklösterle Gravettian 387 Canine Complete Possibly scraping +
widening
Used: rounding/polish/striae
(lateral sides)
Geißenklösterle Gravettian 286 Canine Complete Scraping/grooving +
widening Not diagnostic
Manufacturing Traces
As in Hohle Fels, at Geißenklösterle we identified two main production techniques:
(1) scraping/grooving/gouging and (2) drilling.
1.
Scraping/grooving/gouging. Scraping and grooving appeared in eight cases, some-
times combined with gouging. In all cases, this technique was performed bifacially.
Scraping is represented by a bundle of close and narrow, long and short striations,
while grooving is attested by deep isolated long grooves (Figure 7a–c). In the case
of ID 391, the perforation was opened by bifacial gouging and superficial carving of
the root, whose traces are evident in sectors 12 and 6 (Figure 7d). The perforations
associated with this technique are elongated or sub-oval.
2.
Drilling. Drilling is attested at Geißenklösterle from only two teeth, both showing
traces of scraping to prepare the surface of the root. Clear circular and concentric
drilling traces are visible at low and high magnification. The irregularity of the
perforations suggests the use of free hand drills (Figure 7e,f). Interestingly, the root of
ID 487 is worked in two ways, suggesting two different attachment methods. The root
was previously scraped and subsequently drilled in order to obtain a circular hole,
which later broke (Figure 7f,g). Underneath the perforation, the root is carved on both
lateral sides to create two deep incisions, which might have served to secure a thread
(Figure 7f,g). Unfortunately, we could not establish whether the perforation broke
ab antiquo or was a result of post-depositional events. Moreover, we were unable to
perform observations at higher magnification in search of use-related traces on the
carved lateral sides, as the piece could not leave the museum. We thus cannot establish
whether the root was carved after the perforation broke to re-facilitate attachment or
if it was a stylistic choice.
Quaternary 2023,6, 50 16 of 36
Quaternary2023,6,xFORPEERREVIEW18of38
Figure7.OverviewoftechnologicaltracesrecordedonfoxteethfromGeißenklösterle.(a)ID226,
groovingmarks;(b)ID658.2scrapingmarks;(c)ID360,groovingmark;(d)ID391,carvingtraces
insector12;(e)ID1b,drillingtraces;(f)ID487,drillingtracesandlateralcarvednotches;(g)ID487,
closeupofscrapingmarks.
Figure 7.
Overview of technological traces recorded on fox teeth from Geißenklösterle. (
a
) ID 226,
grooving marks; (
b
) ID 658.2 scraping marks; (
c
) ID 360, grooving mark; (
d
) ID 391, carving traces in
sector 12; (
e
) ID 1b, drilling traces; (
f
) ID 487, drilling traces and lateral carved notches; (
g
) ID 487,
close up of scraping marks.
Use-Wear Related Traces
Out of four complete perforations, three showed polish and shallow long striations
on the internal rim. In particular, ID 658.2 showed a small, polished area in sector 12 with
Quaternary 2023,6, 50 17 of 36
a developed rounding of the internal rim (Figure 8a) while ID 391 and ID 76 exhibited a
smooth polish on sectors 3 and 9 associated with striations (Figure 8b,c).
Quaternary2023,6,xFORPEERREVIEW19of38
Figure8.Overviewofuse-relatedtracesrecordedonfoxteethfromGeißenklösterle.(a)ID658.2,
polishdevelopedinsector12;(b)ID38F,smoothpolishdevelopedinsector9;(c)ID391,smooth
polishwithshortshallowstriationsdevelopedinsector9.
3.2.3.Hohlenstein-Stadel
PerforatedfoxteetharerepresentedatHohlenstein-Stadelby13specimensincluding
sevencanines,threeincisors,andtwopremolars.Arootfragmentshowingacomplete
perforationcouldnotbeassignedtoaspecifictoothtype(Table5).Thestateofpreserva-
tionoftheentiresampleisgood.Mostteethdisplayedcompleteperforations,andonly
fourwerebrokenintheupperpartoftheeyelet.Unfortunately,10teethshowedvarnish
aroundtheperforations,andsomehadtracesofglueinsidetheholes,whichhampered
observationathighmagnification.
ManufacturingTraces
OnlyonetechniquewasattestedatHohlenstein-Stadelforperforatingfoxteeth:
scraping.All13teethwerelongitudinallyscrapedtoflattenandthintheroot,themarks
ofthispreparationbeingvisibleaslongandirregularstriations(Figure9a,b,d,e).Insev-
eralcases,theextensionofthestriationsexceededtheareaofthehole,reachingthe
enamel.Bifaciallongitudinalscrapingwasperformeduntilperforationswereachieved,
Figure 8.
Overview of use-related traces recorded on fox teeth from Geißenklösterle. (
a
) ID 658.2,
polish developed in sector 12; (
b
) ID 38F, smooth polish developed in sector 9; (
c
) ID 391, smooth
polish with short shallow striations developed in sector 9.
3.2.3. Hohlenstein-Stadel
Perforated fox teeth are represented at Hohlenstein-Stadel by 13 specimens including
seven canines, three incisors, and two premolars. A root fragment showing a complete
perforation could not be assigned to a specific tooth type (Table 5). The state of preservation
of the entire sample is good. Most teeth displayed complete perforations, and only four
were broken in the upper part of the eyelet. Unfortunately, 10 teeth showed varnish around
the perforations, and some had traces of glue inside the holes, which hampered observation
at high magnification.
Quaternary 2023,6, 50 18 of 36
Table 5. List of archeological fox teeth analyzed in this study from Hohlenstein-Stadel.
Site Culture Find
Number (ID) Tooth Integrity of
Perforation Technique of Perforation Use Traces
Hohlenstein-Stadel Aurignacian 1.0380 Canine Complete Scraping + widening Most likely
used: rounding
Hohlenstein-Stadel Aurignacian 2.0380 Canine Complete Scraping + carving Most likely
used: rounding
Hohlenstein-Stadel Aurignacian 3.0380 Canine Complete Scraping Most likely
used: rounding
Hohlenstein-Stadel Aurignacian 4.0380 Canine Fragmentary Scraping + grooving Not diagnostic
Hohlenstein-Stadel Aurignacian 5.0380 Canine Complete Scraping Not diagnostic
Hohlenstein-Stadel Aurignacian or
Magdalenian 14.1 Canine Fragmentary Grooving Not diagnostic
Hohlenstein-Stadel Aurignacian or
Magdalenian 141.1 Canine Fragmentary Scraping + possibly
widening Not diagnostic
Hohlenstein-Stadel Aurignacian or
Magdalenian 167.1 Root
fragment Complete Possibly abrasion/
scraping + widening Most likely
used: rounding
Hohlenstein-Stadel Aurignacian or
Magdalenian 178.1 Premolar Complete Possibly abrasion/
scraping + widening Most likely
used: rounding
Hohlenstein-Stadel Aurignacian or
Magdalenian 33.1 Premolar Complete Carving + widening Most likely
used: rounding
Hohlenstein-Stadel Aurignacian or
Magdalenian 36.1 Incisor (?) Complete Possibly abrasion/
scraping + widening Used: rounding/
smoothing
Hohlenstein-Stadel Aurignacian or
Magdalenian 9000.1 Incisor (?) Complete Possibly abrasion/
scraping + possibly
widening
Most likely
used: rounding
Hohlenstein-Stadel Aurignacian or
Magdalenian 9001.1 Incisor (?) Fragmentary Possibly abrasion/
scraping + possibly
widening Not diagnostic
Manufacturing Traces
Only one technique was attested at Hohlenstein-Stadel for perforating fox teeth:
scraping. All 13 teeth were longitudinally scraped to flatten and thin the root, the marks of
this preparation being visible as long and irregular striations (Figure 9a,b,d,e). In several
cases, the extension of the striations exceeded the area of the hole, reaching the enamel.
Bifacial longitudinal scraping was performed until perforations were achieved, resulting
in perforations with an elongated form (Figure 9b,d,e). However, we recognized traces
corresponding to the widening of the hole, likely to achieve a more rounded and regular
shape (Figure 9c).
It is interesting to notice that the five canines likely associated with the Aurignacian
horizon show a common singular technological trait: the upper part of the root is strongly
bent, almost to form a step between the end of the perforation and the apex of the root
(Figure 9d–f). This was due to the deep and strong scraping resulting in an almost carved
surface on the apical side of the perforation, well visible in tooth ID 2.380 (Figure 9e).
Hohlenstein-Stadel is the only site in the Swabian Jura to report perforated fox premo-
lars. Interestingly, two premolars and three incisors show flat facets around the perforation
in association with striations, which differ from those on the canines in being wider, more
regular, deeper, and in an oblique direction (Figure 9g,h). In those cases, abrasion against a
coarse surface might better explain these traces than scraping with a lithic tool. However,
they are subsequently worked following the same scheme, namely by flattening the surface
until achieving the perforation and later widening the hole with bifacial semi-rotation.
Quaternary 2023,6, 50 19 of 36
Quaternary2023,6,xFORPEERREVIEW21of38
Hohlenstein-
Stadel
Aurignacianor
Magdalenian9001.1Incisor(?)Fragmen-
tary
Possiblyabrasion/scraping
+possiblywideningNotdiagnostic
Figure9.OverviewoftechnologicaltracesrecordedonfoxteethfromHohlenstein-Stadel.(a)ID
3.380,scrapingmarks;(b)ID1.380,scrapingandgroovingmarks;(c)ID178.1,wideningbysemi-
rotationaldrilling;(d)ID3.380,deepscrapingmarksresultinginabentupperroot;(e)ID2.380,
Figure 9.
Overview of technological traces recorded on fox teeth from Hohlenstein-Stadel. (
a
) ID
3.380, scraping marks; (
b
) ID 1.380, scraping and grooving marks; (
c
) ID 178.1, widening by semi-
rotational drilling; (
d
) ID 3.380, deep scraping marks resulting in a bent upper root; (
e
) ID 2.380,
striations and carving marks; (
f
) ID 33.1, deep carving resulting in a bent upper root; (
g
) ID 167.1,
abrasive/scraping marks; (h) ID 36.1, flat facet and abrasive/scraping marks.
Quaternary 2023,6, 50 20 of 36
Use-Wear Related Traces
Among the teeth with complete perforations, five showed a well-developed rounding
of their internal and external rim, especially localized in sectors 12, 3, and 9 (Figure 10a–d).
On five better-preserved teeth subjected to observation at higher magnification, we did
not identify any diagnostic use-related microwear. The rest of the assemblage exhibited
traces of varnish and glue and, consequently, it was not possible to perform more detailed
microscopic observations.
Quaternary2023,6,xFORPEERREVIEW22of38
striationsandcarvingmarks;(f)ID33.1,deepcarvingresultinginabentupperroot;(g)ID167.1,
abrasive/scrapingmarks;(h)ID36.1,flatfacetandabrasive/scrapingmarks.
Figure10.Overviewofuse-relatedtracesrecordedonfoxteethfromHohlenstein-Stadel.(a)ID
178.1,roundingandsmoothingoftheinnerandouterrim;(b)ID36.1,roundingandsmoothingof
therim;(c)ID9001.1roundingoftherim;(d)ID178.1,roundingoftheinnerwalloftheperforation.
3.2.4.Brillenhöhle
AtBrillenhöhle,threeperforatedcaninescomefromtheGravettian.Twoshowcom-
pleteperforationsandgoodpreservationofthesurfaces,whileoneisbrokenontheapical
sideoftheroot(Table6).Wecouldonlymakemacroscopicobservationsoftheseteeth
andwereunabletoperformobservationsathighermagnificationbecausethespecimens
couldnotleavethemuseum.
ManufacturingTraces
Theperforationsofallthreeornamentswereachievedbyscraping/grooving.How-
ever,scrapingwasnotperformedasapreliminarythinningtreatment,butrathertodi-
rectlyachievetheperforation.OnID130,theremaininglowerpartoftheperforationin
sector6isV-shaped,suggestinganoriginalelongatedholemadebygrooving/carving
(Figure11a).SpecimenID132exhibitedunifacialshortgroovesandadepressiononthe
upperpartoftheholeproducedbydeeplygroovingtheroot.Ontheotherface,theper-
forationshowedtracesofwideningforregularizingtheshapeofthehole(Figure11b).
ToothID131carriesafewtechnologicaltracesandexhibitsaroundandsmoothperfora-
tion.Duetothepresenceofencrustedsoilparticlesontheinnerwall,itwasnotpossible
toascertainiftheperforationwasmadebydrillingorbysemi-rotationforwideningthe
hole.Noclearrotationaltracesarevisibleeveniftheroundnessoftheholesuggestsa
Figure 10.
Overview of use-related traces recorded on fox teeth from Hohlenstein-Stadel. (
a
) ID 178.1,
rounding and smoothing of the inner and outer rim; (
b
) ID 36.1, rounding and smoothing of the rim;
(c) ID 9001.1 rounding of the rim; (d) ID 178.1, rounding of the inner wall of the perforation.
3.2.4. Brillenhöhle
At Brillenhöhle, three perforated canines come from the Gravettian. Two show com-
plete perforations and good preservation of the surfaces, while one is broken on the apical
side of the root (Table 6). We could only make macroscopic observations of these teeth and
were unable to perform observations at higher magnification because the specimens could
not leave the museum.
Manufacturing Traces
The perforations of all three ornaments were achieved by scraping/grooving. How-
ever, scraping was not performed as a preliminary thinning treatment, but rather to directly
achieve the perforation. On ID 130, the remaining lower part of the perforation in sector 6
is V-shaped, suggesting an original elongated hole made by grooving/carving (Figure 11a).
Specimen ID 132 exhibited unifacial short grooves and a depression on the upper part of
Quaternary 2023,6, 50 21 of 36
the hole produced by deeply grooving the root. On the other face, the perforation showed
traces of widening for regularizing the shape of the hole (Figure 11b). Tooth ID 131 carries a
few technological traces and exhibits a round and smooth perforation. Due to the presence
of encrusted soil particles on the inner wall, it was not possible to ascertain if the perforation
was made by drilling or by semi-rotation for widening the hole. No clear rotational traces
are visible even if the roundness of the hole suggests a complete rotational movement.
A depression in the upper part of the root might suggest a scraping/grooving activity
(Figure 11c).
Table 6. List of archeological fox teeth analyzed in this study from Brillenhöhle.
Site Culture Find Number
(ID) Tooth Integrity of
Perforation
Technique of
Perforation
Use Traces on
Perforation
Brillenhöhle Gravettian 132 Canine Complete Grooving/carving +
widening
Most likely used:
rounding/smoothing
Brillenhöhle Gravettian 131 Canine Complete Scraping/grooving +
widening or drilling
Most likely used:
rounding/smoothing
Brillenhöhle Gravettian 130 Canine Fragmentary Scraping + grooving Not diagnostic
Quaternary2023,6,xFORPEERREVIEW23of38
completerotationalmovement.Adepressionintheupperpartoftherootmightsuggest
ascraping/groovingactivity(Figure11c).
Table6.ListofarcheologicalfoxteethanalyzedinthisstudyfromBrillenhöhle.
SiteCultureFindNumber
(ID)ToothIntegrityof
PerforationTechniqueofPerforationUseTracesonPerfo-
ration
BrillenhöhleGravettian132CanineCompleteGrooving/carving+widen-
ing
Mostlikelyused:
rounding/smoothing
BrillenhöhleGravettian131CanineCompleteScraping/grooving+widen-
ingordrilling
Mostlikelyused:
rounding/smoothing
BrillenhöhleGravettian130CanineFragmentaryScraping+groovingNotdiagnostic
Figure11.OverviewoftechnologicaltracesrecordedonfoxteethfromBrillenhöhle.(a)ID130,
carvingmarks,(b)ID132,groovingmarks;(c)ID131,groovingmarkintheupperperforation.
4.Discussion
TheuseofanimalteethasornamentsduringtheUpperPaleolithicprovidesvaluable
insightsintothecreativity,symbolism,andculturalpracticesofourancestors,givingus
aglimpseintotheirwayoflifeandhowtheyinteractedwiththenaturalandanimal
world.Indeed,theuseofanimalremainsasasourceoffood,aswellasarawmaterial
(e.g.,teethforornaments,furforclothes,bonesfortools),evidencesanintimaterelation-
shipbetweenhumansandanimals.Thisisparticularlytrueforanimalslikefoxes,which
Figure 11.
Overview of technological traces recorded on fox teeth from Brillenhöhle. (
a
) ID 130,
carving marks, (b) ID 132, grooving marks; (c) ID 131, grooving mark in the upper perforation.
Quaternary 2023,6, 50 22 of 36
4. Discussion
The use of animal teeth as ornaments during the Upper Paleolithic provides valuable
insights into the creativity, symbolism, and cultural practices of our ancestors, giving us a
glimpse into their way of life and how they interacted with the natural and animal world.
Indeed, the use of animal remains as a source of food, as well as a raw material (e.g.,
teeth for ornaments, fur for clothes, bones for tools), evidences an intimate relationship
between humans and animals. This is particularly true for animals like foxes, which exhibit
behaviors showing their charisma, adaptiveness, and ability to influence their surroundings
and shape human communities. The human–fox relationship—today, as in the past—is
often characterized by the commensal behavior of foxes and their coexistence with humans
and their ability to thrive in human-constructed environments [
38
,
44
,
54
,
55
,
89
–
91
]. The
result of this relationship is still observed in urban and suburban environments today,
where foxes often live in close proximity to humans [92,93].
This human–fox relationship also existed in prehistory, as evidenced by the frequent
occurrence of fox bones and teeth in the archaeological record, many of which were used
to produce ornaments [
55
,
94
]. Although foxes are not the most common taxa in the
archeological record of the Swabian Jura [
55
], their perforated teeth are a particularly
informative class of artifacts with the potential to shed light on the role foxes played in past
human societies.
4.1. Production of Fox Tooth Ornaments in the Swabian Jura
The results presented in this study show that canines were the preferred tooth type for
ornamentation made from foxes in the Swabian Jura. In our assemblage, Hohlenstein-Stadel
exhibits ornaments made from other teeth, namely incisors (one of which is also present
at Geißenklösterle) and premolars, albeit in lesser quantities than canines. Based on the
available published data, only a handful of other sites share this variability in tooth choice,
one of which is the Magdalenian site of Petersfels (Germany), a few hundred kilometers
south of the Ach and Lone valleys in the Swabian Jura (see Table S1) [
95
–
97
]. Given the
uncertain context of some of the Hohlenstein-Stadel ornaments as either Aurignacian or
Magdalenian, it is possible that the use of premolars and incisors indicates a Magdalenian
origin for these five ornaments, like those represented at Petersfels.
From a traceological point of view, we noticed that five Hohlenstein-Stadel canines
(Aurignacian) were processed by scraping with a lithic tool, while the two premolars
and three incisors (Aurignacian or Magdalenian) were possibly abraded to achieve the
perforations. This difference might suggest a deliberate cultural choice in technique.
However, the other two canines from the uncertain context also show traces in line with
the five Aurignacian canines, suggesting the chosen technique was more dependent on
tooth type and likely morphology rather than a cultural or individual preference. Tooth
morphology also influences the practical aspects of perforation. Canines, for example, are
larger with a greater root surface area than other teeth, making ornament production easier
and perhaps providing a specific aesthetic function regarding their shape, size, and luster.
Despite the likely morphological influence on production technique, some of the
Hohlenstein-Stadel ornaments may still belong to the Magdalenian, given their uncertain
stratigraphic context. However, a cultural explanation for tooth preference is complicated
by the broader record of fox tooth ornaments throughout the Upper Paleolithic. Ornaments
made from fox incisors and premolars are found at numerous sites across Europe in the
Magdalenian but also the Gravettian [
34
,
86
,
94
–
96
,
98
,
99
] (Table S1). Retracing the context of
the Hohlenstein-Stadel ornaments, as well as traceological comparison with other ornament
assemblages, may thus illuminate the potential hunter-gatherer networks that influenced
tooth choice in the Swabian Jura.
Regarding the perforation techniques, our four sites show a common trend: the
preferred technique was bifacial scraping and grooving combined with the widening of
the hole to regularize the perforation. In most of the analyzed teeth, the scraping and
grooving technique was not performed to thin the root as a surface preparation, but rather
Quaternary 2023,6, 50 23 of 36
to make the perforation itself. Only two specimens (one from Hohle Fels and one from
Geißenklösterle) were instead scraped before being rotationally drilled. In one case, we
recorded deep gouging around the perforation (ID 2.0380), while in a few others, the root
was scraped and carved producing a hollowed facet around the hole (ID 1228, ID 33-1).
According to our experimental reproductions, scraping and grooving was an efficient
method of perforation that did not require any prior surface treatment. For teeth from
larger animals with thicker roots, this might not be the case. By analyzing perforated
teeth from other animal species at Hohle Fels and Geißenklösterle, we noticed that horse
incisors always exhibit a scraped surface before the perforation is achieved by drilling (F.V.,
personal communication). This reflects the need to adjust perforation techniques according
to the constraints of the raw material.
In our assemblage, we did not notice any correlation between a specific technique
with a particular cultural horizon or site. This might reflect our singular focus on fox teeth.
Comparing the perforated teeth from different animal species at these sites (for an overview
of numbers and percentages see Table S6) may highlight variations in techniques revealing a
tooth functional-dependent choice, rather than cultural or individual preferences. However,
it is worth noting that the five canines assigned to the Aurignacian at Hohlenstein-Stadel
show the same technological trait of bent upper roots (Figure 9d,f). This signature is
a distinct marker for the site and strengthens the attribution of the teeth to the same
Aurignacian horizon. Moreover, we suggest here the possibility that these five teeth were
worked by the same person using a distinctive and repetitive technical gesture, which was
not recorded at the other sites.
Another aspect to discuss is if and how the teeth were worn. Microtraces on teeth
are always very ambiguous because of their naturally shiny surfaces and animal life
history, especially considering the various post-depositional processes that can also impact
preservation. However, macro traces such as rounding and smoothing localized on the
inner and outer rim of perforations are a clear marker of use. More difficult is to reconstruct
when teeth served as hanging pendants or when they were sewn to clothes or other objects.
In our assemblage, we considered all well-preserved teeth with complete perforations
to be most likely used, based on the presence of developed and localized rounding and
smoothing of the perforations (Tables 3–6). For the teeth exhibiting use-related polishes
and striations, we do not feel confident in assigning a specific attachment mode or binding
method. However, the presence of striations recorded on some specimens may be related
to the use of abrasive strings like nettle or other rough vegetal/animal materials. Strings
treated with colorant such as ochre can also develop smoother polish and bundles of
striations. Indeed, we recorded traces of red pigment in seven teeth from Hohle Fels and
one from Geißenklösterle. Other than ID 1713 (Figure 5c), which shows reddish traces
inside the inner rim in association with rounding, polish, and striations likely from ochred
strings, we interpreted most red coloration as post-depositional, given its presence within
recent fractures, in the cavity of the root, and/or mixed with the adhering sediment.
4.2. Selection of Red versus Arctic Fox
The relationship between humans and a specific fox species—red or arctic—is often
impossible to determine in the archaeological record, given the nondiagnostic nature of
isolated faunal elements. Differentiation in loose teeth is particularly tricky (see Supple-
mentary Materials). Although some teeth are distinguishable based on size, canines are
more ambiguous [
100
], and prior attempts at metric differentiation [
101
] exhibit notable
flaws that decrease the reliability of species determinations [
39
,
102
–
105
]. For this reason,
a geometric morphometric approach is advantageous and a potential improvement on
existing morphological and metric methods.
With the help of our modern comparative sample, we successfully determined the
species of 17 (71%) ornaments from our archaeological sample. Although some ornaments
plotted outside the recent fox ranges, their position within the PCAs still shows tendencies
toward a specific classification. The larger range of the archaeological sample may indicate
Quaternary 2023,6, 50 24 of 36
noise in landmark placements, which is expected in archaeological specimens due to post-
depositional damage resulting in alterations to the cervical outline. Other factors, such as
variation between individuals, sexes, and populations also impact crown heights.
Our results show that both red and arctic foxes were used to produce ornaments in
the Swabian Jura. On a site level, there is no clear preference for species; however, the total
assemblage includes slightly more arctic than red foxes. Between cultural periods, there is
also no strong preference for species—the Aurignacian is represented by five red foxes and
six arctic foxes, while the Gravettian includes two red foxes and four arctic foxes.
Concerning the broader Upper Paleolithic record of fox tooth ornaments, few studies
provide species-specific identifications. Our literature review revealed limited information
from Aurignacian assemblages, but more from the Gravettian and the Magdalenian. In
the Gravettian, two notable sites report high numbers of perforated arctic fox teeth: 42
at the Czech site of DolníVˇestonice I (alongside 31 indeterminate) and 41 at the Russian
site of Zaraysk (Table S1) [
99
,
106
,
107
]. For the Magdalenian, 16 perforated arctic fox
teeth are reported from Petersfels (alongside 25 indeterminate), at least 5 come from
Kniegrotte [
95
–
97
], and 162 were recovered from Wilczyce in Poland (Table S1) [
86
]. Few
sites report perforated red fox canines, with Dolni Vˇestonice I being the only clear example
from our literature review [99,106,107].
Despite the overwhelming abundance of perforated arctic fox canines, both arctic and
red foxes were available to hunter-gatherers across Europe. In the Swabian Jura, red and
arctic foxes were even found within the same layers at sites like Geißenklösterle [
39
]. Al-
though both species were likely sympatric during the Upper Paleolithic (see Supplementary
Materials), their abundance may have varied. In modern sympatric contexts, arctic foxes
are observed in smaller numbers [
108
–
111
]; if the same were true for the Paleolithic, hunter-
gatherers would have encountered red foxes more frequently. As acquisition methods like
traps and snares were non-selective, the ratio of captured animals would thus reflect this
natural abundance. However, setting traps at known fox dens, perhaps dens used by a
single species, may have enabled selective acquisition (see Supplementary Materials).
It is thus possible that the relative scarcity of arctic foxes on the landscape increased
their value as a motif for ornamentation and influenced their preferential exploitation across
Europe. However, local paleoenvironments and faunal assemblages should be considered.
Although foxes were likely sympatric in a broad sense, our modern understanding of
interspecific fox relationships may be a poor analog for the Paleolithic. Additionally, climate
and environmental conditions varied significantly throughout this period, sometimes
within a short timeframe, potentially influencing the degree of species overlap. The
resolution of the archaeological record may also limit our ability to parse moments in
time when conditions were more or less hospitable to a particular species. A closer look at
the ratio of red and arctic foxes in the broader faunal assemblages would further illuminate
their nuanced relationship within the local paleoenvironment. Biomolecular techniques,
such as aDNA and ZooMS [
112
,
113
], would also increase the reliability of these distinctions.
In the Swabian Jura, some information on the ratio of foxes in the broader faunal
assemblages is available [
39
,
46
,
51
,
55
]. As with the ornaments, there is no overwhelming
preference for arctic versus red fox. This near-equal distribution suggests the ornament
ratio likely reflects the hunted ratio and perhaps the ratio of animals on the landscape.
Butchery traces indicate foxes were used as food and for their fur. Concerning the former,
other species, namely ungulates, provided a far greater contribution to hunter-gatherer diet.
However, fox fur may have served a unique purpose. Arctic foxes are uniquely valued
amongst modern arctic hunter-gatherer populations for their small size and bright, white
color [
114
,
115
]. This then may be a convincing argument for the relative overabundance of
arctic fox canines elsewhere in Europe—an aesthetic or functional preference for their fur.
A final consideration is the method of identification. Arctic fox canines, due to their
small size, are far more recognizable with simple morphological and metric characteristics.
Those in the area of overlap may go unidentified, or are identified as arctic fox by default,
and few specimens are large enough to be confidently identified as red fox. Unfortunately,
Quaternary 2023,6, 50 25 of 36
few studies report their methods of species determination and the commonly applied metric
method [
101
] may be biased towards arctic foxes [
39
]. Future application of geometric-
morphometric methods, like those applied in this study, would help to clarify this point and
allow further investigation of the other factors influencing red versus arctic fox abundance
in the Upper Paleolithic.
4.3. Fox Tooth Ornaments in the Broader European Context
We report the data from our literature review of fox tooth ornaments across the
European Upper Paleolithic in Table S1. These ornaments first appear in small numbers
during the Châtelperronian at sites such as La Grotte du Renne [
20
,
116
]. Increasing in the
Aurignacian, fox tooth ornaments dominate various German, French, and Russian sites [
10
].
The richest French sites are those of Isturitz, Saint-Jean-de-Verges, and La Quina, showing
13, 14, and 28 perforated canines, respectively [
25
,
117
,
118
]. In Russia, Kostenki hosts at least
37 pierced fox teeth from the Initial and Early Upper Paleolithic layers, the richest for the
Aurignacian [
119
]. Comprehensive studies on the investigation of perforation techniques
are generally rare, but at Isturitz fox canines were first thinned by fine abrasion and then
bifacially scraped to open a perforation. Deep carving of the root followed by scraping and
bifacial semi-rotational perforation is also attested [25].
Foxes continued to be an important ornament resource during the Gravettian, espe-
cially in Central and Eastern Europe. Hunter-gatherers also began to exploit a greater
variety of tooth types, including incisors and premolars in addition to canines. In Czechia,
two sites in Moravia recorded a high number of fox teeth used as pendants. At Pavlov
I, a total of 284 perforated fox teeth were discovered, among which 146 are incisors, 105
are canines, and 33 are premolars [
94
]. The nearby site of DolníVˇestonice I reports 73
perforated teeth, of which 42 are canines [
106
,
107
]. Moving eastward, in Russia, the site of
Zaraysk exhibits only 32 incisors and 9 premolars for a total of 41 teeth, all assigned to arctic
fox and found together as a “necklace” [
99
]. The funerary site of Sunghir in Russia offers
the best glimpse into the complex worldview of some of the first anatomically modern
humans in Europe. Within the several burials, a large number of fox tooth ornaments (>300)
were discovered in situ and in close association with human remains, offering a snapshot of
how personal ornaments were worn by this group of hunter-gatherers [
17
,
120
]. Numerous
perforated fox teeth were found on the forehead of an adult male (Sunghir 1), >40 on the
cranial vault and >250 on the belt of another individual (Sunghir 2), one in association
with an isolated cranium (Sunghir 5), and at least two found in association with a final
individual (Sunghir 10). In addition to heavy adornment, the individuals were also covered
with a large quantity of red ochre and other grave goods [17].
Foxes are also well-represented in Magdalenian ornament assemblages from Germany
and France with 41 perforated teeth from Petersfels [
95
–
97
] and 45 from Gönnersdorf in
Germany [
31
,
121
,
122
] as well as at least 35 from the Magdalenian layers of Isturitz [
85
]
and La Vache [
123
], 12 from La Madeleine [
124
], and 13 from Bruniquel [
125
,
126
] in France.
Although canines were the preferred tooth type, Petersfels also exhibits several incisors
and premolars. With 162 fox teeth, the Polish site of Wilczyce is the richest with 149 pierced
incisors alongside 13 canines. These were all acquired from the upper jaws of a minimum
of 31 arctic foxes and may represent the remains of a necklace associated with a neonatal
burial [86].
These data show that the Upper Paleolithic sites of the Swabian Jura are in line with
the general trends observed across Europe concerning the use of fox teeth as ornaments.
Their exploitation does not represent a unique signature or a key cultural marker for the
region, but a shared and well-established behavior amongst European Upper Paleolithic
groups. Canines are the preferred tooth type in both the Swabian Jura and the whole of
Europe, while incisors and premolars are less frequently used, although their presence
might represent a cultural preference or individual choice within certain groups of hunter-
gatherers. The techniques of perforation used in the analyzed sites compare well with
other European sites; although, scraping seems more common to achieve the perforation
Quaternary 2023,6, 50 26 of 36
itself rather than as a surface preparation method. The ubiquitous presence of perforated
fox teeth throughout the Upper Paleolithic in Central Europe attests to the important role
and intimate relationship this animal established with prehistoric human communities.
Although rare, evidence of fox butchery in the Swabian Jura suggests they were hunted for
food, for their furs, and as a raw material for ornamentation. The diversity of resources
provided by foxes along with their commensal behavior likely granted them a special
significance in the ecological, dietary, functional, and symbolic realms of Upper Paleolithic
hunter-gatherers.
5. Conclusions
Our results provide insight into the role of foxes during the Upper Paleolithic, espe-
cially regarding human subsistence, cultural expression, and ornament production.
Focusing on the latter, we cannot overlook the fact that foxes were one of many
species exploited for ornamentation in the Swabian Jura. Other taxa whose teeth were
used as ornaments include various herbivores like horses, reindeer and red deer, as well
as carnivores such as wolves, bears, and lions (Table S6). However, like the other species,
foxes had their own unique relationship with humans. As previously discussed, their
opportunistic nature likely drew foxes to human food waste, bringing both species closer
together in commensal relationships, which are attested to throughout the Upper Paleolithic.
Once connected, foxes played at least three key roles: as a resource, as a symbol, and as
a companion animal [
127
]. The latter two are less tangible in the archaeological record,
while study of material culture derived from foxes allows us to illuminate the economic
and symbolic significance of foxes in the Upper Paleolithic.
Although butchery traces on fox remains are rare, cut marks suggest foxes were likely
consumed as food in the Swabian Jura. Other marks indicate their use as a raw material,
particularly for their fur, a valuable resource for clothing and other sewn goods [
53
,
55
]. The
striking color and waterproof quality of their pelts likely played a role in the selection and
use of foxes for this resource [
121
]. While direct evidence is lacking, fox bones may have
also been used for tools (e.g., awls). Otherwise, fox tooth ornaments are the best example
of the use of foxes as a raw material.
The study of fox tooth ornaments helps us to unravel the symbolic importance of foxes
among Upper Paleolithic hunter-gatherers. This is because ornaments are social objects,
capable of reflecting otherwise unsaid aspects of the wearer, conveying messages on ethnic,
gender, social, and/or personal identity [
14
,
15
,
128
,
129
]. We can thus assume foxes, both as
a raw material and as a species, had a symbolic value in the lives of the hunter-gatherers
who wore their teeth.
Despite their clear symbolic importance, it is surprising that foxes are rarely depicted
in parietal or figurative art, other than a cave engraving at the Aurignacian site of Altxerri
B Cave in Spain [
130
]. In comparison, ungulates and larger predators, such as lions or
bears, are regularly depicted. This appears to change, however, during the Pre-Pottery
Neolithic and Natufian, when foxes increase in abundance, are commonly depicted in relief
(e.g., at Göbekli Tepe, Turkey) [
131
], and are buried alongside humans (e.g., at Uyun al-
Hammam, Jordan) [
132
]. A continued strengthening of the human–fox relationship is also
evidenced in the Neolithic [
131
,
133
], with the first possible evidence of fox domestication
in the Early-Middle Bronze Age [134].
To conclude, there is no doubt that the relationship between humans and foxes was
significant and started as early as the Initial Upper Paleolithic. The effect of this long-lasting
link still persists today in the folklore and mythology of many cultures, which depict foxes
as beautiful, clever, cunning, and magical beings [135,136].
Supplementary Materials:
The following supporting information can be downloaded at: https:
//www.mdpi.com/article/10.3390/quat6030050/s1, Figure S1: Overview of experimental technolog-
ical traces by drilling, Figure S2: Overview of experimental technological traces by scraping/grooving,
Figure S3: Overview of experimental technological traces by scraping or abrading and drilling,
Figure S4: Example of various ornament replicas and modes of attachment during the experi-
Quaternary 2023,6, 50 27 of 36
mentation, Figure S5: Pre- and post-use macrographs of perforations, Figure S6: Micrographs of
polish recorded on the perforations and corresponding location, Figure S7: Error margins for the
16 landmarks placed along the cervical outline, Table S1: Summary of the literature review on
Upper Paleolithic fox tooth ornaments, Table S2: List of archaeological fox canines included in the
geometric-morphometric analysis, Table S3: List of modern fox canines included in the geometric-
morphometric analysis, Table S4: Experimental fox canine replicas and related technological results,
Table S5: Experimental fox canine replicas with wear information and related recorded traces,
Table S6: Perforated animal teeth from the Swabian Jura (Hohle Fels, Geißenklösterle, Brillenhöhle,
Bockstein, Hohlenstein-Stadel, and Vogelherd). Ornaments unidentifiable to species and ornaments
from uncertain stratigraphic contexts are not included [137–248].
Author Contributions:
Conceptualization, F.V., M.J.M. and S.W.; Methodology, F.V. and M.-L.O.;
Formal analysis, F.V. and M.-L.O.; Investigation, F.V. and M.-L.O.; Data curation, F.V., M.J.M. and
M.-L.O.; Writing—original draft preparation, F.V., M.J.M. and M.-L.O.; Review and editing, F.V.,
M.J.M., N.J.C., M.-L.O. and S.W.; Visualization, F.V. and M.-L.O.; Project administration, F.V., M.J.M.
and S.W.; Funding acquisition, S.W.; Resources, N.J.C. and S.W. All authors have read and agreed to
the published version of the manuscript.
Funding:
This research was funded by the Senckenberg Center for Human Evolution and Palaeoen-
vironment at the University of Tübingen.
Data Availability Statement:
All relevant data are within the manuscript and its Supplementary
Materials files.
Acknowledgments:
We are grateful to Ran Barkai for inviting us to contribute to this Special Issue.
We thank Chris Baumann for providing the recent fox canines from his private collection, Alexandros
F. Karakostis and Gabriel Ferreira for their support during CT-scans, and Julia Zastrow for her help
during the GM-data collection. We would also like to thank Viviane Bolin and Nadine Rabovsky from
Museum Ulm, Fabian Haack from Landesmuseum Württemberg Stuttgart and the Archäologisches
Landesmuseum Baden-Württemberg, Zentrales Fundarchiv Rastatt for their assistance and for
providing access to the teeth under curation analyzed in this work. Thanks are also due to our
colleague Rudolf Walter for his active contribution to the experimental program. We are finally
grateful to the three anonymous reviewers for their constructive comments which contributed to
improve this manuscript.
Conflicts of Interest: The authors declare no conflict of interest.
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