Sorting the Sheep from the Goats: Morphological Distinctions between the Mandibles and Mandibular Teeth of AdultOvis and Capra

Abstract

Diagnostic criteria are described for the taxonomic discrimination of adult mandibles of sheep (Ovis aries) and goat (Capra hircus). These criteria, based on morphological distinctions in the mandible, permanent premolars (P3–P4) and molars (M1–M3), have been successfully tested on a large and diverse body of modern reference material. In conjunction with published work on deciduous premolars, the new criteria enable complete and fragmentary mandibles of young, adult and elderly individuals to be identified to sheep or goat with acceptable levels of accuracy.

Full text

Journal of Archaeological Science (2002) 29, 545–553
doi:10.1006/jasc.2001.0777, available online at http://www.idealibrary.com on
Sorting the Sheep from the Goats: Morphological Distinctions
between the Mandibles and Mandibular Teeth of Adult Ovis
and Capra
P. Halstead and P. Collins
Department of Archaeology and Prehistory, University of Sheffield, Northgate House, West Street, Sheffield
S1 4ET, U.K.
V. Isaakidou
Institute of Archaeology, University College London, 31–34 Gordon Square, London WC1H 0PY, U.K.
(Received 24 May 2001, revised manuscript accepted 16 November 2001)
Diagnostic criteria are described for the taxonomic discrimination of adult mandibles of sheep (Ovis aries) and goat
(Capra hircus). These criteria, based on morphological distinctions in the mandible, permanent premolars (P3–P4) and
molars (M1–M3), have been successfully tested on a large and diverse body of modern reference material. In
conjunction with published work on deciduous premolars, the new criteria enable complete and fragmentary mandibles
of young, adult and elderly individuals to be identified to sheep or goat with acceptable levels of accuracy.
2002 Elsevier Science Ltd. All rights reserved.
Keywords: ARCHAEOZOOLOGY, MANDIBLE, TEETH, SHEEP, OVIS, GOAT, CAPRA, IDENTIFICATION.
Introduction
Although sheep and goat may be herded together
and raised for the same goals, these two species
exhibit significant differences in environmental
tolerances, feeding preferences, ease of control, repro-
ductive characteristics, carcase quality, and range of
secondary products (e.g., Payne, 1973;Redding, 1981,
1984). Archaeozoological analyses of mortality pat-
terns for the combined category ‘‘sheep/goat’’ thus risk
masking important contrasts in management between
these species or, worse still, creating an illusory com-
posite picture which is valid for neither species. These
problems are particularly acute in areas, such as the
Mediterranean and Near East, where both sheep and
goat have long been major constituents of livestock
populations. Young ovicaprid mandibles, containing
deciduous premolars (dp3, dp4) and/or very lightly
worn first or second molars (M1, M2) and representing
animals up to c. 2 years of age, can be speciated
following Payne (1985). The analysis of adult mortality
patterns, however, has largely been dependent on the
problematic evidence of postcranial epiphyseal fusion;
in addition to the usual limitations of fusion data (e.g.,
Payne, 1972;Watson, 1978), the identification of post-
cranial material to sheep or goat tends to be much
more difficult for juvenile than for adult specimens
(Boessneck, Mu¨ller & Teichert, 1964). Recently,
Helmer (2000) has published diagnostic criteria for the
permanent premolars, P3 and P4, of sheep and goat,
which greatly aid separation of adult mandibles of
these two species. The present paper describes tested
diagnostic criteria for the separation of adult mandi-
bles (with deciduous premolars replaced by permanent)
of sheep and goat. These criteria partly overlap with
those of Helmer, but also cover the molars (M1–M3)
and parts of the mandible and can thus be applied to a
broader range of fragmentary material. They can also
be used to separate younger mandibles in which appli-
cation of Payne’s published criteria is precluded by loss
of dp3 and dp4 and degree of wear of M1 and M2 (in
our experience, beyond wear stage 7a of Payne, 1987:
610 Figure 1).
Materials and Methods
Diagnostic criteria were initially sought, tested, revised
and refined on adult mandibles of 43 sheep and 41
goats held in the modern reference collection of
the Department of Archaeology and Prehistory,
University of Sheffield. In addition, also from the
545
0305–4403/02/$-see front matter 2002 Elsevier Science Ltd. All rights reserved.

Sheffield collection, young mandibles (but with M1
erupting or erupted) were examined of 20 sheep and 28
goats, in order to explore further the speciation of M1
and M2 in early wear. Because the ultimate aim of this
project was the archaeozoological analysis of mortality
patterns rather than taxonomic differentiation per se,
the search for diagnostic criteria was restricted to the
mandibular cheek tooth row (P2–M3) and to the
adjacent parts of the mandible. Useful diagnostic cri-
teria were found on P3, P4, M1, M2 and M3 and on
the parts of the mandible immediately adjacent to
P2–P4 and posterior of M3.
The revised and refined criteria were then tested in
two stages, with the aim of evaluating their diagnostic
value and reliability both individually and in combina-
tion. First, the Sheffield mandibles on which the diag-
nostic criteria were originally developed, were
examined ‘‘blind’’ (mandible by mandible, rather than
tooth by tooth or criterion by criterion). Plainly, a
blind test on an independent set of mandibles would
have been preferable, but this re-examination was
conducted some months after the material was last
handled and the collection is sufficiently large to mini-
mize the risk that the analyst can ‘‘cheat’’ by recogniz-
ing the colour, size or wear state of individual
specimens. For each modern mandible, the following
information was recorded:
(1) accession number;
(2) species attribution and, if known, provenance,
age, sex, and breed;
(3) the state of eruption and wear of P3–M3;
(4) the scoring of each of 22 diagnostic criteria
as ‘‘sheep’’, ‘‘goat’’, ‘‘possible sheep’’, ‘‘possible
goat’’, ‘‘indeterminate’’ (i.e., intermediate,
ambiguous or resembling neither) or ‘‘unsuitable
for diagnosis’’ (i.e., broken, missing, insufficiently
erupted, too worn);
(5) the overall attribution of the mandible to
‘‘sheep’’, ‘‘goat’’, ‘‘possible sheep’’, ‘‘possible
goat’’ or ‘‘indeterminate’’, taking acount not only
of the scores for each criterion but also of the
reliability and asymmetry of each criterion (see
below).
Secondly, a similar examination was conducted, and
similar records were made, of 31 adult and 40 young
sheep and 107 adult and 24 young goats held by the
British Museum of Natural History in London, the
National Museum of Wales in Cardiff(Noddle collec-
tion), the Museum d’histoire naturelle in Paris, and Dr
Ingrid Mainland of Bradford University. The London,
Cardiff, Paris and Mainland collections have the
advantage of being independent of the material on
which the criteria were developed, but could not prac-
ticably be used for blind testing because the material is
stored by taxon.
The provenance information for these modern speci-
mens is of varying quality (particularly for earlier
accessions), but the material examined for both species
includes individuals drawn from wild or feral popula-
tions and several domestic breeds, with a broad geo-
graphical range (Table 1), and ranging in age from
juveniles (with deciduous premolars), through young
adults (with permanent premolars erupting or
unworn), to elderly individuals (with all premolars and
molars heavily worn or lost). Thus intra-specific
variability, though doubtless not exhausted, has been
explored fairly thoroughly.
Description of Diagnostic Criteria
Most dental criteria are described in occlusal view and
for teeth in medium wear; criteria may be unclear or
misleading in unworn/lightly worn and heavily worn
teeth. It is useful to view also the buccal or lingual face
of the tooth for criteria P3.1–2, P4.1, M1.1–3, M2.1–3
and M3.1–4, especially if the occlusal surface has not
yet been worn flat. It is essential to view the buccal face
for criteria M1.4 and M3.5. Note that the terms
‘‘buccal’’, ‘‘lingual’’, ‘‘mesial’’ and ‘‘distal’’ are used to
describe the location of diagnostic features within
individual teeth, while ‘‘anterior’’ and ‘‘posterior’’ refer
to the orientation of these features or to the location of
features within the mandible (Figure 1).
Most of the criteria described below are ‘‘symmetri-
cal’’, in the sense that alternative forms provide means
Figure 1. The terms ‘‘mesial’’, ‘‘distal’’, ‘‘buccal’’ and ‘‘lingual’’ are
used to denote different parts of the tooth; the terms ‘‘anterior’’ and
‘‘posterior’’ are used to denote the orientation of particular features
relative to the mandible as a whole. Thus, the buccal edge of the
bucco-distal cusp (shown by a thickened line) points in a posterior
direction in the tooth on the right but not in that on the left.
546 P. Halstead et al.

of identifying both sheep and goat, with more or less
even reliability. It is important to note, however, that
some criteria are ‘‘asymmetrical’’ in that, while one
form may positively identify goats, the alternative form
is compatible with both sheep and goats.
P3 (Figure 2a)
P3.1. A vertical ridge in the middle of the lingual face
is more strongly developed in sheep. As a result, the
lingual edge of the occlusal face is usually clearly
‘‘stepped’’ in sheep and usually forms a more or less
straight line, inclining buccally in a posterior–anterior
direction, in goat.
P3.2. The vertical ridge on the disto-buccal corner
tends to be more pronounced in sheep. As a result, the
distal part of the buccal edge forms a relatively distinct
and deep hollow in sheep, but a more or less shallow
and indistinct hollow in goat.
P3.3. The mesial part of the buccal face slopes inwards
(lingually) in a posterior–anterior direction more
strongly in goat than sheep; the mesial face often slopes
anteriorly in a bucco-lingual direction in goat, but is
typically perpendicular to the axis of the mandible in
sheep. As a result, the mesio-buccal quarter of the
tooth tends towards a right angle in sheep, but towards
a more open angle (in extreme cases, almost approach-
ing a straight line) in goat. N.B. the mesio-buccal
‘‘corner’’ may be rounded or angled in both species.
P3.4. Overall, the tooth tends to be longer and slen-
derer in goat, shorter and broader in sheep, but these
dimensions may be altered by tooth wear and masked
by tooth crowding.
P4 (Figure 2b)
P4.1. The mesio-lingual corner is typically marked by a
vertical rib, projecting lingually, in sheep; this feature is
typically weak or absent in goat. (There is a similar,
but less consistent, difference between sheep and goat
on the disto-lingual corner.)
P4.2. The mesio-buccal quarter of the tooth forms an
open angle (clearly more than 90 degrees) in goat, but
is closer to a right angle (sometimes rounded, some-
times with the ‘‘corner’’ emphasized by a vertical rib) in
sheep.
P4.3. Overall, the tooth tends to be longer and slen-
derer in goat, usually shorter and broader in sheep, but
these dimensions may be altered by tooth wear and
masked by tooth crowding.
M1 and M2
M1.1 and M2.1 (Figure 2c).The mesial part of the
buccal edge (ignoring the buccally projecting flange on
Table 1. Sources of modern comparative material consulted
a. Nos of individuals studied, by species and age group, in each collection
Collection
Sheep Goat
TotalYoung
1
Adult
2
Young
1
Adult
2
Sheffield 20 43 28 41 132
BMNH, London 0 2 9 40 51
NMW, Cardiff0 0 15 42 57
MNHN, Paris 0 12 0 25 37
Mainland 40 17 0 0 57
Total 60 74 52 148 334
1
With deciduous premolars in place.
2
With permanent premolars erupting/erupted.
Table b. Country of origin and species/breed/population (where known)
3
Domestic sheep Greenland (Northern Short-Tail), Iceland, Scotland (incl. Scottish Blackface), England
(incl. Herdwick, Portland, White-Faced Woodland), France, Dahomey, Greece (local and
improved breeds), Albania, Spain
Wild/feral sheep England (farmed Soay, mouflon [Ovis musimon])
Domestic goats England (incl. Bagot, Golden Guernsey), Sardinia, Greece (incl. local and improved breeds),
Spain, Egypt, Sudan (incl. Dwarf Nilotic), Nigeria, Tanzania, Comores, Madagascar,
Nepal, India, Tahiti
Wild/feral goats Scotland (Rhum feral, Arran feral), Greece (Cretan feral), Azores (?feral), Iran (C.
aegagrus), Oman (C. aegagrus)
3
Attributions to wild, feral or domestic populations, based on museum records, may be inaccurate in the case of
some early accessions.
Morphological Distinctions between Mandibles and Mandibular Teeth 547

the mesial corner) is typically convex in sheep, but
often concave in goat. Note that the mesial part of the
buccal edge may be somewhat hollow (or goat-like) in
unworn M1 and M2 and also in heavily worn M1 and
M2 (especially those beyond the ‘‘mature’’ wear stage
9A [Payne, 1987: 610 Figure 1]), even in sheep; criteria
M1.3 and M2.3 (below) may aid correct identification
in such cases.
M1.2 and M2.2 (Figure 2c).The buccal edge of the
disto-buccal cusp of M1 and M2 often points strongly
in a posterior direction in goat, while typically it is
relatively symmetrical in sheep. Thus, a strong pos-
terior orientation is indicative of goat, but a slight or
no posterior orientation, although more typical of
sheep, is compatible with either sheep or goat. Note
that the distal margin of unworn and lightly worn M1
and M2 (up to wear stages 6A/7A [Payne, 1987: 610
Figure 1]) flares in a posterior direction so that, in such
cases, the buccal edge of the disto-buccal cusp may be
symmetrical (or sheep-like) even in goats.
M1.3 and M2.3 (Figure 2c).The buccal edge of M1 and
M2 overall tends to a pointed ‘‘triangular’’ appearance
in goat and to a rounded ‘‘arcaded’’ appearance in
sheep. This criterion in part subsumes criteria M1.1–2
and M2.1–2.
M1.4 (Figure 3). A ‘‘caprine pillar’’ occurs fairly
frequently on the buccal face of goat M1 and very
rarely in sheep (Payne, 1985: 143). The presence of a
pillar is thus strongly suggestive of goat, but the
absence of a pillar has no diagnostic value. A similar
pillar was observed in M2 and M3 of goat, but so
rarely as to be of very limited routine diagnostic value.
M3 (Figure 2d)
M3.1. The mesial part of the buccal edge of the
mesio-buccal cusp of M3 (ignoring the buccally pro-
jecting flange on the mesio-buccal corner) is typically
convex in sheep, but often concave or flat in goat. Note
that the mesial part of the buccal edge may be some-
what concave (or goat-like) in heavily worn M3
(beyond wear stage 11G [Payne, 1987: 610 Figure 1]),
even in sheep; criterion M3.3 (below) may aid correct
identification in such cases.
M3.2. The buccal edge of the centro-buccal cusp of M3
often points strongly in a posterior direction in goat,
Sheep Goat
P3.1
P3.3 P3.2
P3.4
P4.3
P4.1
P4.2
P3.4
P3.2
P3.3
P3.1
P4.1
P4.2
P4.3
M1.1
/M2.1 M1.2
/M2.2
M1.3/M2.3 M1.3/M2.3
M1.1
/M2.1
M1.2
/M2.2
M3.6
M3.1 M3.2 M3.4
M3.5
M3.3
M3.3
M3.2
M3.1
M3.4
M3.5
M3.6
Figure 2. Morphological criteria for distinction between mandibular
teeth of sheep and goat, in occlusal view. Orientation: mesial(/
anterior) left, distal(/posterior) right, lingual above, buccal below.
The numbering of each criterion (P3.1, P3.2, etc.) follows the
description in the text. (a)—P3, (b)—P4, (c)—M1 and M2, (d)—M3.
Figure 3. Morphological criteria for distinction between mandibles
of sheep and goat, in lateral view. The numbering of each criterion
(MD.1, MD.2) follows the description in the text. Note that, in goat,
MD.1 may be anterior of P2 (as shown) or absent altogether. The
dental criterion M1.4 is also shown.
548 P. Halstead et al.

while it is relatively symmetrical in sheep. Thus, a
strong posterior orientation is indicative of goat, but a
slight or no posterior orientation is compatible with
sheep or goat. Note that the distal margin of the
centro-buccal cusp of unworn and lightly worn M3 (up
to wear stages 6A/7A [Payne, 1987: 610 Figure 1])
flares in a posterior direction so that, in such cases, the
buccal edge of this cusp may be symmetrical (or
sheep-like) even in goats.
M3.3. The buccal edge of the mesial and central parts
of M3 overall tends to a pointed ‘‘triangular’’ appear-
ance in goat and to a rounded ‘‘arcaded’’ appearance
in sheep. This criterion in part subsumes criteria
M3.1–2. Thus, a flat and pointed profile is indicative of
goat; conversely, a rounded profile is more typical of
sheep than goat, but is of less diagnostic value than the
corresponding criteria in M1 and M2 (i.e., M1.3 and
M2.3).
M3.4. The buccal edge of the distal cusp of M3 is often
more or less pointed in goat and typically rounded in
sheep. Thus, a pointed edge is suggestive of goat, but a
rounded edge is compatible with sheep or goat.
M3.5. The distal margin of the distal cusp of M3 often
has a buccally defined ‘‘flute’’ in sheep, rarely so in
goat. Thus, the presence of a flute is strongly suggestive
of sheep, while its absence is suggestive of goat. Note
that a flute defined only lingually is of no diagnostic
value and that a buccally defined flute may be clearer in
buccal than in occlusal view. (The diagnostic value of a
distal flute was suggested to the authors independently
by Profs H.-P. Uerpmann and A. J. Legge).
M3.6. The flange on the mesial face of M3 tends to be
broad in sheep and narrow in goat, but this feature is
heavily influenced by, and must be judged relative to,
the degree of occlusal wear. Thus, a broad flange is
suggestive of sheep in a lightly worn M3 (up to, say,
wear stage 10G/10H [Payne, 1987: 610 Figure 1]) and
less securely so in an M3 of medium wear (at the
long-lasting stage 11G), but may be compatible with
either species in a heavily worn M3 (from, say, wear
stage 12G/12H onwards). Conversely, a narrow flange
is strongly suggestive of goat in an M3 of medium or
heavy wear (from, say, stage 11G onwards) and less
securely so in a lightly worn M3.
The mandible (Figure 3)
MD.1. On the lateral face of the mandible, a foramen is
frequently found below P2–4 in sheep, but is typically
absent or, less commonly, present but anterior of P2 in
goat.
MD.2. Immediately posterior of M3, the lateral face of
the mandible usually has a more or less pronounced
hollow in goat, which is typically slight or absent in
sheep. In sheep, a single obvious sinus is common,
whereas in goat there tends to be no sinus or two or
more indistinct sinus. This criterion cannot be evalu-
ated until M3 is fully erupted and the mandible has
more or less achieved full size.
The Reliability of Individual Criteria
Taking together all the modern collections examined,
334 mandibles of known species were studied. For
individual criteria (Table 2), the number of specimens
potentially attributable to species (i.e., excluding those
‘‘unsuitable for diagnosis’’ because of breakage, loss,
insufficient eruption or excessive wear) ranged from a
minimum of 183 (55%) for M3.5 (requiring full erup-
tion of the distal cusp of M3 for reliable attribution) to
a maximum of 331 (99%) for MD.1 (usually available
for inspection even when teeth were unerupted, broken
or shed). Of these potentially attributable cases, the
number classified as ‘‘indeterminate’’ (i.e., intermedi-
ate, ambiguous or atypical) on a particular criterion
ranged between 1 and 39 (0–15%).
The accuracy of identifications to species varied
between criteria. Moreover, while some criteria (e.g.,
all those for P3 and P4) were equally able to identify
sheep and goats, other ‘‘asymmetrical’’ criteria (see
descriptions for M1.2, M2.2, M3.2, M3.3, M3.4, M3.6
and especially M1.4) could only positively identify
goats. On individual criteria, 84–100% of attributions
to goat were correct; the corresponding figures for
sheep are only 43–95%, but are much higher, at
83–95%, if the ‘‘asymmetrical’’ criteria (Table 2) are
excluded. As all the criteria used are qualitative rather
than quantitative, and as the distinction between
‘‘possible sheep’’ or ‘‘possible goat’’, on the one hand,
and ‘‘indeterminate’’ or ‘‘unsuitable for diagnosis’’, on
the other hand, is somewhat subjective, the percentage
figures for correct attribution should not be taken as
precise measures of reliability. They do, however,
suggest that, with due allowance for any ‘‘asymmetry’’,
each criterion, even in isolation, has considerable
diagnostic value.
Helmer’s study of mandibular P3 and P4 of sheep
and goats (Helmer, 2000) draws attention to criteria
essentially comparable with our P3.1, P3.3 and P4.2,
although defined in slightly different terms. He studied
40 modern mandibles not examined for the present
study and his results were only seen, in draft form,
after completion of the present study. The compatibil-
ity of the results of these two, entirely independent,
studies must be regarded as very encouraging.
The Reliability of the Criteria in Combination
When the diagnostic criteria were used in combination,
the ease of identification of modern mandibles of
known species was still variable. In some mandibles,
teeth were of intermediate appearance or different
Morphological Distinctions between Mandibles and Mandibular Teeth 549

criteria for the same tooth were contradictory. In
others, some teeth were typical of sheep and others of
goat; indeed rare examples of each tooth (P3–M3) in
mandibles of both sheep and goat closely resembled the
wrong species. In many cases, however, most or all
criteria and most or all teeth were consistently typical
of either sheep or goat (Figure 4). In general, identifi-
cation was easiest for mandibles with all teeth in
moderate wear, because the greatest number of criteria
could be assessed. It was harder to identify mandibles
with erupting or heavily worn teeth and hardest for
senile mandibles with missing teeth, because fewer
criteria could be assessed (Figure 4c,f). In identifying
each mandible, therefore, an overall judgement was
made of the strength of evidence in favour of each
species, taking account of the state of preservation
(and hence assessibility) of relevant cusps, the degree of
similarity to the ‘‘typical’’ forms, defined here, and the
‘‘asymmetry’’ of certain criteria; identification was not
based on a simple tally of the number of criteria scored
to each species and indeed, some mandibles were
(correctly, as it transpired) assigned to the species
represented by fewer criteria. The attribution of man-
dibles, using a combination of observable criteria in
this way, was remarkably successful.
When the Sheffield adult mandibles, on which the
criteria were originally developed, were re-examined
‘‘blind’’, all 84 mandibles were correctly identified to
species; young mandibles were excluded from this
exercise to avoid the risk of a successful identification
being based on Payne’s established criteria for dp3–dp4
or lightly worn M1–M2. The London, Cardiff, Paris
and Mainland collections were not examined ‘‘blind’’,
but identification based on the new diagnostic criteria
was ‘‘incorrect’’ in only three out of 138 adult mandi-
bles. Moreover, in two of these cases (one each in
London and Paris), the contentious mandible could be
securely matched with a cranium which demonstrated
clearly that our mandibular identification was correct
and the museum accession records incorrect. In the
third case (BMNH no. 3048), the accuracy of the
accession records is at least open to doubt: labelled as
a goat collected in 1962 on the Scottish island of
Rhum, this accession in fact comprises at least two
individuals, including one mandible attributed by our-
selves to goat and a pair of mandibles attributed to
sheep. In sum, the diagnostic criteria presented here
have been tested with remarkable (probably 100%)
success against a large and varied body of modern
control specimens.
While examining these modern mandibles, the op-
portunity was taken also to test the validity of Payne’s
criteria for identification of dp3–dp4 and of unworn/
lightly worn M1–M2 (Payne, 1985). Mandibles of 60
young sheep and 52 young goats were examined in
the Sheffield, London, Cardiff, Paris and Mainland
Table 2. Number and percentage of correct attributions to species for individual diagnostic criteria in modern
mandibles (all collections combined)
Attributions to sheep
1
Attributions to goat
2
Indet. Total
3
Sheep Goat
%
correct Sheep Goat
%
correct
P3.1 82 14 85 139 9 94 10 254
P3.2 40 2 95 133 8 94 11 194
P3.3 47 4 92 132 5 96 6 194
P3.4 38 8 83 112 7 94 28 193
P4.1 89 13 87 145 5 97 5 257
P4.2 58 3 95 137 3 98 5 206
P4.3 45 7 87 115 6 95 31 204
M1.1 77 6 93 133 8 92 38 262
M1.2 G 100 66 60 111 10 92 12 299
M1.3 103 15 87 152 7 96 26 303
M1.4 G 113 149 43 28 2 93 1 293
M2.1 99 8 93 145 7 95 15 274
M2.2 G 98 64 60 89 10 90 17 278
M2.3 101 9 92 120 6 95 39 275
M3.1 67 15 82 114 4 97 12 212
M3.2 G 66 31 68 89 6 94 10 202
M3.3 (G) 69 22 76 81 0 100 31 203
M3.4 G 55 52 51 67 7 91 10 191
M3.5 49 10 83 105 15 88 4 183
M3.6 G 59 33 64 90 13 87 15 210
MD.1 93 5 95 186 36 84 11 331
MD.2 67 11 86 107 0 100 12 197
1
Including specimens scored as either ‘‘sheep’’ or ‘‘possible sheep’’.
2
Including specimens scored as either ‘‘goat’’ or ‘‘possible goat’’.
3
Attributions to sheep+ attributions to goat+‘‘indeterminate’’ (i.e., excluding ‘‘unsuitable for diagnosis’’).
G ‘‘Assymetrical’’ criterion suitable for positive identification of goat only.
550 P. Halstead et al.

Figure 4. Mandibles of modern adult sheep and goats in occlusal view, illustrating more or less ‘‘typical’’ examples of various criteria.
Orientation as in Figure 1. (a) Sheep (Sheff. 0711; from Derbyshire, U.K.): all dental criteria are classified as ‘‘sheep’’, except M3.6 (‘‘possible
sheep’’) and P3.3 (‘‘indeterminate’’); both mandibular criteria (not shown) are classified as ‘‘sheep’’. (b) Sheep (Sheff. 0506; from C. Macedonia,
Greece): all dental criteria are classified as ‘‘sheep’’, except M3.4 (‘‘possible goat’’) and M1.1 (‘‘unsuitable for diagnosis’’); both mandibular
criteria (not shown) are classified as ‘‘sheep’’. (c) Female sheep (Sheff. 0482; from C. Macedonia, Greece): P3 and P4 are missing and M1 is too
heavily worn for diagnosis; dental criteria for M2 and M3 are classified as ‘‘sheep’’ (the ‘‘flute’’ of M3.5 is clearer in buccal view); both
mandibular criteria (not shown) are classified as ‘‘sheep’’. (d) Male goat (Sheff. 0787; from C. Macedonia, Greece): all dental criteria are
classified as ‘‘goat’’, except M1.2, M1.4 and M2.2 (all ‘‘sheep’’); both mandibular criteria (not shown) are classified as ‘‘goat’’. (e) Female goat
(Sheff. 0513; from C. Macedonia, Greece): all dental criteria are classified as ‘‘goat’’, except P4.2 and P4.3 (‘‘indeterminate’’), M1.1 (‘‘unsuitable
for diagnosis’’) and M3.6 (‘‘possible sheep’’); both mandibular criteria (not shown) are classified as ‘‘goat’’. (f) Female goat (Sheff. 0585f; from
C. Macedonia, Greece): M1 is too worn for diagnosis; dental criteria for P3, P4, M2 and M3 are classified as ‘‘goat’’, except P4.2 and P4.3 (both
‘‘possible goat’’) and P4.1 and M3.6 (both ‘‘unsuitable for diagnosis’’); both mandibular criteria (not shown) are classified as ‘‘goat’’.
Morphological Distinctions between Mandibles and Mandibular Teeth 551

collections. Although representing a considerably
larger and more heterogeneous control sample than
that originally used by Payne, his published diagnostic
criteria were thoroughly vindicated, with difficulties
encountered only in the case of teeth insufficiently
erupted for thorough inspection.
Application of the Criteria to Archaeological
Mandibles of Unknown Species
To date, the new criteria have been applied by the
authors to archaeological mandibles from a range of
prehistoric sites in Greece (Late Neolithic Makrigialos
and Late Bronze Age Toumba Thessalonikis in Central
Macedonia, Final Neolithic Doliana in Epirus, Late
Neolithic–Early Bronze Age cave of Zas in the Cy-
clades, Early Neolithic and Early Bronze Age Knossos
and Early Bronze Age Poros on Crete, Late Bronze
Age Pylos in Messenia), Ireland (Mediaeval Dublin)
and Scotland (Mediaeval Dun Vulan in the Outer
Hebrides). As with the modern material, the archaeo-
logical assemblages include mandibles in which all
criteria are consistently suggestive of either sheep or
goat, others in which some criteria suggest sheep and
some suggest goats, and others in which some or
all criteria are ambiguous. Identification is further
hindered by the fragmentation of the archaeological
assemblages, although identification of loose teeth on
these criteria did speed up the strewing and mending of
fragmented mandibles. These adult mandibular criteria
‘‘work’’ well in the sense that most of the material
examined could be identified with some confidence,
with the greatest difficulties being posed by incomplete
or heavily worn specimens. The proportion of indeter-
minate specimens is probably higher than that encoun-
tered by the present authors in applying Payne’s
criteria for dp3–dp4 to young mandibles, but far
lower than that normally encountered in postcranial
material (following Boessneck, Mu¨ ller & Teichert,
1964;Prummel & Frisch, 1986) because of the difficulty
of identifying immature postcranial specimens and the
vulnerability of postcranial specimens to postdeposi-
tional attrition. These archaeological applications are,
of course, strictly untested, although not untestable (cf.
Loreille et al., 1997), but it is encouraging that loose
teeth and mandible fragments, independently identified
to the same species using these criteria, were frequently
found to belong to the same mandible during mending.
Conclusions
The criteria presented here, in combination with those
published by Payne (1985), enable sheep/goat mandi-
bles of all ages to be identified to species. Speciation is
most difficult in very young mandibles, with unerupted
teeth, and in very old mandibles, with heavily worn or
shed teeth. Overall, however, teeth tend to be better
preserved than postcranial material and so these cri-
teria are widely applicable. All of the criteria exhibit
intra-specific variability, but no more so than the
widely used postcranial criteria. As with the postcra-
nial criteria, these mandibular criteria are more reliable
in suites than individually and are best interpreted with
the aid of modern reference specimens. The importance
has been stressed of distinguishing between ‘‘symmetri-
cal’’ criteria, equally able to identify both sheep and
goats, and ‘‘asymmetrical’’ criteria which only allow
positive identification of one species (in this case goat);
similar caution is needed with some of the standard
postcranial criteria for speciating sheep and goats.
Acknowledgements
The initial impetus for this study was a drawing on a
beer-mat at Troy by Hans-Peter Uerpmann of a cri-
terion (our M3.5) which was subsequently and inde-
pendently suggested also by Tony Legge. In building
up the Sheffield collection of modern mandibles, we are
particularly grateful to Francis Pryor and Maisie
Taylor, for Soay sheep and mouflon carcases from
Flag Fen Visitor Centre, in England, and to Stelios
Moukhalis of Assiros, in Greek Macedonia, for heads
of local sheep and goats; Yannis Tzelepis and Panagi-
otis Miaouras kindly provided burial space in Assiros
and Makrigialos, respectively, for our Greek speci-
mens, while Rocky Hyacinth and Glynis Jones helped
with burial and cleaning. The collection and initial
analysis of modern reference material in Greece was
undertaken in tandem with archaeozoological study of
material from Makrigialos, funded by the British
Academy and the Institute for Aegean Prehistory; our
diagnostic criteria were refined with the help of AHRB
funding for faunal analysis at Makrigialos.
For access to collections at the BMNH in London,
Natural History Museum in Cardiffand Museum
d’histoire naturelle in Paris, we are grateful to Paula
Jenkins and Richard Sabin, to Peter Howlett, and to
Jean-Denis Vigne, respectively. Louise Martin and
Ingrid Mainland have provided a variety of valuable
comments on the project and the latter also made
accessible a large assemblage of sheep mandibles from
Scotland and Greenland. We are indebted to Jenny
Marchant and Colin Merrony respectively for original
drawing and final composition of Figures 1–3,toRob
Craigie for production of Figure 4, and to Glynis
Jones, Ingrid Mainland, Louise Martin and Jean-Denis
Vigne for comments on the clarity of the text. Finally,
we thank Jean-Denis Vigne for bringing to our
attention the parallel study by Daniel Helmer.
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