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Martin Street, Hannes Napierala and Luc Janssens
Abstract
The association of dogs with early food producing
peoples has often been documented, in some cases as
far back as the terminal Pleistocene. Their presence in
Holocene hunter-gatherer contexts such as the Euro-
pean Mesolithic or the Japanese early Jomon has also
long been accepted. More recently, dogs or “dog-like”
canids have increasingly been described and in some ca-
ses discussed controversially from Upper Palaeolithic
contexts.
One of the first Palaeolithic sites at which dog re-
mains were recognized and generally accepted was the
Pleistocene late glacial site of Bonn-Oberkassel in the
northern Rhineland. Here fragmentary remains of a
canid recovered with two human skeletons are inter-
preted as a dog interred as part of an Upper Palaeolithic
human double burial.
Introduction
One hundred years ago, on the eve of the First World
War, two relatively complete human skeletons were
discovered during basalt quarrying at Oberkassel close
to Bonn in the northern German Rhineland. The dis-
covery of the human remains and an associated “hair
pin” was subsequently reported to the University of
Bonn, but by the time experts could examine the finds,
the bones were out of primary context and stored in a
workmen’s hut (Verworn et al. 1914). The finds were
described as coming from “a reddish cultural layer ly-
ing upon and in a sandy loam” (translations in quota-
tions MS).
From the first examination of the material, the en-
countered situation was interpreted as the remains of
a burial dating to the “Diluvial” Age and, indeed, was
already attributed by the first published description to
the “Renntierperiode” (reindeer period) (Verworn et al.
1914: 645). This first report also mentions animal bones
discovered in the “prolongation of the red-coloured
cultural layer towards the basalt rock face,” among
them “a right mandible of a wolf, a cave bear tooth and
bones of roe deer.”
An exhaustive report on the Oberkassel finds fol-
lowing the end of the First World War (Verworn et al.
1919) briefly adds further detail on the canid mandible,
described as being of “normal size with the teeth C, P4,
M1” and assigns other material (“…various vertebrae
and bone fragments of Canis lupus, the wolf”) to this
animal (Verworn et al. 1919: 9).
Following publication of the exhaustive monograph
describing the Oberkassel site, the Palaeolithic human
remains and associated “cultural objects” were stored
together in the archive of the Rheinisches Landesmuse-
um Bonn (today the LVR-LandesMuseum Bonn). Here
with some exceptions (e.g., Breuil and Kühn 1927; Ge-
rasimov 1964; Kahrs 1935; Mollison 1928; Szombathy
1920) the finds remained largely ignored during the
following 50 years. By contrast with this “cultural”
material the associated faunal remains and some geo-
logical samples at some point became partitioned into
different groups, and while some material, including
the canid mandible, remained associated with the hu-
man remains, other finds were stored separately in the
Geological Collections of the University of Bonn and
their provenience apparently forgotten.
Following this period of comparative disregard,
Bonn-Oberkassel became the subject of renewed in-
terest in the late 1970s when Gerhard Bosinski assig-
ned the site as a study object to Erwin Cziesla, then a
prehistory student at the University of Cologne (events
summarized in Cziesla 2013). This led to the “disco-
very” in the University of Bonn’s palaeolontological
and geological collections of faunal and other materi-
al from Oberkassel, which was thus reunited with the
finds stored at the Landesmuseum. A reappraisal of the
canid mandible (Fig. 1) carried out during the study
The late Palaeolithic dog from
Bonn-Oberkassel in context
Martin Street, Hannes Napierala and Luc Janssens2
led to its re-identification as a domesticated animal and
subsequent publication as a Palaeolithic dog by Günter
Nobis (Nobis 1979, 1981).
Context and chronology
The fresh awareness generated by these events led to
a reappraisal of the Oberkassel site as a whole, leading
to a new analysis of the human remains by Winfried
Henke (Henke 1984) and an overview presenting the-
se alongside a description of the dog mandible (Henke
1986; Nobis 1986). A fresh description of the archaeo-
logical context of the Oberkassel finds, including the
associated cultural objects and their interpretation,
which was planned to accompany the anthropological
and faunal reports, remained unfinished and the the-
me instead became the subject of a Master‘s Thesis by
Birgit Wüller which was subsequently presented at the
University of Cologne (Wüller 1992).
Well into the 1990s it was generally accepted that
the Oberkassel burial should be assigned to the Midd-
le Magdalenian (e.g., Bosinski et al. 1974; Veil 1978),
an age estimate based on the stylistic interpretation of
a carved bone or antler object as equivalent to French
Middle Magdalenian contours découpés (Verworn et al.
1919: 188; Bosinski 2014). The find depicts an animal
body, today generally accepted to be an elk (see Molli-
son 1928), although an appraisal as a wild pig was also
suggested (Kahrs 1935).
The Middle Magdalenian would have been earlier
than any of the German Magdalenian sites dated until
then (synthesis in Street, Baales and Weninger 1994),
and the Oberkassel site was discussed as representing
early recolonisation of more northern parts of Euro-
pe from the Franco-Cantabrian region following the
height of the last glaciation. Bone samples from the site
were therefore included in a programme to establish
the chronology of this recolonisation process and AMS
radiocarbon dated at the Oxford ORAU Laboratory
in 1994.
Results for both of the human skeletons and the
dog were younger than the stylistic estimate and da-
ted Oberkassel to the late glacial (Housley et al. 1997;
Street and Wüller 1998). They were subsequently con-
firmed by three further AMS dates for the dog obtained
at Kiel in cooperation with Norbert Benecke (Baales
and Street 1998; Street and Baales 1999) and by dating
results of the most recent resubmission of material to
Oxford (Higham et al. 2015; Street and Jöris 2015).
The absolute dates for Oberkassel clearly negate a
stylistic interpretation of the carving as Middle Mag-
dalenian. More convincing parallels in keeping with its
more recent age and interpretation as an elk are found
in younger contexts (Baales and Street 1998; Tinnes,
Giemsch and Schmitz 2014). More importantly, con-
firmation of an early Late Glacial Interstadial age for
Oberkassel (GI 1e) permits meaningful discussion of
the site in its correct climatic, environmental and archa-
eological context (Street and Jöris 2014, 2015).
The Bonn-Oberkassel dog
An unexpected side effect of sampling the Oberkassel
fauna for radiocarbon dating was the identification of
Fig. 1 Mandibula of the Bonn-Oberkassel dog seen in the medial/lingual aspect. (Photo: J. Vogel, LVR-LandesMuseum Bonn).
5 cm
The late Palaeolithic dog from Bonn-Oberkassel in context 3
2002
Original
label
Catalogue RLMB Catalogue
GeoI.-Pal.
Identification (*find already
described by Nobis (1986)
Staining
D 1001a
D 1001a
D 1001a
D 1001b
RLMB D001001, 01
RLMB D001001, 01
RLMB D001001, 01
RLMB D001001, 01
RLMB D001001, 01
RLMB D001001, 01
RLMB D001001, 01
RLMB D001001, 01
RLMB D001001, 01
RLMB D001001, 01
RLMB D001001, 21 OB 48
*right maxilla with incisors I
2
und I
3
*right mandible with C, P
4
, M
1
, M
2
*left mandible, processus coronoideus
*right maxillary molar M
1
*left maxillary premolar P
1
*left maxillary premolar P
3
*right mandibular incisor I
1
*right mandibular incisor I
2
*right mandibular incisor I
3
*left mandibular premolar P
2
os occipitalis
ochre
ochre
none (cleaned?)
ochre
ochre
ochre
ochre
ochre
none (cleaned?)
D 1001d
RLMB D001001, 10
RLMB D001001, 10
RLMB D001001, 10
RLMB D001001, 11
RLMB D001001, 21
RLMB D001001, 10
RLMB D001001, 10
RLMB D001001, 04
OB 21
OB 11
OB 45
OB 12, OB 13,
OB 14, OB 15
OB 59
OB 4
OB 6
left caudal epiphysis of an atlas CV 1
fragment of cervical vertebra CV 2, fits OB 45
fragment of cervical vertebra CV 2, fits OB 11
vertebrae (OB 12 - 14 lumbar vertebrae)
unfused cranial epiphysis of a lumbar vertebra, fits OB 14
anterior left rib (costa 2/3/4?)
left rib (costa 1?)
four small rib fragments
ochre
ochre
ochre
weak ochre
ochre
ochre
ochre
ochre
D 1001a
D 1001a
D 1001c3
RLMB D001001, 01
RLMB D001001, 14
RLMB D001001, 01
RLMB D001001, 09
RLMB D001001, 03
RLMB D001001, 03
RLMB D001001, 16
RLMB D001001, 08
OB 23
1
OB 3
2
OB 1
3
OB 31
4
OB 2
*right scapula
*epiphysis of a left humerus
*shaft of a right humerus
*shaft fragment of a left radius
*shaft fragment of a left ulna
*shaft fragment of a left ulna
*right ulna
*left metacarpus IV
ochre
ochre
ochre
ochre
ochre
ochre
ochre
Skull
Axial
Forelimb
Table 1 Bone and tooth fragments assigned to the Bonn-Oberkassel dog.
3
Two fragments (OB 1 and D 1001 c) refit to form a sec-
tion of the left ulna shaft (inventory RLMB D 001001, 03). They
are stored with a note bearing the words “Ulna-Diaphyse vom
Luchs” and were originally published as “probably lynx” (Nobis
1986).
4
A fragment (OB 31) of the right dog ulna (inventory
RLMB D00 1001, 16) was originally published as lynx (Nobis
1986).
1
A left proximal humerus epiphysis (inventory RLMB
D 001001, 14) was originally identified as “probably lynx” (No-
bis 1986).
2
A left radius fragment (inventory RLMB D 001001, 09) is
stored with a note dated to 27.2.1914 which describes the find as
„cervus capreolus L. ulna“. A more recent undated note corrects
the description to „Rad. Diaphys. dextr. Reh?“, an identification
which was subsequently published (Nobis 1986).
Martin Street, Hannes Napierala and Luc Janssens4
material previously described as roe deer and lynx as
further fragments of the dog (Street 1995), and a revisi-
on of the highly fragmented assemblage was initiated.
Several more small fragments of bone and tooth were
assigned to the dog, showing that appreciably more of
the skeleton was represented (Fig. 2) than previously
believed (Baales and Street 1998; Henke et al. 2006;
Street 2002, 2014). Overall, several elements of the
head, both of the forelimbs, vertebrae and ribs were
identified (Table 1). The fact that no elements are du-
plicated, along with the age estimates for the few spe-
cimens which provide such information, makes it very
likely that an entire single dog was originally interred
as part of the burial. Recent examination of the material
by the authors of the present paper provided new spe-
cific information on body type, age and pathology of
the animal (Janssens, Napierala and Street 2015).
Dogs in the Palaeolithic
Dogs are demonstrated convincingly by studies of re-
cent and ancient DNA (aDNA) to be a domesticated
form of wolf Canis lupus and to have no admixture
of other species such as jackal Canis aureus. Wolves
were the first species to become commensal with and
subsequently domesticated by humans, possibly due
to their pre-existing cognitive and social structures
(Scheider et al. 2011; Shipman 2009, 2012; Téglas et al.
2012). Major questions still exist as to the mechanisms
and date of their domestication, the answers to which
presuppose rigorous identifications and definitions of
the Pleistocene canids claimed to be involved in the
process (e.g., Boudadi-Maligne 2010; Pionnier-Capi-
tan 2010).
Until quite recently the oldest plausible claims for
early dogs appeared to be in association with people
living in western Eurasia at around the time of transi-
tion to a food producing lifestyle (Table 2). Dogs ap-
peared to be a quite regular phenomenon at Middle
and Near Eastern early Holocene and even terminal
Pleistocene sites with the earliest agricultural evidence
(Clutton-Brock 1979 [Jericho]; Davis and Valla 1978
[Ain Mallaha]; Dayan 1994 [Kebara]; Lawrence and
Reed 1983 [Jarmo]; Tchernov and Valla 1997 [Ain Mal-
laha, Hayonim]; Turnbull and Reed 1974 [Palegawra]).
The presence of dogs in Holocene hunter-gatherer
settlement contexts has also long been accepted (Ta-
ble 3), for example in the European Mesolithic (e.g.,
Benecke 1987; Chaix 2000; Clutton-Brock and Noe-
Nygaard 1990; Degerbøl 1961; Detry and Cardoso
2010; Henriksen et al. 1976; Street 1989, 1991; Vigne
and Marinval-Vigne 1988) or the northeastern Asian
Jomon (Shigehara and Hongo 2000).
Recently, morphologically distinct canids found at
a number of Late Pleistocene, mainly Upper Palaeo-
lithic contexts, have been described as “Palaeolithic
dogs” or interpreted as representing incipient wolf do-
mestication (e.g., Germonpré et al. 2009, 2012, 2013,
2014, 2015; Ovodov et al. 2011; Sablin and Khlopachev
2002), claims which have been met with scepticism by
a number of authors (Crockford and Kuzmin 2012;
Boudadi-Maligne and Escarguel 2014; Morey 2014;
Drake et al. 2015). Other morphological and isotope
studies discussing vanished specialized Pleistocene
wolf ecomorphs in terms of dietary adaptation (Flow-
er and Schreve 2014; Bocherens et al. 2014; Sansalone
et al. 2015) and their potential relationship to the ice
age megafauna and its extinction (Leonard et al. 2007)
might be relevant to this debate.
Genetic studies of dog DNA have also argued for
ancient domestication of wolves (e.g., Druzhkova et
al. 2013; Pang et al. 2009; Savolainen et al. 2002; Vilà et
al. 1997; vonHoldt et al. 2010, 2011; Wang et al. 2013),
although interpretations derived from the genetic
studies have varied widely regarding the age and geo-
graphical location of the origins of the domestic dog.
Analyses of aDNA show loss of some of the genetic
variation identified among Pleistocene canids, and the
absence in present-day wolves of specific haplotypes
of the dog genome suggests extinction of the progeni-
tor wolf population (Thalmann et al. 2013; Freedman
et al. 2014; Larson and Bradley 2014).
It has been suggested that answers to the question
of wolf domestication will ultimately only be found
by a synthesis of the genetic and the archaeological
evidence (Larson et al. 2012). Clearly, both the mor-
phological and genetic studies on the origins of the
domestic dog form part of a broader debate on the
motivation, mechanisms, timing and implications of
wolf domestication by humans and must clearly be
seen against the backdrop of the undoubtedly emotive
role played by “man’s best friend” in contemporary
western society.
It is perhaps representative of the ongoing debate
on wolf domestication that the authors of the present
paper also agree to differ in their individual assessment
of current evidence for the earliest reported associa-
tions of humans and canids.
However, in the case of claims for more recent Up-
per Palaeolithic dogs there does appear to be a gro-
wing consensus of acceptance (e.g., Pionnier-Capitan
2010; Pionnier-Capitan et al. 2011; Morey 2014), one
which is shared by the present authors.
The late Palaeolithic dog from Bonn-Oberkassel in context 5
Bonn-Oberkassel was indeed one of the very first
Upper Palaeolithic localities at which dog remains
were first recognized on morphological grounds (No-
bis 1979, 1981, 1986; Benecke 1987; Street 2002; Henke
et al. 2006; Cziesla 2013). Subsequent analysis of the
mtDNA of the Bonn-Oberkassel canid has confirmed
the specimen unambiguously as a dog, assigning it to
clade C of the modern dog genome (Thalmann et al.
2013), and today the Bonn-Oberkassel dog can be seen
to form part of a small group of late Upper Palaeoli-
thic dogs from a number of Magdalenian and slightly
younger sites across Western Europe (Table 4).
The valid status of these late Upper Palaeolithic dogs
has been accepted by a number of researchers (sum-
mary in Morey 2014) and need not be repeated here.
It is however worth reviewing the occurrence of these
animals against their regional and supra-regional archa-
eological context, something which may be relatively
ignored by studies arising from zoomorphological or
palaeogenetic interests.
A number of the sites have produced canid remains
which are morphologically not clearly distinct from
wolves and are regarded at most as transitional forms,
possibly representing an initial phase of taming. Among
these are remains of large canids described by Rudolf
Musil from the Magdalenian sites Kniegrotte, Teufels-
brücke and Oelknitz (Musil 1974, 1980, 1985) which
together were considered by him as probably derived
from domestic dogs (Musil 2000). An early comprehen-
sive review of claims for Palaeolithic dogs by Norbert
Benecke pointed out that the Kniegrotte canid show-
ed great similarities with zoo-kept wolves and sugge-
sted that such specimens might reflect wolves kept by
humans at an initial stage of domestication (Benecke
1987). Interestingly the majority of reliable 14C dates
for the eastern German Magdalenian sites with “incipi-
ent dogs” places them early in the phase of late glacial
reoccupation of Central Europe (Housley et al. 1997),
well before the sudden warming episode of Greenland
Interstadial II (Street and Jöris 2015 in this volume).
Most of the other finds under discussion are less ambi-
guous and more clearly identifiable as dogs and often also
associated with somewhat younger absolute dates (Table 4).
A single record from Eralla in Spain represents the
only presence of dog south of the Pyrenees. It is un-
fortunate that the age of the specimen is unclear, being
estimated at either ~20,000–19,000 or ~14,500 years old
(Altuna and Mariezkurrena 1985; Vigne 2005; García-
Moncó 2005). The younger estimate would seem to be
more compatible with the other dates for Palaeolithic
Fig. 2 The identified bones of the Bonn-Oberkassel dog skeleton. Elements of the skull are at top left; bones of
the forelimb are at centre and below; the axial skeleton (ribs, vertebrae) extend across the top of the photograph.
(Photo: J. Vogel, LVR-LandesMuseum Bonn).
10 cm
Country Site Calendar age
BP
Elements Description Reference
Kazakhstan Botai 5,550–5.050 Several bones Benecke and von den Driesch 2003
~5,650 Complete skeleton Olsen 2000
Turkmenistan Jeitun 8,200–7,700 Multiple elements Harris et al. 1993
Turkey Çayönü 9,200–9,100 Complete skeleton Özdoğan 1999
Cafer Höyük 9,500–8,300 Complete skull,
14 other elements
Digested bones and gnawing marks indicate
that dogs were living in the village.
Cauvin et al.1999;
Helmer 2008
Cyprus Klimonas 11,120–10,615 One phalanx consistent
with a very young dog
Gnaw marks on the ungulate bones at the site
confirms that these dogs lived together with humans.
Vigne 2011a;
Vigne et al. 2011
Shillourokambos 12,400–12,300 Multiple elements The dog bones are associated with the earliest layers
of the site (early phase A) and clearly represent very
small dogs.
Guilaine et al. 2011;
Vigne 2011b; Vigne et al. 2011
Israel Ain Mallaha ~11,500 One juvenile (puppy)
skeleton, one adult and
one partial mandible
Co-burials with humans. The puppy is buried with a
woman whose hand is resting on the dog. The second
complete individual is an adult positioned underneath
a human burial.
Davis and Valla 1978;
Tchernov and Valla 1997
Hayonim Cave &
Terrace
12,000–11,000 Complete skeleton Tchernov and Valla 1997
Kebara Cave 12,500–12,000 Mandible Dating of the site and determination of the mandible
are uncertain. Contemporary remains of elements
more confidently assigned to domestic dogs are
found nearby.
Dayan 1994
Table 2 (Continuation see next page) Western Eurasian dogs associated with early food producing and transitional contexts.
Country Site Calendar age
BP
Elements Description Reference
Syria Aswad-Damascus 10,200–9,400 A few tens of elements Helmer and Gourichon 2008;
Stordeur et al. 2010
Tell Mureybet 11,500–11,300 Skull and left and
right mandibles
Morphological features and metrical comparison
with Canis lupus pallipes confirms the domestic
status of this specimen.
Évin and Stordeur 2008;
Gourichon and Helmer 2008
Iraq Jarmo 9,000–8,500 53 cranial and mandibular
fragments, 18 of which
positively identifiable as dog
The dog bones were compared with remains from
the local wolf, Canis lupus pallipes.
Lawrence and Reed 1983
Palegawra ~13,000 Mandible Tooth size reduction and crowding in a smaller jaw. Turnbull and Reed 1974
Table 3 (Continuation see next page) Northern Eurasian early Holocene dogs from hunter-gatherer contexts and / or burials.
All calibrations of
14
C dates were performed in August 2015 using the online OxCal programme https://c14.arch.ox.ac.uk/oxcal/OxCal.html
Country Site Lab. Code
14
C Age BP Calibrated age
BP
Dated material
Russia, N. Siberian
Islands
Zhokhov 8,480–8,175
Russia (Cis-Baikal) Shamanka II E2008.175 OxA-20561 6,430 ± 35 7,425–7,281
Lokomotiv wolf
H2003.704
GIN-8841a
TO-11558
7,230 ± 40
7,320 ± 70
8,161–7,971
8,317–8,002
Latvia Zvejnieki 7,434–7,200
Sweden Skateholm ~7,000
Denmark Svaerdborg 8,400–7,800
Germany Bedburg KN-3998
KN-3999
9,600 ± 100
9,780 ± 100
11,210–10,672
11,600–10,773
Plant remains from layer containing bones
(range, 9 dates) COL-2669.2.1
COL-2672.1.1
10,006 ± 43 -
10,115 ± 48
12,000 –11,280 Bos cranium, humerus, metacarpus (8 dates)
UK Star Carr KIA-37034
OxA-V-994-33
9,342 ± 41
9,680 ± 55
10,685–10,428
11,224–10,789
Seamer Carr OxA-1030 9,940 ± 100 11,807–11,196
Italy Romanelli Beta ???
Beta ???
9,670 ± 40
9,860 ± 50
11,205–10,798
11,393–11,198
Tibia (sample PIC 3)
Metapodial (sample PIC 2)
France Noyen-sur-Seine ~10,200
Saint-Thibaud-de-Couz OxA-4405 10,050 ± 100 11,980–11,260
Portugal Cabeco da Arruda Beta-152956 7,070 ± 40 7,973–7,801 Ribs
Country Site Description References
Russia, N. Siberian
Islands
Zhokhov 2 mandibles, maxilla, canine, radius, tibia Pitul‘ko/Kasparov 1996
Russia (Cis-Baikal) Shamanka II
E2008.175
Neolithic human burial with dog skeleton. Both dog and wolf also at several nearby sites Losey et al. 2011; 2013
Lokomotiv wolf
H2003.704
Neolithic? „tundra wolf“ buried with human crania. Both dog and wolf also found at several nearby sites Bazaliiskiy/Savelyev 2003; Losey et al. 2011; 2013
Latvia Zvejnieki Complete skeletons. Burials. Löugas 2006
Sweden Skateholm Burials of complete skeletons. 10 canine graves, some with ochre. Larsson 1990
Denmark Svaerdborg 14 fragments of canid bones suggested to be three types of dogs: large Greenland-like dogs, medium- and
small-sized dogs like Lappish Spitz
Henriksen et al. 1976
Germany Bedburg Cranium, humerus and tibia. Street 1991
(range, 9 dates) Street/Gehlen in prep.
UK Star Carr Subadult skull fragment; single tooth, femur, tibia. Bones identified as both dog and wolf are found at this site. Clutton-Brock/Noe-Nygaard 1990; Schulting/Richards 2009
Seamer Carr 6 vertebrae Clutton-Brock/Noe-Nygaard 1990
Italy Romanelli Verginelli et al. 2005
France Noyen-sur-Seine 2 complete skulls, as big as wolf skulls, but with domesticated features; suggested by Vigne as likely derived
from local population of wolves. Several post-cranial bones.
Vigne/Marinval-Vigne 1988; Mordant/Mordant 1992
Saint-Thibaud-
de-Couz
Skull, right mandible, atlas, axis, some teeth, left humerus. Chaix 2000; Pionnier-Capitan et al. 2011
Portugal Cabeco da
Arruda
Near-complete dog skeleton burial. Detry/Cardoso 2010
Country Site Lab. Code
14
C Age BP Calibrated age
BP
Dated material Dog remains References
Germany Kartstein OxA-9032
OxA-9031
9,995 ± 65
10,220 ± 75
11,751 –11,256
12,378 – 11,615
bone, bird
bone, reindeer
Phalanges Baales 1996; Street et al. 1994
Bettenroder Berg ~13,000 Right juvenile mandible with pd3, d4 Staesche 1994
Bonn-Oberkassel
KIA-4163
KIA-4161
KIA-4162
OxA-4793
OxA-29868
11,620 ± 60
12,110 ± 45
12,210 ± 60
12,270 ± 100
12,390 ± 55
13,569 – 13,319
14,122 – 13,799
14,337 – 13,861
14,773 – 13,860
14,809 – 14,140
ulna dext., dog
maxilla dext., dog
humerus dext., dog
ulna sin., dog
ulna sin., dog
Mandible, fragmentary maxilla, Street 2002, 2014; Street/Jöris 2015
Oelknitz (range,
12 dates)
OxA-5711
OxA-5716
12,050 ± 110 –
12,790 ± 110
15,673 – 13,604 bone, reindeer and horse Phalanges, metapodials and part of distal humerus and tibia Musil 1985; 2000; Housley et al. 1997
Teufelsbrücke
(range, 5 dates)
OxA-5726
OxA-5723
12,640 ± 130 –
13,080 ± 140
16,085 – 14,279 bone, reindeer, horse and ibex? Proximal metapodial fragment and first phalange Musil 1980; 2000; Housley et al. 1997
Kniegrotte
(range, 8 dates)
OxA-4853
OxA-4832
13,520 ± 130 –
13,310 ± 110
16,772 – 15,277 bone, reindeer, horse, saiga and fox Partial maxillary fragment with teeth Musil 1974; 2000; Housley et al. 1997
Switzerland Kesslerloch Cave KIA-33350 12,225 ± 45 14,287 – 13,976 maxilla dext., dog Partial skull fragment:
maxilla / zygo / palatine and a few teeth
Napierala/Uerpmann 2012
Champréveyres
(range, 13 dates)
UZ-2287
UZ-2285
12,510 ± 145 –
13,050 ± 155
16,082 – 14,137 charcoal Metatarsal and two teeth, second phalange Morel/Müller 1997; Müller 2004; Pionnier-Capitan et al. 2011
Monruz (range,
10 dates)
ETH-6414
ETH-6413
12,840 ± 120 –
13,330 ± 110
16,345 – 14,952 charcoal Deciduous dentition (possibly dog) Müller 2006; 2013
France Pont d‘Ambon GifA-99102 10,730 ± 100 12,806 – 12,428 bone, dog 39 skull, limb, mandible, vertebrae and tooth fragments Célérier/Delpech 1978;
Célérier/Tisnerat/Valladas 1999; Pionnier-Capitan et al. 2011
Le Closeau,
Locus 46 (range,
5 dates)
AA-41882
AA-41881
12,248 ± 66 –
12,423 ± 67
14,944 – 13,953 bone, pig, red deer, horse and lion 7 fragments: mandible, metapodials, phalanges Bémilli 2000; Pionnier-Capitan et al. 2011
Abri Morin OxA-23628
OxA-23627
12,450 ± 55
12,540 ± 55
14,975 – 14,227
15,127 – 14,429
mandibular M1, dog
maxillary I3, dog
32 remains of a small canid Boudadi-Maligne et al. 2012
Montespan - - 15,500 – 13,500 „bad collagen“ 1 atlas, 1 femur, 1 baculum Pionnier-Capitan et al. 2011
Spain Erralla ~20,000 – 19,000
or ~14,500
bones 1 humerus of a small dog Altuna/Mariezkurrena 1985; Vigne 2005; García-Moncó 2005
Table 4 (Continuation see next page) Late Pleistocene dogs and wolves claimed to be transitional to domestication from
the Western and Central European late Upper Palaeolithic. All calibrations of
14
C dates were performed in August 2015
using the online OxCal programme https://c14.arch.ox.ac.uk/oxcal/OxCal.html
Country Site Lab. Code 14C Age BP Calibrated age
BP
Dated material Dog remains References
Germany Kartstein OxA-9032
OxA-9031
9,995 ± 65
10,220 ± 75
11,751 – 11,256
12,378 – 11,615
bone, bird
bone, reindeer
Phalanges Baales 1996; Street et al. 1994
Bettenroder Berg ~13,000 Right juvenile mandible with pd3, d4 Staesche 1994
KIA-4163 11,620 ± 60 13,569 – 13,319 ulna dext., dog Mandible, fragmentary maxilla, Street 2002, 2014; Street/Jöris 2015
KIA-4161 12,110 ± 45 14,122 – 13,799 maxilla dext., dog vertebrae, ribs, scapula,
Bonn-Oberkassel KIA-4162 12,210 ± 60 14,337 – 13,861 humerus dext., dog humerus, radius, ulna
OxA-4793 12,270 ± 100 14,773 – 13,860 ulna sin., dog metapodial
OxA-29868 12,390 ± 55 14,809 – 14,140 ulna sin., dog
Oelknitz (range,
12 dates)
OxA-5711
OxA-5716
12,050 ± 110 –
12,790 ± 110
15,673 – 13,604 bone, reindeer and horser Phalanges, metapodials and part of distal humerus and tibia Musil 1985; 2000; Housley et al. 1997
Teufelsbrücke
(range, 5 dates)
OxA-5726
OxA-5723
12,640 ± 130 –
13,080 ± 140
16,085 – 14,279 bone, reindeer, horse and ibex? Proximal metapodial fragment and first phalange Musil 1980; 2000; Housley et al. 1997
Kniegrotte
(range, 8 dates)
OxA-4853
OxA-4832
13,520 ± 130 –
13,310 ± 110
16,772 – 15,277 bone, reindeer, horse, saiga and fox Partial maxillary fragment with teeth Musil 1974; 2000; Housley et al. 1997
Switzerland Kesslerloch Cave KIA-33350 12,225 ± 45 14,287 – 13,976 maxilla dext., dog Partial skull fragment:
maxilla / zygo / palatine and a few teeth
Napierala/Uerpmann 2012
Champréveyres
(range, 13 dates)
UZ-2287
UZ-2285
12,510 ± 145 –
13,050 ± 155
16,082 – 14,137 charcoal Metatarsal and two teeth, second phalange Morel/Müller 1997; Müller 2004; Pionnier-Capitan et al. 2011
Monruz (range,
10 dates)
ETH-6414
ETH-6413
12,840 ± 120 –
13,330 ± 110
16,345 – 14,952 charcoal Deciduous dentition (possibly dog) Müller 2006; 2013
France Pont d‘Ambon GifA-99102 10,730 ± 100 12,806 – 12,428 bone, dog 39 skull, limb, mandible, vertebrae and tooth fragments Célérier/Delpech 1978;
Célérier/Tisnerat/Valladas 1999; Pionnier-Capitan et al. 2011
Le Closeau,
Locus 46 (range,
5 dates)
AA-41882
AA-41881
12,248 ± 66 –
12,423 ± 67
14,944 – 13,953 bone, pig, red deer, horse and lion 7 fragments: mandible, metapodials, phalanges Bémilli 2000; Pionnier-Capitan et al. 2011
Abri Morin OxA-23628
OxA-23627
12,450 ± 55
12,540 ± 55
14,975 – 14,227
15,127 – 14,429
mandibular M1, dog
maxillary I3, dog
32 remains of a small canid Boudadi-Maligne et al. 2012
Montespan - - 15,500 – 13,500 „bad collagen“ 1 atlas, 1 femur, 1 baculum Pionnier-Capitan et al. 2011
Spain Erralla ~20,000 – 19,000
or ~14,500
bones 1 humerus of a small dog Altuna/Mariezkurrena 1985; Vigne 2005; García-Moncó 2005
Country Site Dog remains References
Germany Kartstein Phalanges Baales 1996; Street et al. 1994
Bettenroder Berg Right juvenile mandible with pd3, d4 Staesche 1994
Mandible, fragmentary maxilla, Street 2002, 2014; Street/Jöris 2015
vertebrae, ribs, scapula,
Bonn-Oberkassel humerus, radius, ulna
metapodial
Oelknitz (range,
12 dates)
Phalanges, metapodials and part of distal humerus and tibia Musil 1985; 2000; Housley et al. 1997
Teufelsbrücke
(range, 5 dates)
Proximal metapodial fragment and first phalange Musil 1980; 2000; Housley et al. 1997
Kniegrotte
(range, 8 dates)
Partial maxillary fragment with teeth Musil 1974; 2000; Housley et al. 1997
Switzerland Kesslerloch Cave Partial skull fragment:
maxilla / zygo / palatine and a few teeth
Napierala/Uerpmann 2012
Champréveyres
(range, 13 dates)
Metatarsal and two teeth, second phalange Morel/Müller 1997; Müller 2004; Pionnier-Capitan et al. 2011
Monruz (range,
10 dates)
Deciduous dentition (possibly dog) Müller 2006; 2013
France Pont d‘Ambon 39 skull, limb, mandible, vertebrae and tooth fragments Célérier/Delpech 1978;
Célérier/Tisnerat/Valladas 1999; Pionnier-Capitan et al. 2011
Le Closeau,
Locus 46 (range,
5 dates)
7 fragments: mandible, metapodials, phalanges Bémilli 2000; Pionnier-Capitan et al. 2011
Abri Morin 32 remains of a small canid Boudadi-Maligne et al. 2012
Montespan 1 atlas, 1 femur, 1 baculum Pionnier-Capitan et al. 2011
Spain Erralla 1 humerus of a small dog Altuna/Mariezkurrena 1985; Vigne 2005; García-Moncó 2005
Country Site Lab. Code
14
C Age BP Calibrated age
BP
Dated material Dog remains References
Germany Kartstein OxA-9032
OxA-9031
9,995 ± 65
10,220 ± 75
11,751 –11,256
12,378 – 11,615
bone, bird
bone, reindeer
Phalanges Baales 1996; Street et al. 1994
Bettenroder Berg ~13,000 Right juvenile mandible with pd3, d4 Staesche 1994
Bonn-Oberkassel
KIA-4163
KIA-4161
KIA-4162
OxA-4793
OxA-29868
11,620 ± 60
12,110 ± 45
12,210 ± 60
12,270 ± 100
12,390 ± 55
13,569 – 13,319
14,122 – 13,799
14,337 – 13,861
14,773 – 13,860
14,809 – 14,140
ulna dext., dog
maxilla dext., dog
humerus dext., dog
ulna sin., dog
ulna sin., dog
Mandible, fragmentary maxilla,
Street 2002, 2014; Street/Jöris 2015
Oelknitz (range,
12 dates)
OxA-5711
OxA-5716
12,050 ± 110 –
12,790 ± 110
15,673 – 13,604 bone, reindeer and horse Phalanges, metapodials and part of distal humerus and tibia Musil 1985; 2000; Housley et al. 1997
Teufelsbrücke
(range, 5 dates)
OxA-5726
OxA-5723
12,640 ± 130 –
13,080 ± 140
16,085 – 14,279 bone, reindeer, horse and ibex? Proximal metapodial fragment and first phalange Musil 1980; 2000; Housley et al. 1997
Kniegrotte
(range, 8 dates)
OxA-4853
OxA-4832
13,520 ± 130 –
13,310 ± 110
16,772 – 15,277 bone, reindeer, horse, saiga and fox Partial maxillary fragment with teeth Musil 1974; 2000; Housley et al. 1997
Switzerland Kesslerloch Cave KIA-33350 12,225 ± 45 14,287 – 13,976 maxilla dext., dog Partial skull fragment:
maxilla / zygo / palatine and a few teeth
Napierala/Uerpmann 2012
Champréveyres
(range, 13 dates)
UZ-2287
UZ-2285
12,510 ± 145 –
13,050 ± 155
16,082 – 14,137 charcoal Metatarsal and two teeth, second phalange Morel/Müller 1997; Müller 2004; Pionnier-Capitan et al. 2011
Monruz (range,
10 dates)
ETH-6414
ETH-6413
12,840 ± 120 –
13,330 ± 110
16,345 – 14,952 charcoal Deciduous dentition (possibly dog) Müller 2006; 2013
France Pont d‘Ambon GifA-99102 10,730 ± 100 12,806 – 12,428 bone, dog 39 skull, limb, mandible, vertebrae and tooth fragments Célérier/Delpech 1978;
Célérier/Tisnerat/Valladas 1999; Pionnier-Capitan et al. 2011
Le Closeau,
Locus 46 (range,
5 dates)
AA-41882
AA-41881
12,248 ± 66 –
12,423 ± 67
14,944 – 13,953 bone, pig, red deer, horse and lion 7 fragments: mandible, metapodials, phalanges Bémilli 2000; Pionnier-Capitan et al. 2011
Abri Morin OxA-23628
OxA-23627
12,450 ± 55
12,540 ± 55
14,975 – 14,227
15,127 – 14,429
mandibular M1, dog
maxillary I3, dog
32 remains of a small canid Boudadi-Maligne et al. 2012
Montespan - - 15,500 – 13,500 „bad collagen“ 1 atlas, 1 femur, 1 baculum Pionnier-Capitan et al. 2011
Spain Erralla ~20,000 – 19,000
or ~14,500
bones 1 humerus of a small dog Altuna/Mariezkurrena 1985; Vigne 2005; García-Moncó 2005
Country Site Dog remains References
Germany Kartstein Phalanges Baales 1996; Street et al. 1994
Bettenroder Berg Right juvenile mandible with pd3, d4 Staesche 1994
Bonn-Oberkassel
Mandible, fragmentary maxilla,
vertebrae, ribs, scapula,
humerus, radius, ulna
metapodial
Street 2002, 2014; Street/Jöris 2015
Oelknitz (range,
12 dates)
Phalanges, metapodials and part of distal humerus and tibia Musil 1985; 2000; Housley et al. 1997
Teufelsbrücke
(range, 5 dates)
Proximal metapodial fragment and first phalange Musil 1980; 2000; Housley et al. 1997
Kniegrotte
(range, 8 dates)
Partial maxillary fragment with teeth Musil 1974; 2000; Housley et al. 1997
Switzerland Kesslerloch Cave Partial skull fragment: maxilla / zygo / palatine and a few teeth Napierala/Uerpmann 2012
Champréveyres
(range, 13 dates)
Metatarsal and two teeth, second phalange Morel/Müller 1997; Müller 2004; Pionnier-Capitan et al. 2011
Monruz (range,
10 dates)
Deciduous dentition (possibly dog) Müller 2006; 2013
France Pont d‘Ambon 39 skull, limb, mandible, vertebrae and tooth fragments Célérier/Delpech 1978;
Célérier/Tisnerat/Valladas 1999; Pionnier-Capitan et al. 2011
Le Closeau,
Locus 46 (range,
5 dates)
7 fragments: mandible, metapodials, phalanges Bémilli 2000; Pionnier-Capitan et al. 2011
Abri Morin 32 remains of a small canid Boudadi-Maligne et al. 2012
Montespan 1 atlas, 1 femur, 1 baculum Pionnier-Capitan et al. 2011
Spain Erralla 1 humerus of a small dog Altuna/Mariezkurrena 1985; Vigne 2005; García-Moncó 2005
Martin Street, Hannes Napierala and Luc Janssens12
dogs younger than the Late Glacial Maximum. However,
an older age cannot be ruled out should the process of
wolf domestication have its origins in the southwest Eu-
ropean refugial heartland of the Magdalenian culture, in
which case Eralla might indeed potentially represent the
oldest known individual of the lineage leading to modern
dogs.
In France the occurrence of Final Pleistocene canids
accepted as dogs extends over a large area from the Py-
renees to the Paris Basin (Boudadi-Maligne et al. 2012;
Pionnier-Capitan et al. 2011). The archaeological context
of the finds ranges from the Middle and Late Magdaleni-
an (Pionnier-Capitan et al. 2011 [Montespan]; Boudadi-
Maligne et al. 2012 [Le Morin]) to the succeeding Azilian
(Bémilli 2000 [Le Closeau]; Célérier and Delpech 1978;
Célérier, Tisnerat and Valladas 1999 [Pont d’Ambon]).
The small number of finds shows no fine patterning bet-
ween region and chronology.
Certain or potential finds of late Pleistocene dogs are
reported from the Swiss Magdalenian (Napierala and
Uerpmann 2012 [Kesserloch]; Morel and Müller 1997;
Müller 2004 [Champréveyres], Müller 2006, 2013 [Mon-
ruz]). While the latter two open-air sites are well dated
to the final part of the stadial preceding Greenland In-
terstadial GI II, the direct date on the Kesslerloch dog is
the same as those for Oberkassel which would suggest a
younger Magdalenian context at this site than at Champ-
réveyres and Monruz.
Other than at Oberkassel, probable or possible dog
remains have been reported from two other, somewhat
younger Late Glacial sites in Germany (Staesche 1994
[Bettenroder Berg]; Baales 1996 [Kartstein]). Both sites
are rockshelters, the former dated by stratigraphy to the
final part of the Allerød Interstadial (~GI 1 a) and the lat-
ter to the Younger Dryas Stadial GS I.
In summary, the chronological and geographical pat-
terning of domestic dog occurrence shows that dogs were
present in several parts of northwestern Eurasia during
the late Pleistocene following the Late Glacial Maximum.
This suggests, together with genetic information obtained
from recent dogs and from aDNA, that the process of
wolf domestication which resulted in lineages surviving
in the dog genome until the present day probably began
in Western Europe in the context of the Magdalenian and
that it had already produced morphologically and geneti-
cally “modern dogs” by around 14,500 years ago.
The dog in the grave
The importance of dogs for early human societies
perhaps finds its clearest expression in their interment
in graves, usually together with humans, but sometimes
as specific dog burials (Larsson 1989, 1990, 1994; Losey
et al. 2011, 2013). Ritual dog burials are known from
several past time periods but Oberkassel is, to the best
of our knowledge, the oldest directly dated case of this
practise.
The Oberkassel canid is altered sufficiently anatomi-
cally from a wolf that it in itself presents a strong argu-
ment for the domestication process, probably due to
conscious or unconscious selection for certain qualities
by humans. However, what might these qualities have
been? What was the role of dogs in this late Palaeolithic
society in general and how did humans regard and in-
teract with this dog specifically?
Numerous practical functions have been described
as contributing to the usefulness of early dogs for hu-
mans, perhaps foremost among them as hunting aids
(e.g., Ruusila and Pesonen 2004; Lupo 2011; Koster
and Tankersley 2012). Some authors indeed suggest
that the efficiency of this combination may have had
remarkably far reaching implications for the broader
ecology (Fiedel 2005; Shipman 2014, 2015). Other dis-
cussed roles are as transport or guard dog (e.g., dis-
cussion in Germonpré et al. 2012), food source (White
1955), waste disposal (Street 1989) or perhaps simply
as companions. The pathology of the Bonn-Oberkas-
sel dog suggests that it died young after debilitating
illness (Janssens, Napierala and Street 2015 in this vo-
lume) and will therefore not have been in a position to
take on any of these potential roles. The implication
is rather that the dog needed to be fed by humans to
survive.
Does this imply a form of ownership or at least re-
sponsibility? While a concept of property is perhaps
not appropriate in the context of what are generally
believed to be broadly “egalitarian” societies, it is
perhaps ineviTable that dogs generally will have had
a structured relationship to people within any exis-
ting social hierarchies and that the Oberkassel dog in
particular will have had a closer relationship to cer-
tain people who must clearly have empathized with
the animal.
The Oberkassel dog skeletal remains were stained
with red ochre, and it is proposed that they were re-
covered from a context so close to the buried human
skeletons interpreted as a double inhumation that the
dog too formed part of the Oberkassel burial. This is
suggestive of the existence of emotional or symbolic
ties between humans and their dogs, perhaps generally
or at least specifically between this dog and these bu-
ried humans. Even though no answers may be possib-
le, it is perhaps legitimate to speculate what these ties
The late Palaeolithic dog from Bonn-Oberkassel in context 13
may have been and why the animal was buried with
the humans?
It is not known whether the dog was intentionally
killed to be buried together with the humans, the cause
of whose death is also unknown. If the dog had still
been reliant upon intense care for survival (Janssens,
Napierala and Street 2015 in this volume), the removal
of this by the death of its carers would in itself have
been a death sentence. This would of course imply that
the buried humans were the dog’s carers, for which we
have no evidence. A natural death (sickness, accident,
starvation, etc.) of all three individuals must also be
considered a possibility.
Did the dog perhaps take on a function in the after
world as a symbolic companion, for example as a hun-
ting partner or a guard, or indeed as a spirit guide into
the after world? The fact that the Oberkassel dog must
have suffered repeated episodes of illness might suggest
that some of these roles would have been “inappropria-
te.” On the other hand, an animal potentially subject to
fits (Janssens, Napierala and Street 2015 in this volume)
might have been regarded as particularly suitable for
some “supernatural” role.
The preceding thoughts on the motivation for inclu-
ding a dog in the Oberkassel double inhumation remain
firmly in the realm of speculation. However, the well-
documented presence of dogs in burial contexts shows
that they played a role in human life transcending the
purely utilitarian (Byrd et al. 2013; Morey 2006, 2010;
Losey et al. 2013). The fact that intentional burial ext-
ends to canid species other than domestic dogs (Hale
and Salls 2000; Bazaliiskiy and Savelyev 2003; Losey et
al. 2011; Prates 2014) suggests that a heterogeneous set
of motivations may be involved in the inclusion of the-
se animals in human burials, not all of them necessarily
equivalent to or even directly related to the processes
which had led to the domestication of wolves and the
creation of the first domestic animal.
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Martin Street, Hannes Napierala and Luc Janssens22
Contact
MONREPOS Archaeological Research Centre and Museum for Human Behavioural Evolution
Schloss Monrepos, Neuwied, D-56567, Germany
street@rgzm.de
Altrathausstrasse 5, 72511 Bingen, Germany
Department of Archaeology
Einsteinweg 2, 2333 CC, Leiden, The Netherlands
coati1@icloud.com
