The Innominate Bone Sample from Krapina

Abstract

The Croatian site of Krapina has yielded a large collection of human fossils
attributed to the archaic Neandertals. The sample includes fourteen inno-
minate bone specimens, minimum number of seven individuals (MNI=7).
Among them, it is possible to distinguish two fully adults (one female, one
male), two late adolescent or young adults (both males) and three children
of unknown sex. Metric analysis reveals the Krapina hip bones to be
characterized by relatively small vertical acetabular diameter compared to
the classic Neandertals, and a long and remarkably slender pubis relative to
living humans. Morphologically, the Krapina specimens are included with-
in the Neandertal variation, showing a narrow, rounded and/or tilted bone
surface between the coronal portion of the greater sciatic notch, and a
distinctive morphology of the superior pubic ramus, the latter agreeing with
the metric data. On the other hand, they can be distinguished from the
modern human innominate bone in aspect related with the anterior inferior
iliac spine, the topography of the posterior wall of the acetabulum, the
supraacetabular sulcus and some traits of the superior pubic ramus.

INTRODUCTION

Excavations carried out by Prof. Dragutin Gorjanovi}-Kramberger,
between 1899–1906 in the Hu{njakovo rock-shelter close to the

village of Krapina (Croatia), yielded a large collection of faunal remains,
stone tools and human fossils (1–5). The human fossil sample from
Krapina is the largest Neandertal collection from a single site, with
some 874 human remains present (6). The human fossils derive from
seven of the nine stratigraphic levels (7) identified by Gorjanovi}-
Kramberger in 1899, and are associated with a Middle Palaeolithic
stone tool technology (8, 9). The human fossil sample has been dated to
130±10 kyr (10) by combined ESR and U-series techniques on tooth
enamel from associated faunal remains, and they have been widely
considered as displaying a fully Neandertal suite of anatomical cha-
racteristics (11, 12). Importantly, the collection includes a large number
of postcranial remains (6, 13–18), including a large sample of in-
nominate bones (6, 13), one of the least represented portions of the
human skeleton in the fossil record. Therefore, the Krapina sample can
provide important insights into the evolution of the pelvis in Pleisto-
cene Homo.

Prior studies have catalogued the innominate bone collection from
Krapina (6, 13), and this sample has frequently been included as part of
the comparative sample in studies of the evolution of this region of the
body (19–22). However, to date, no detailed study has focused exclusively
on this collection. The present study provides an in-depth inventory of

BONMATÍ A.1,2

ARSUAGA J. L.1,2

1 Centro de Investigación (UCM-ISCIII) de
Evolución y Comportamiento Humanos
c/ Sinesio Delgado, 4
28029 Madrid Spain

2 Dpto. de Paleontología
Facultad de Ciencias Geológicas
Universidad Complutense de Madrid,
Ciudad Universitaria
28040 Madrid Spain

Correspondence:
Bonmatí A.
Centro de Investigación (UCM-ISCIII) de
Evolución y Comportamiento Humanos
c/ Sinesio Delgado, 4
28029 Madrid Spain
E-mail: abonmati@isciii.es

Key words: Innominate Bone, Krapina,
Neandertal, Pubis

Received June 1, 2007.

PERIODICUM BIOLOGORUM UDC 57:61
VOL. 109, No 4, 335–361, 2007 CODEN PDBIAD

ISSN 0031-5362

Original scientific paper



the hip bone collection, estimates the minimum number
of individuals represented within the sample, and at-
tempts to determine their corresponding ages at death,
sex and body mass. In addition, anthropological mea-
surements as well as the expression of morphological
features in the Krapina specimens are compared with
both Neandertals and modern humans to establish the
morphological affinities of the Krapina specimens.

MATERIAL AND METHODS

The innominate bone sample from Krapina is com-
posed of fourteen elements, ranging from almost com-
plete specimens to more fragmentary remains. The first
inventory of the Krapina remains was made at some
point after 1924 (6) by the director of the excavations,
Dragutin Gorjanovi}-Kramberger, who distinguished and
numbered six innominate bones as Coxal 1 to 6. These
labels, still visible today, were written on the bone in
black ink. Based on this first inventory, the curator and
subsequent director of the Croatian Natural History
Museum, Josip Poljak, carried out an inventory of the
human and faunal remains which included most of the
currently-known sample. Radov~i} et al. (6) have noted
that this must have happened in the early 1930s. Poljak´s
inventory was based on a fractional numbering system,
in which the most complete elements received individual
numbers whereas fragmentary remains were provided
with fractional ones, both labeled in red ink to distin-
guish them from the original black label assigned by
Gorjanovi}. Several workers subsequently reviewed parts
of the sample (13–18), and a complete catalogue was
finally published in 1988 (6), updating that of Poljak by
including the most fragmentary remains and those iden-
tified within the faunal sample that were not previously
catalogued, together with the original Gorjanovi} nota-
tion. According to Radov~i} et al. (6), the entire innomi-
nate collection was already labelled by Poljak and any

modification in the numbering system was done later.
Therefore, in the present study we have adopted the
catalogue number figured in Trinkaus (13) and Radov~i}
et al. (6), referring to both the catalogue number and the
coxal number, when appropriate.

Data for the Krapina sample was collected on the ori-
ginal fossil collection housed at the Croatian Natural
History Museum (CNHM) of Zagreb, where measure-
ments, photographs and observations were undertaken
on the 14 innominate bones. For comparative purposes, a
sample of hominin fossils has been used, relying on data
collected on original specimens and from high quality
casts of the originals or from the literature. In addition, a
large sample of recent humans has been used from the
Beira litoral region of Portugal, housed at the Instituto de
Antropologia of the Universidade de Coimbra (N=448,
209 females and 241 males) (Table 1).

To summarize and compare the fossil collection, a
comprehensive inventory has been carried out, including
a brief description of the element, which side it comes
from and the approximate age at death. The age catego-
ries used in the present study are based on the ossification
pattern of the different pelvic elements in modern hu-
mans, and include: child (no evidence of ossification of
the triradiate cartilage), juvenile (close to or complete
ossification of the triradiate cartilage, rest of epiphyses
unfused), late adolescent (complete ossification of the
triradiate cartilage and nearly complete ossification of
the rest of the epiphyses) and adult (no evidence of
unfused sutures, some degenerative processes may be
evident). Based on the recent review of the developmen-
tal pattern of the pelvis in living humans (23), we have
proposed a tentative age at death for some specimens. In
the case of adult individuals, the age at death estimations
were based on the degenerative modifications of the auri-
cular surface of the sacroiliac joint (24, 25) and on the
appearance of the articular surface of the acetabulum

336 Period biol, Vol 109, No 4, 2007.

A. Bonmatí Krapina Innominate Bones

TABLE 1

Comparative sample consulted in the present study.

Taxon Sample size Specimens (source)

Australopithecus 4 AL 288-1 (cast), Sts 14 (cast), Stw 431 (cast), MLD 25 (cast)

Paranthropus 3 SK 50 (cast), SK 3155 (cast), TM 1605 (cast)

Homo ergaster 4 KNM WT 15000 (cast), KNM ER 1808 (cast), KNM ER 3228 (cast), OH 28 (cast)

Middle Pleistocene Homo 14 Arago 44 (cast), Sima de los Huesos sample (fossil), Broken Hill E 719 (cast)

Neandertals 10 Villafamés 2 (cast, 83), Le Prince 1 (79), L’Hortus 45 (20),
La Chapelle-aux-Saints 1 (79,80), La Ferrassie 1 (79), La Ferrassie 6 (66),
La Ferrassie 8 (66), Neandertal 1 (cast), Amud 1 (fossil), Kebara 2 (fossil),

Shanidar 1 (21, 70), Shanidar 3 (70), Tabun C1 (69)

Fossil Homo sapiens 7 Qafzeh 8 (fossil), Qafzeh 9 (fossil), Qafzeh 10 (fossil), Qafzeh 13 (fossil),
Qafzeh 21 (fossil), Skhul 4 (cast), Skhul 5 (69)

Recent Humans 451 Pooled sample
(242 Males

209 Females)

Instituto de Antropologia de la Universidade de Coimbra

Numbers in parentheses indicate source of comparative data (see reference list)



(26). However, a recently published test (27) of one of the
auricular surface aging methods (25) revealed this tech-
nique to provide poor estimates of the ages at death.
Therefore, estimates based on this technique must be
considered with caution. Further, it is possible that the
postcranial skeleton in Neandertals followed a somewhat
different developmental pattern from that of modern
humans, as has been shown to characterize the dentition
(28). Finally, the minimum number of individuals (MNI)
has been estimated based on developmental criteria, sex
determination and repetition of anatomical parts.

At the same time, the hip joint is a weight-bearing
articulation, and is closely correlated with the body weight.
Several correlations (29–31) have been established be-
tween the femoral head diameter (FHD) and the body
mass (BM). Since the hip joint is a ball and socket arti-
culation, the head of the femur (the ball) and the diame-
ter of the acetabulum (the socket) are highly correlated as
shown by several studies (30, 32, 33), making it possible
to estimate the body mass from the vertical acetabular
diameter.

The hip bone is the most reliable skeletal element for
sexual determination (34–40), and the anthropological
and forensic literature since the late 19th century has
provided an extensive list of sexually dimorphic features
of the hip bone. There are two main approaches, mor-
phological and metric, in sex determination. The mor-
phological traits have been traditionally grouped in two
different complexes, the pubic (39, 41–44) and the sacroi-
liac (45–54), each of them composed of different charac-
ters considered diagnostic in assigning sex. In addition,
different techniques have tried to evaluate the sex based
on both complexes (34–36, 54–56). Further, there are
some features related to body size (i.e. robusticity and
muscles attachments) that are useful in sexual determi-
nation in extant human populations. At the same time,
the metric approach is based on quantifying both the
above-mentioned traits and the dimensions related with
body size. In extant populations, indices and discrimi-
nant functions have been developed to attempt to classify
individuals according to sex based on this metric infor-
mation (36, 37, 39, 57–65).

Period biol, Vol 109, No 4, 2007. 337

Krapina Innominate Bones A. Bonmatí

TABLE 2

Inventory of the Krapina sample.

Specimen Side Age category Age at death Sex
Attribution

Body Mass†

(1)
Body Mass†

(2)
Mean Body

Mass

Cx.1. 207* R Late
Adolescent

c. 20 yrs M 62.5 kg 67.3 kg 64.9 kg

Cx.2. 208* L Middle-Aged
Adult

30–40 yrs M 65.1 kg 69.8 kg 67.5 kg

Cx.3+Cx.6.
209+212*

L Old Adult c. 50 yrs F 63.8 kg 68.5 kg 66.2 kg

Cx.4. 210 L Adult – ? – – –

Cx.5. 211 R Late
Adolescent/
Young Adult

c. 25 yrs M? – – –

255.1 L Adult – ? – – –

255.3* L Child 6–14 yrs ? – – –

255.4* L Child 6–14 yrs ? – – –

255.5* L Child 6–14 yrs ? – – –

255.6 L Juvenile/Late
Adolescent

– ? – – –

255.7 R Adult – ? – – –

255.8* L Late
Adolescent

– M? – – –

255.9 L Adult – ? – – –

255.10 R Adult – F? – – –

* Elements representing minimum number of individuals (MNI)
† Estimate of femoral head diameter (FHD) from vertical acetabular diameter (ACET) derived by us from raw data used in
reference (33):

FHD = 0.9465ACET-5.6467; r = 0.949 (n = 143, pooled-sex)

(1) Estimated according to the regression formula; BM = 2.239FHD-39.9 r = 0.98 (pooled-sex) (29)

(2) Estimated according to the regression formula; BM = 2.268FHD-36.5 r = 0.92 (pooled-sex) (30)



Sex determination in a fossil population is compli-
cated by a number of factors. In some cases, it is not clear
that the sex-specific patterns of anatomical variation seen
in extant populations are expressed similarly in the fossil
specimens. In addition, the often fragmentary nature of
fossil specimens severely limits the amount of morpholo-
gical and metric information preserved, and particular

diagnostic regions may not be represented. This is the
case with many of the specimens from the Krapina sample.
Fortunately, some specimens do preserve diagnostic sex-
-related features. Nevertheless, these sexual determina-
tions should be considered with caution, since the pro-
bability for a correct assessment decreases when just one
shape or metric character is considered. In addition,

338 Period biol, Vol 109, No 4, 2007.

A. Bonmatí Krapina Innominate Bones

Figure 1. Krapina 207 (Cx. 1). Medial view (a) showing the distinguishable AIIS morphology in ventral (b) and medioventral (c) orientations. The
ventral margin of the sacropelvic region and the anterior margin of the sciatic notch form a single arc (1). In lateral view (d) it is possible to observe
the acetabulospinal buttress (2), the acetabulocristal buttress (3), the aperture of the greater sciatic notch (4), the interspinous notch (upper arrow)
and the supra-acetabular sulcus (lower arrow). Scale bar = 5 cm.

Figure 2. Krapina 207 (Cx. 1). Details of the ilium (a) and ischium (b), showing unfused sutures on the iliac crest (arrow, a) and ischial tuberosity
epiphysis (arrows, b). Scale bar = 2 cm.



sexual attribution becomes much more complicated in
immature individuals. Furthermore, although there is an
extensive amount of literature addressing this problem, it
is widely held that sexual dimorphism is not clearly
expressed until puberty (23). Hence, no attempt has been

made to sex the specimens in the youngest (child) age
category.

Finally, the relevant anthropological measurements
and morphological traits preserved in the adult Krapina
specimens have been described and compared with Ne-
andertals and modern humans in order to assess the
population affinities of the Krapina fossils, and therefore,
to place them in the context of the evolution of the
innominate bone within Pleistocene Homo.

INVENTORY

As discussed above, the Krapina innominate bones
were numbered from 1 to 6 by Gorjanovi}-Kramberger
in 1924. The present inventory is based on these labels,
together with the numbering system subsequently ap-
plied by Poljak. More detailed descriptions of the pre-
servation of the individual Krapina specimens can be
found elsewhere (6, 13). Table 2 summarizes the infor-
mation related to the inventory.

Krapina Cx 1. 207. (Figures 1, 2, 20). This is a right in-
nominate bone with a mostly complete ilium and ischium.
Ossification has begun in the anterior portion of the iliac
crest epiphysis, with its pelvic aspect in advance of the
gluteal aspect (Figure 2a). In addition, all borders of the
ischial tuberosity have almost completed fusion (Figure
2b). Based on modern human standards, we estimate an
age at death of around 20 years (23). Further, the billow-
ing of the auricular surface supports a late adolescent age
at death for this individual.

Krapina Cx 2. 208. (Figures 3, 15–17). This is a left
innominate bone with large portions preserved of the
pubis, the ischium and the iliac body. The complete

Period biol, Vol 109, No 4, 2007. 339

Krapina Innominate Bones A. Bonmatí

Figure 3. Krapina 208 (Cx. 2). Lateral view, showing the supra-ace-
tabular sulcus (arrow). Scale bar = 5 cm.

Figure 4. Krapina 209+ 212 (Cx. 3+Cx. 6). Lateral view (a) with details of the anterior inferior iliac spine and the morphology of the iliopsoas
sulcus (b). Scale bar = 5 cm.



fusion of all the secondary ossification centres indicates
an age at death of greater than 23 years in living humans
(23). In addition, the acetabular margin is rounded with
a smooth depression on its internal border, no osteophyte
is present on either the anterior or posterior horns of the
lunate surface, the acetabular fossa is slightly deeper than
the lunate surface and some portions of its perimeter are
transforming into trabecular bone. All these features in-
dicate that this individual was probably between 30–40
years old (26).

Krapina Cx 3+Cx 6. 209+212. (Figures 4, 5, 17, 20).
This is a left innominate bone with the complete aceta-
bulum, most of the superior pubic ramus and the com-
plete auricular surface. The secondary growth centres
have completed ossification, indicating an age older than
23 years in modern populations. The modifications of
the auricular surface yield a mean age of 51 years old,

according to the aging method of Buckberry et al. (25).
However, the loss of the billowed pattern characteristic of
immature individuals, together with some granular and
transverse striations (Figure 5b), are conditions which
are more commonly found in the fourth decade (24).
Changes in the morphology of the acetabulum (Figure
5a), including a pronounced groove on the external mar-
gin of the lunate surface, the porosity on the posterior
wall of the acetabulum and the micro- and macroporo-
sity accompanied with bony activity on more than three
quarters of the acetabular fossa, all suggest an age close to
50 years (26).

Krapina Cx 4. 210. (Figures 6, 15). This is a fragment
of the right ischial body with part of the acetabulum.
Based on the appearance of the lunate surface, this indi-
vidual is most consistent with an adult morphology.

Krapina Cx 5. 211. (Figure 7). This is a left dorsal
fragment of the ilium. The compact, very dense and
smooth sacroiliac joint, together with the presence of an
easily recognized transversal organization (billowing) in
the inferior caudal ramus of the auricular surface (Figure
7b), suggest a maximum age of around 25 years (24).

Krapina 255.1. (Figure 8). This is a fragment of the left
iliac body, with only the lateral cortical bone preserved.
The anterior inferior iliac spine (AIIS) is fused, and there
are no signs of immaturity in the articular surface of the
acetabulum, suggesting an adult age for this individual.

Krapina 255.3. (Figure 9a). This is a fairly complete
left ilium. None of the secondary epiphyseal centres
show any signs of ossification, indicating an age at death
younger than 11–14 years (23). Their values for the »ante-
rior inferior iliae spine (AIIS)-ilioauricular point dia-
meter« (Va 9 = 44.5 mm, Table 7) and the »AIIS greater
sciatic notch diameter (Va 10 = 45.3 mm, Table 7) are
higher than in La Ferrassie 8 [Va 9 = 22.1 mm; Va 10 =
21.2 mm (66)], La Ferrassie 6 [Va 9 = 28.6 mm; Va 10 =

340 Period biol, Vol 109, No 4, 2007.

A. Bonmatí Krapina Innominate Bones

Figure 5. Krapina 209+212 (Cx 3+Cx 6). Appearance of the acetabulum (a) and striations of the auricular surface (arrow, b) suggest an adult age
for this individual. Not to scale.

Figure 6. Krapina 210 (Cx. 4). Lateral view of the ischial body and
preserved portion of the acetabulum. Scale bar = 2 cm.



Period biol, Vol 109, No 4, 2007. 341

Krapina Innominate Bones A. Bonmatí

Figure 7. Krapina 211 (Cx.5). Medial view (a), showing the dense and smooth auricular surface (b) with transverse undulations pattern (billowing)
at the sciatic notch margin (arrow, b) and the detail of the anterior fossa of the postauricular sulcus at the sacropelvic region (arrow, c). Not to same
scale. Scale bar = 2 cm.

Figure 8. Krapina 255.1. Lateral view of the iliac body and lunate surface, with the path of the supra-acetabular sulcus drawn over the specimen.
Scale bar = 2 cm.

Figure 9. Krapina 255.3 (a), 255.4 (b) and 255.5 (c). Lateral views. Scale bar = 5 cm.



30.5 mm (66)], Qazfeh 21 (Va 9 = 29.3 mm; Va 10 =
30.0 mm), Qafzeh 10 (Va 9 = 35.3 mm; Va 10 = 38.9
mm), Lagar Velho I [Va 9 = 31.2 mm; Va 10 = 29.2 mm
(66)] (among whom the oldest is 6 years old) and a
modern human sample between 3–6 years of age [Va 9 =
33.1±4.0mm; Va 10 = 30.8±2.8 (67)]. Therefore, we
can estimate an age for Krapina 255.3 between 6 years
and 14 years old, based on the dimensions achieved by
this individual at the moment of death.

Krapina 255.4. (Figure 9b). This is the left iliac body
of an immature individual. The portion preserved suggests
an age at death close to the Krapina 255.3, considering its
similar epiphyseal developmental stage and dimensions
(Va 10 = 47.8 mm, Table 7). The value for the »AIIS-
greater sciatic notch diameter« is slightly higher than
Krapina 255.3 and any of the fossil and modern imma-
ture comparative specimens cited above. Hence, we have
also estimated an age between 6 and 14 years old for this
specimen, but perhaps slightly older than Krapina 255.3.

Krapina 255.5. (Figure 9c). This is a left sciatic notch
and anterior auricular surface of a child probably of
similar age as Krapina 255.4 and 255.5 (6 and 14 years
old), according to the size of the portions preserved and
the developmental evidence. However, no osteometric
standard measurement could be taken to compare with
the comparative sample.

Krapina 255.6. (Figure 10). This is the anterior su-
perior portion of the left iliac blade. Its cranial margin is
probably coincident with the iliac suture between the
crest and the blade; therefore, we suggest that the iliac
crest was either unfused or only weakly fused with the
blade, indicating a juvenile but most probably a late
adolescent age at death for this individual.

Krapina 255.7. (Figure 11). This is a fragment of a
right ischial body, with part of the lunate and the poste-
rior wall of the acetabulum. The preserved region shows
a fully adult morphology with no trace of immaturity.

Krapina 255.8. (Figure 12). This is a left ilium, with
preserved portions of the iliac body and anterior margin,
together with the cranial portion of the acetabulum. The

epiphysis for the AIIS is complete, although bone forma-
tion is still present at the articular surface of the acetabulum
(Figure 12c). This corresponds to a late adolescent in-
dividual.

Krapina 255.9. (Figures 13, 16). This is a fragment of a
fairly complete left ischium. The lunate surface and the
ischial tuberosity show a mature appearance without any
traces of periostitic processes, suggesting an adult aged
individual.

Krapina 255.10. (Figures 14, 17). This is a fragment of
a right superior pubic ramus with part of the acetabulum
attached. It does not preserve any evidence of immaturity,
and therefore, we estimate an adult age for this individual.

Minimum number of individuals

A minimum number of 7 individuals (MNI = 7) was
established for the sample based on the ilium (greater
siciatic notch) and relying on the number of repeated
elements, the developmental ages of the specimens, anti-
mere symmetry and size compatibility. These 7 individuals
are represented by: Cx.1 (207), Cx.2 (208), Cx.3+Cx.6
(209+212), 255.3, 255.4, 255.5 and 255.8 (Table 3). We
have discarded an association based on antimere sym-
metry between 255.8 (R) and Cx.2 (L), Cx.3+Cx.6 (L)
and Cx.5 (L), according to the thickness of the sciatic
notch at its deepest point [Cx.3+Cx.6 = 19.1 mm; Cx.5
= 18.6mm; 255.8 = 21.5 mm], the cross-sectional shape
of the arcuate line on both 255.8 (sharper) and Cx.3+Cx.6
(rounded) and the width of the lunate surface from the
acetabular point of Genoves (35) to the AIIS (Cx.2 =
22.9 mm; 255.8 = 31.2 mm).

Possible associations

Among the remainder of the elements it is possible to
establish some potential associations, as well as, incom-
patibilities (Table 3).

§Krapina Cx.4 is a close morphological antimere of
Cx.2 at the level of both the lunate surface and the
tuberoacetabular sulcus (Figure 15). Thus, values of the
minimum width of this sulcus (68) are virtually identi-
cal: Cx.2 = 17.1mm; Cx.4 = 17.7mm. Therefore, we agree
with Trinkaus (13) in associating these two elements.

§Although Krapina Cx.5 is incompatible with 6 of the
7 individuals used to calculated the MNI, we can not
discard association between this element and Krapina
Cx.1, based on the similarity in developmental age, mor-
phology and dimensions (Thickness of sciatic notch at its
deepest point: Cx.1 = 18.3 mm; Cx.5 = 18.6 mm. Maxi-
mum thickness of the iliac blade at the level of the
sacroiliac joint: Cx.1 = 25.7 mm; Cx.5 = 25.8 mm).
Accordingly, these two specimens could be associated to
the some individual.

§Krapina 255.1 is symmetrically incompatible with
Krapina Cx.1 and Krapina 255.8, and its age at death is
significantly older than that of Krapina 255.3, Krapina
255.4 and Krapina 255.5. In contrast, Krapina Cx.3+
Cx.6 and Krapina 255.1 show a morphological affinity in

342 Period biol, Vol 109, No 4, 2007.

A. Bonmatí Krapina Innominate Bones

Figure 10. Krapina 255.6. Lateral view, showing the region running
from the anterior superior corner of the ilium to the iliac tubercle.
Scale bar = 2 cm.



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Krapina Innominate Bones A. Bonmatí

Figure 11. Krapina 255.7. The articular surface can be seen in medial view. Scale bar = 2 cm.



those common portions preserved, as well as a similar
value of the minimum width of the ilium (35) (255.1 =
54.6 mm; Cx.3 = 55.6 mm aprox.). Therefore, we agree
with Trinkaus (13) in associating these two fragments to
the same individual.

§The area of the left ilium preserved in Krapina 255.6
makes it anatomically incompatible with Krapina Cx.1,
which also preserves this region. Although it was not
possible to directly articulate this specimen with Krapina
255.8, we cannot discard their association.

The similarity in appearance of the articular surface
of the acetabulum in both Krapina 255.7 and 255.8 makes
them ontogenetically compatible. The morphology and
developmental similarity between Krapina Cx.1 and Kra-
pina 255.7 also makes its association possible. However,
due to the fragmentary nature of Krapina 255.7, it was
not possible to obtain standard osteometric measure-
ments to support these associations. On the other hand,
this specimen is clearly incompatible based on symmetry
and developmental age with Cx.2, Cx.3+Cx.6, 255.3,
255.4, 255.5 and 255.6.

§Krapina 255.9 could be developmentally compatible
with Krapina Cx.3+Cx.6. However, it is clearly incom-
patible, due to lack of symmetry, with Krapina Cx.2
(Figure 16) and due to repetition of anatomical parts
with Krapina Cx.1 and Krapina Cx.4. At the same time,
it is developmentally incompatible with Krapina 255.3,
Krapina 255.4 and Krapina 255.5, which represent im-
mature individuals.

§Krapina 255.10 could theoretically be associated with
either Krapina Cx.2 or Krapina Cx.3+Cx.6. However,
morphologically it is clearly more similar to Krapina
Cx.3+Cx.6 (Figure 17). In addition, measurement over
the depth (69) and thickness of the superior pubic ramus
(70) agrees with this a priori visual inspection (Cranio-
Caudal Depth: Cx.2 = 8.9 mm; Cx.3+Cx6 = 8.2 mm;
255.10 = 8.2 mm. Dorso-Medial to Ventro-Lateral Width:
Cx.2 = 21.9 mm; Cx.3+Cx.6 = 17.3 mm; 255.10 = 16.9
mm). Therefore, we consider an association between
255.10 and Cx.3+Cx.6 possible (contra 13).

Body Mass

Table 2 summarize body mass estimations for Krapina
Cx.1, Krapina Cx.2 and Krapina Cx.3+Cx.6 specimens.
Regressions formulae from three different studies (29, 30,
33) have been used to calculate the body mass from the
acetabular vertical diameter (see Material and Method).
These equations are based on diverse and combined sex
modern human samples. Mean values of the two esti-
mates show a range of body mass from 64.9 kg to 67.5 kg.
Although slightly different values in body mass estimates
of the Krapina specimens were obtained by Ruff et al.(29 –
supplementary information), our results fall within 2 stan-
dards deviation from the mean body mass obtained by
these authors for early Late Pleistocene populations (67.7
± 2.4 kg) (29 – Table 1).

SEXUAL ASSESSMENT

Only the elements preserving portions that are rele-
vant for sex assessment have been considered (Cx.1, Cx. 2,

344 Period biol, Vol 109, No 4, 2007.

A. Bonmatí Krapina Innominate Bones

Figure 12. Krapina 255.8. Lateral aspect (a) with the interspinal notch (upper arrow) and the supra-acetabular sulcus above the acetabular margin
(lower arrow). The lunate surface (b) shows the margins of the ossifying sheet of bone covering the joint socket of the acetabulum (arrows). Scale bar
= 2 cm.

TABLE 3

Minimum number of individuals (MNI) and associa-

tions between elements from the Krapina site

MNI Left Side Right Side

1 Cx.5.211 Cx.1.207

2 Cx.2.208 Cx.4.210

3 Cx.3+Cx.6.
206+212

255.10

4 255.3 –

5 255.4 –

6 255.5 –

7 255.8 –



Period biol, Vol 109, No 4, 2007. 345

Krapina Innominate Bones A. Bonmatí

Figure 13. Krapina 255.9. Fragment of a nearly complete left ischium. Scale bar = 2 cm.



Cx.3+Cx.6, 255.8, 255.10 and 255.11). We have approach-
ed the sexual assessment from the morphological traits as
well as from osteometric data. A summary of the sex
attribution of the Krapina specimens is shown in Table 2.

Morphological features for sex
determination

Krapina Cx.1. 207. The morphology of the greater
sciatic notch (Figure 1d4), closely resembles closely to the
stage 5 proposed by Walker (53 – Figure 1) a category
which encompasses, 90% of males. In addition, when the
hip bone is oriented with the internal side facing the
observer, the anterior margin of the sciatic notch and the
anterior margin of the sacroiliac joint describe a single

imaginary arc [i.e. absence of the composite arc, ac-
cording to Genoves (35)], and therefore, there is no space
between these two margins. This trait is more commonly
found in males individuals (1 – Figure 1). Hence, mor-
phological data suggests a male sex for this individual (in

agreement with 13)

Krapina Cx.2. 208. Neither the sacroiliac nor the pu-
bic diagnostic regions are preserved in this individual.
Nevertheless, dimensions related with body size (aceta-
bular diameter) and robusticity (pubic width and depth,
size of the ischial tuberosity) (Table 7) are at the top of
the range of variation within the Krapina sample. Thus,
we propose a probable male sex for this individual.

346 Period biol, Vol 109, No 4, 2007.

A. Bonmatí Krapina Innominate Bones

Figure 14. Krapina 255.10. A superior pubis ramus in cranial view, showing the wide pectineal surface and the eroded but sharp pectineal crest. Scale
bar = 2 cm.

Figure 15. Krapina Cx.2 (208) and Cx.4 (210). Lateral (a) and dorsal (b) views showing morphological similarity in those regions preserved in both
specimens. Scale bar = 2 cm.



Krapina Cx.3+Cx.6. 209+212. Based on the defini-
tion proposed by Genoves (35), the composite arc is
present when the outline of the anterior margin of the

sacroiliac joint and that of the anterior margin of the
sciatic notch are not part of the same arc. This is the
condition of Cx.3+Cx.6. In modern humans, this mor-

Period biol, Vol 109, No 4, 2007. 347

Krapina Innominate Bones A. Bonmatí

Figure 16. Krapina Cx.2 (208) and 255.9. Lateral (a) and dorsal (b) views showing the asymmetrical anatomy of the ischial body in these two
specimens. Scale bar = 2 cm.

Figure 17. Krapina Cx.2 (208), Cx.3+Cx.6 (209+212) and 255.10. Lateral (a) and medial (b) views. According to the morphology and robusticity,
255.10 resembles the anatomy of Cx.3+Cx.6 more than it does Cx.2. Scale bars = 2 cm.



phology is usually seen in female individuals. In ad-
dition, the preserved portion of the pubis in C.3+Cx.6 is
very long and slender (Figure 17). These traits have been
widely noted as classic Neandertal features (19, 21, 71).
Modern humans are characterized by sturdier and short-
er pubic bones; nevertheless, there is significant sexual
dimorphism in the pubic length between extant males
and females (35, 37 – among others). Although none of
the Krapina remains preserve a complete superior pubic
ramus, Krapina Cx.2 and Krapina Cx.3+Cx.6 are pre-
served approximately to the same point along the su-
perior margin of the obturator foramen; the Cx.3+Cx.6
specimen seems to be longer than Cx.2 (Table 7). Furt-
her, the Cx.3+Cx.6 pubis is relatively (compared with
the vertical acetabular diameter) longer than Cx.2 (Table
7). Regarding the pubic thickness, all the Krapina speci-
mens have a notably flattened pubis compared with mo-
dern humans (21), although Krapina Cx.3+Cx.6 and
Krapina 255.10 exhibit higher values for the flattening
index (see above) than Cx.2. However, this index has not
been proven to be a reliable indicator of sexual dimor-
phism in modern humans (16, 37). Thus, all the evi-
dence seems to suggest female affinities for Cx.3+Cx.6
and we consider this individual to be most probably a
female (in agreement with 13).

Cx.5. 211. We consider this element to belong to a
male individual based on its anatomical similarity, and
consequently its association (see above), with Krapina
Cx.1.

255.8. The anterior margin of the auricular surface
forms a single arc with the contour of the sciatic notch
(i.e. absence of the composite arc). Further, the acute and
prominent arcuate line and the large size of the pre-
served areas are indicative of robusticity. Consequently,
we tentatively consider this individual as male.

255.10. We consider this element to belong to a female
individual based on its anatomical similarity, and conse-
quently its association (see above), with Krapina Cx.3+
Cx.6 (Figure 17).

Quantitative approach to sex
determination

As commented above, multiple methods based on
indices and discriminant analysis have emerged and have

348 Period biol, Vol 109, No 4, 2007.

A. Bonmatí Krapina Innominate Bones

TABLE 5

Discriminate analysis classification matrix in the mo-

dern human sample.

M
(p=0.54187)

F
(p=0.45813)

% correct
classification

Males 184 36 83,64

Females 36 150 80,65

Total 220 186 82,27

Rows indicate observed classifications, columns predicted
classifications

T
A

B
L
E

4

D
is

c
ri
m

in
a
n
t
a
n
a
ly

s
is

s
u
m

m
a
ry

.

W
ilk

s’
Pa

rt
ia

l

Va
ri

ab
le

s
M

al
es

Fe
m

al
es

N
L

am
bd

a
L

am
bd

a
F

–
re

m
ov

e
(1

,4
04

)
p-

le
ve

l
To

le
ra

nc
e

C
la

ss
ifi

ca
tio

n
fu

nc
tio

ns

M
ea

n
(±

s.
d.

)
(r

an
ge

)
(n

)
M

ea
n

(±
s.

d.
)

(r
an

ge
)

(n
)

M
al

es
(p

=
0.

54
18

7)
Fe

m
al

es
(p

=
0.

45
81

3)

“M
in

im
um

w
id

th
of

th
e

ili
um

”
/

“S
up

ra
ac

et
ab

ul
ar

-I
lio

au
ri

cu
la

r
di

am
et

er
”

In
de

x

89
.4

7
(±

5.
80

)
(1

04
.5

-7
0.

9)
(2

20
)

78
.8

(±
5.

94
)

(9
6.

9-
64

.8
)

(1
86

)
40

6
1,

00
00

0
0,

54
84

3
33

2,
65

45
0

0,
00

00
0

1,
00

00
0

2,
59

4
2,

28
44

C
on

st
an

t
–

–
–

–
–

–
–

–
–1

16
,6

5
–9

0,
79

24

Se
e

Ta
bl

e
7

fo
r

va
ri

ab
le

de
fin

iti
on

s.



been tested on extant humans. Although these quanti-
tative approaches could theoretically be used on fossil
populations, the state of preservation of the Krapina
sample limits the collection of standard osteological mea-
surements. Therefore, we have decided to use an index
which we believe reflects sexual dimorphism in extant
humans, and can be established in some specimens of the
Krapina sample (n=3). We have calculated this index
according with the following formula: (Minimum width
of the illium (35) / Supraacetabular – Ilioauricular chord
(35)*100) (Tables 7–9). This index is a measure of the
relative distance between the anterior margin of the sciatic
notch and the anterior margin of the sacropelvic region
and is highly correlated with the size of the posterior
space of the pelvic inlet which is significantly larger in
modern females than in males. Thus, females tend to
show lower values for this index than males (Table 4).

First, we performed a standard discriminant function
analysis to assess the accuracy of this index in sex deter-
mination in our modern human sample (Table 4). The

value of the Partial Lambda (0.548425) indicates an in-
termediate discriminatory power for this index. The
correct sex assignment for males is 83.64% (a priori clas-
sification probability (p) =0.54187), whereas the accu-
racy for females is 80.65% (p =0.45813) (Table 4). Next,
we have investigated the probability that a modern hu-
man with the corresponding index value to that of the
Krapina specimens (n=3) would be a male or female.
We have also performed this analysis on the specimens of
the fossil sample that preserve the corresponding ana-
tomical landmarks (n=19). The probabilities for each of
the specimens of the fossil sample, including Krapina,
are presented in Table 6. Regarding the Krapina sample,
two individuals (Krapina Cx.1 and Cx.3+Cx.6) show a
higher probability of being assigned to the female sex,
whereas the third one exhibits a higher probability of
being a male individual (Krapina 255.3). As determined
above, Krapina Cx.1 was morphologically attributed to a
male individual, whereas for Krapina Cx.3+Cx.6 both
visual determination and the metric data are in agree-
ment.

Period biol, Vol 109, No 4, 2007. 349

Krapina Innominate Bones A. Bonmatí

TABLE 6

Discriminant analysis posterior probabilities in the fossil human sample.

Specimen M (p=0.54187) F (p=0.45813) Most Probable Sex
Determination

AL 288-1 (“Lucy”) 0,007598 0,992402 F

Sts 14 0,029990 0,970010 F

SK 3155b 0,012667 0,987333 F

KNM ER 3228 0,499510 0,500490 ?

AR 44 0,036187 0,963813 F

OH 28 0,150541 0,849459 F

Broken Hill E 719 0,716370 0,283630 M

Krapina 207 Cx. 1 0,041441 0,958559 F

Krapina 209+212 Cx. 3/6 0,014804 0,985196 F

Krapina 255.3 0,877959 0,122041 M

Neandertal 1 0,111759 0,888241 F

Amud 1 0,060729 0,939271 F

Kebara 2 (R) 0,077733 0,922267 F

Qafzeh 9 (L) 0,974072 0,025928 M

Skhul 4 0,988360 0,011640 M

AT Pelvis 1 (L) 0,489938 0,510062 ?

AT Pelvis 1 (R) 0,531139 0,468861 ?

AT 800 0,259834 0,740166 F

AT 1004 0,028178 0,971822 F

AT 500+AT 501+AT 708 0,431229 0,568771 ?

AT 3454+AT 3819+AT 3856 0,277668 0,722332 F

AT 3809+AT 3807+AT 3808 0,301613 0,698387 F

See Table 7 for variable definitions.



350 Period biol, Vol 109, No 4, 2007.

A. Bonmatí Krapina Innominate Bones
T
A

B
L
E

7

L
in

e
a
r

m
e
a
s
u
re

m
e
n
ts

(m
m

)
o
f
th

e
K

ra
p

in
a

in
n
o
m

in
a
te

b
o
n
e
s
.

C
x.

1
C

x.
2

C
x.

3+
C

x.
6

C
x.

4
C

x.
5

25
5.

1
25

5.
3

25
5.

4
25

5.
7

25
5.

8
25

5.
9

25
5.

10

1.
M

ax
im

um
he

ig
ht

(3
5)

(1
78

.4
)

–
–

–
–

–
–

–
–

–
–

–

2.
Il

ia
c

he
ig

ht
(3

5)
(1

11
.3

)
–

–
–

–
–

–
–

–
–

–
–

3.
M

ax
im

um
ili

ac
w

id
th

(3
5)

+
12

6.
2

–
–

–
–

–
–

–
–

–
–

–

4.
Pr

oj
ec

tio
n

of
th

e
A

SI
S

(3
5)

73
.1

–
–

–
–

–
–

–
–

–
–

–

5.
Su

pr
aa

ce
ta

bu
la

r-
Il

io
au

ri
cu

la
r

di
am

et
er

(3
5)

73
.8

79
.4

–
–

–
41

.9
–

–
–

–
–

6.
M

in
im

um
w

id
th

of
th

e
ili

um
(3

5)
54

.2
54

.6
(5

5.
6)

–
–

54
.6

37
.7

(4
4.

6)
–

61
.7

–
–

7.
Su

pr
aa

ce
ta

bu
la

r
po

in
t–

P
II

S
di

am
et

er
*

–
11

4.
2

–
–

–
–

–
–

–
–

–
–

8.
Su

pr
aa

ce
ta

bu
la

r
po

in
t–

A
SI

S
di

am
et

er
(3

5)
47

.1
–

–
–

–
–

–
–

–
–

–
–

9.
A

II
S-

Il
io

au
ri

cu
la

r
po

in
td

ia
m

et
er

(3
5)

+
75

.1
–

80
.7

–
–

–
44

.5
–

–
–

–
–

10
.A

II
S-

G
re

at
er

sc
ia

tic
no

tc
h

di
am

et
er

(8
1)

+
60

.6
–

(6
3.

8)
–

–
60

.1
45

.3
47

.8
–

69
.4

–
–

11
.A

II
S

–
P

II
S

di
am

et
er

(3
5)

–
–

12
2.

6
–

–
–

–
–

–
–

–
–

12
.I

lo
au

ri
cu

la
r

po
in

t–
A

SI
S

di
am

et
er

(3
5)

99
.7

–
–

–
–

–
–

–
–

–
–

–

13
.I

lio
au

ri
cu

la
r

po
in

t–
In

te
rs

pi
na

ln
ot

ch
di

am
et

er
(3

5)
(8

0.
0)

–
–

–
–

–
(4

5.
2)

–
–

–
–

–

14
.I

lio
au

ri
cu

la
r

po
in

t–
G

re
at

er
sc

ia
tic

no
tc

h
di

am
et

er
*

28
.5

–
36

.9
–

–
–

21
.9

–
–

–
–

–

15
.H

ei
gh

to
ft

he
ac

et
ab

ul
oc

re
st

al
bu

tt
re

ss
(6

9)
90

.9
–

–
–

–
–

–
–

–
–

–
–

16
.T

hi
ck

ne
ss

of
th

e
ac

et
ab

ul
oc

re
st

al
bu

tt
re

ss
(8

2)
12

.4
–

–
–

–
–

–
–

–
–

–
–

17
.A

SI
S

–
A

ce
ta

bu
lo

cr
es

ta
lb

ut
tr

es
s

di
am

et
er

(3
5)

54
.7

–
–

–
–

–
–

–
–

–
–

–

18
.M

in
im

um
w

id
th

of
th

e
ili

ac
fo

ss
a

(3
5)

95
.3

–
–

–
–

–
–

–
–

–
–

–

19
.H

ei
gh

to
ft

he
sc

ia
tic

no
tc

h
(5

7)
29

.2
–

(3
6.

2)
–

–
–

+
17

.7
–

–
–

–
–

20
.L

en
gt

h
of

th
e

au
ri

cu
la

r
su

rf
ac

e
(3

5)
+

29
.5

–
50

.0
–

–
–

–
–

–
–

–
–

21
.H

ei
gh

to
ft

he
au

ri
cu

la
r

su
rf

ac
e

(3
7)

+
37

.1
–

25
.3

–
30

.6
–

–
–

–
–

–
–

22
.C

ot
ilo

sc
ia

tic
w

id
th

(5
7)

31
.6

+
27

.7
28

.8
–

–
–

–
–

(2
7.

9)
(3

1.
5)

–
–

23
.P

ub
oa

ce
ta

bu
la

r
w

id
th

(8
1)

(2
1.

9)
29

.3
26

.3
–

–
–

–
–

–
–

–
–

24
.W

id
th

of
th

e
tu

be
ro

ac
et

ab
ul

ar
su

lc
us

(6
8)

11
.8

17
.1

–
(1

7.
7)

–
–

–
–

–
–

14
.2

–

25
.I

sc
hi

al
le

ng
th

(3
5)

74
.6

94
.4

–
–

–
–

–
–

–
–

–
–



When the analysis is extended to include the entire
fossil sample, 14 individuals show a higher probability of
being assigned to the female sex, four to the males, and
another four show a similar probability to be male or
female individuals. Other than Krapina 255.3, the fossil
remains that have shown a higher probability of being
assigned to the male sex, correspond to archaic Homo
sapiens (Skhul and Qafzeh) and Broken Hill E 719.
Interestingly, despite the intermediate discriminatory
power for the index (see Partial Lambda value), the
majority of the fossil specimens have a high probability of
being assigned to the female sex. This could be due to a
genuine sex bias in the present fossil sample and/or to a
different pattern of dimorphism in the extinct hominids.
In the latter case, it would mean that the fossil sample
had a smaller range of variation in the position of the
anterior margin of the sacropelvic margin in relation to
the anterior margin of the greater sciatic notch. As com-
mented above, a lower value index value is correlated
with a larger posterior space of the pelvic inlet, and
therefore, the lower degree of variation in this trait in the
fossils could be indicating a difference in the delivery
process relative to modern humans.

Morphometrical analysis of the Krapina
remains

Studies of the evolution of the hip bone in fossil
humans often suffer from a lack of metric data due to the
scarcity of complete specimens which allow for the tak-
ing of standardized measurements. In the present study,
35 metric variables have been measured in the Krapina
specimens (Table 7). Nevertheless, the comparative ana-
lysis (Tables 8–9) has relied on a subsample of 20 metric
dimensions and is restricted to late adolescent and adult
individuals within the Krapina sample.

However, some portions of the innominate are poorly
characterized metrically due to a preservation bias in the
fossil record. In addition, some traits are difficult to stan-
dardize metrically. In these cases, we have alternatively
decided to describe them morphologically within the
context of the fossil record.

Metric analysis

Raw measurements for the Krapina sample (Table 7),
modern humans (Table 8) and Neandertals (Table 9)
have been standardized using Z-scores to allow a proper
comparison between selected variables. The mean value
of the variable in the pooled-sex modern human com-
parative sample was subtracted from that recorded for
each individual of the Krapina and Neandertal samples,
and the result was divided by the modern human mean
(Table 10). A value greater than or equal to ±2.0 for the
Z-score of any variable (Table 10, in bold) has been taken
to indicate that an individual fossil specimen differs sig-
nificantly from the modern human sample.

§Ilium

Regarding the Krapina sample, the specimen labelled
as Coxal 1 shows low values in the »iliac height« and »the

Period biol, Vol 109, No 4, 2007. 351

Krapina Innominate Bones A. Bonmatí
26

.N
on

-a
rt

ic
ul

ar
is

ch
ia

ll
en

gt
h

(3
7)

(3
7.

2)
54

,5
–

–
–

–
–

–
–

–
–

–

27
.P

ub
ic

le
ng

th
(3

5)
–

+
75

.8
+

86
.2

–
–

–
–

–
–

–
–

–

28
.N

on
-a

rt
ic

ul
ar

pu
bi

c
le

ng
th

(3
5)

–
+

55
.7

+
57

.8
–

–
–

–
–

–
–

–
+

42
.2

29
.D

ep
th

of
th

e
su

pe
ri

or
pu

bi
s

ra
m

us
(6

9)
–

8.
9

8.
2

–
–

–
–

–
–

–
–

8.
2

30
.W

id
th

of
th

e
su

pe
ri

or
pu

bi
s

ra
m

us
(7

0)
–

21
.9

17
.3

–
–

–
–

–
–

–
–

16
.9

31
.D

or
so

-v
en

tr
al

di
am

et
er

of
th

e
su

pe
ri

or
pu

bi
s

ra
m

us
(2

1)
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(3
5)

29
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(2
8.

8)
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–

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Va
lu

es
in

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re

nt
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ic

at
e

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tim

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ed

va
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.+

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te

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in

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um

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e.
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ur
ce

of
th

e
va

ri
ab

le
de

fin
iti

on
si

n
pa

re
nt

he
se

s,
ex

ce
pt

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r*

:“
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pr
aa

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ta

bu
la

rp
oi

nt
-P

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di
am

et
er

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is

m
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po
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(M

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&

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ei

th
,5

9)
to

th
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po
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io

r
in

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ri

or
ili

ac
sp

in
e.

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lio

au
ri

cu
la

r
po

in
t-

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re

at
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tic
no

tc
h

di
am

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is

th
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om

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(G

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th

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no
tc

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ax
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m

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cu
at

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lin

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in

th
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ac
et

ab
ul

ar
m

ar
gi

n.



352 Period biol, Vol 109, No 4, 2007.

A. Bonmatí Krapina Innominate Bones

TABLE 8

Summary statistics of the metrical variables (mm) for modern humans.

Coimbra Modern Human Sample

Mean (± s.d.)
(range) (n)

Variable Males Females Pooled Sample

1. Maximum height (35) 211.9(±9.3)
(238.0-189.0)(208)

196.3(±9.4)
(219.0-166.0)(182)

204.5(±12.3)
(239.0-166.0)(390)

2. Iliac height (35) 124.1(±6.2)
(143.6-104.0)(208)

118.7(±6.7)
(141.0-97.8)(176)

121.6(±7.0)
(143.6-97.8)(384)

3. Maximum iliac width (35) 155.6(±7.9)
(185.0-136.5)(192)

152.9(±7.9)
(174.0-130.0)(165)

154.4(±8.00)
(185.0-130.0)(357)

4. Projection of the ASIS (35) 83.9(±5.1)
(97.4-72.0)(192)

82.6(±5.9)
(100.3-64.9)(165)

83.3(±5.6)
(100.3-64.9)(357)

5. Supraacetabular-Ilioauricular diameter (35) 68.1(±5.0)
(82.9-57.4)(220)

69.7(±6.0)
(87.0-55.2)(186)

68.9(±5.5)
(87.0-55.2)(406)

6. Minimum width of the ilium (35) 60.8(±3.8)
(69.7-51.6)(221)

54.8(±3.4)
(65.0-46.7)(187)

58.0(±4.7)
(69.7-46.7)(408)

7. AIIS-Ilioauricular point diameter (35) 71.8(±5.2)
(85.0-60.2)(213)

72.3(±5.9)
(87.0-58.3)(185)

72.0(±5.5)
(87.0-58.3)(398)

8. AIIS-Greater sciatic notch diameter (81) 70.3(±4.5)
(86.8-55.0)(214)

63.8(±4.1)
(74.0-52.7)(185)

67.3(±5.4)
(86.8-52.7)(399)

9. Height of the sciatic notch (57) 37.1(±4.8)
(51.6-24.0)(212)

41.2(±5.4)
(55.9-29.0)(151)

38.8(±5.4)
(55.9-24.0)(363)

10. Length of the auricular surface (35) 51.7(±4.5)
(66.0-40.8)(211)

48.0(±4.7)
(59.5-35.6)(178)

50.0(±4.9)
(66.0-35.6)(389)

11. Cotilosciatic width (57) 36.9(±3.1)
(45.5-30.0)(218)

33.4(±2.8)
(41.0-26.9)(189)

35.3(±3.5)
(45.5-26.9)(407)

12. Puboacetabular width (81) 26.7(±2.7)
(38.3-17.7)(206)

21.3(±2.9)
(31.3-15.0)(182)

24.2(±3.8)
(38.3-15.0)(388)

13. Ischial length (35) 94.5(±4.8)
(112.0-79.6)(200)

84.9(±4.2)
(94.0-72.0)(155)

90.2(±6.7)
(112.0-72.0)(355)

14. Non-articular ischial length (37) 48.7(±3.1)
(56.1-40.0)(213)

43.1(±2.9)
(54.6-34.2)(185)

46.1(±4.1)
(56.1-34.2)(398)

15. Pubic length (35) 86.5(±5.6)
(99.6-68.1)(187)

87.9(±5.8)
(102.5-71.0)(167)

87.1(±5.7)
(102.5-68.1)(354)

16. Non-articular pubic length (35) 66.9(±4.2)
(79.0-54.6)(188)

69.5(±4.4)
(81.0-56.9)(166)

68.1(±4.5)
(81.0-54.6)(354)

17. Depth of the superior pubis ramus (69) 15.3(±2.4)
(22.9-9.3)(212)

13.8(±2.1)
(18.4-8.8)(184)

14.6(±2.4)
(22.0-8.8)(396)

18. Maximum horizontal acetabular diameter (35) 54.3(±3.0)
(64.2-46.0)(207)

49.1(±2.8)
(59.2-41.8)(183)

51.9(±3.91)
(64.2-41.8)(390)

19. Maximum vertical acetabular diameter (35) 55.2(±2.8)
(62.0-48.2)(213)

49.9(±2.7)
(60.5-41.7)(186)

52.7(±3.8)
(62.0-41.7)(399)

20. Maximum depth of the acetabulum (35) 25.1(±2.5)
(34.5-18.5)(187)

22.8(±2.2)
(29.0-16.6)(173)

24.01(±2.7)
(34.5-16.6)(360)

Source of the variable definitions in parentheses.



projection of ASIS« compared with other Neandertal
specimens (Neandertal 1, Amud 1 and Kebara 2), and in
a lesser degree, with the modern human sample. This
could be partly due to the incomplete fusion of the iliac
crest epiphysis which is important in determining the
value of these variables. When Neandertals are com-
pared with the pooled-sex modern human sample, only
one individual (Amud 1) out of four (Neandertal 1,
Kebara 2, Krapina Cx.3+Cx.6) shows a significantly
different value in the »AIIS-Ilioauricular distance«. In
addition, the Z-score for the »projection of the ASIS« in
Amud 1(2, 76) indicates this individual differs signifi-
cantly from living humans. The »supraacetabular-ilio-
auricular chord« is significantly greater than modern hu-
mans in Neandertal 1 and Amud 1, whereas it is within
the modern human range of variation in Kebara 2, Tabun
C1, Krapina Cx.1 and Krapina Cx.3+Cx.6.

§Ischium

From the Z-score analysis, Krapina Cx.2 shows a
significantly higher value for the »non-articular ischial
length« relative to modern humans and is also very long
compared to other Neandertals (La Ferrassie 1, Nean-
dertal 1 and Kebara 2). Futhermore, La Ferrassie 1 shows
values for the »non-articular ischial length« which are
more than 2 Z-scores below the modern human mean,
whereas the two other individuals (Neandertal 1 and
Kebara 2) show values that fall within the modern hu-
man variation.

The value for the »cotilosciatic width« in the Le Prin-
ce 1 specimen is 2.67 Z-scores below the modern human
mean value, while all eight individuals in both the Kra-
pina and Neandertal samples are within the modern
human range.

§Pubis

All the Neandertal specimens in which the non-arti-
cular pubic length could be measured (La Ferrassie 1,
Kebara 2, Shanidar 1, Shanidar 3, Tabun C1) show signi-
ficantly greater values than the modern human sample.
Regarding the »depth of the superior pubic ramus«, Sha-
nidar 1, Tabun C1, Krapina Cx.2 and Krapina Cx.3+Cx.6
show significantly lower values than modern humans,
whereas Amud 1 and Kebara 2 are within but close to the
lower limits of the modern range of variation.

§Acetabulum

The value of the »vertical acetabular diameter« of the
Krapina specimens (Krapina Cx.1, Cx.2 and Cx.3+Cx.6)
is within the modern human range of variation but is
small compared with some other Neandertals. Specifi-
cally, La Chapelle-aux-Saints 1, La Ferrassie 1 and Nean-
dertal 1 are significantly larger than the modern human
sample. Krapina Cx.3+Cx.6 also shows a small value in
the »maximum depth of the acetabulum« within the
Krapina and Neandertal samples but it is not significant-
ly smaller than in modern humans. On the other hand,
Neandertal 1, Kebara 2 and Krapina Cx.1 show signifi-
cantly higher values for the »depth of the acetabulum«
compared with living humans.

Morphological analysis within the fossil
record

Apart from the metric analysis, some morphological
differences, which are difficult to quantify metrically,
emerge when the Krapina specimens are compared with
both the modern and fossil human samples. These dif-
ferences are found in 6 anatomical regions: the anterior
margin of the ilium, the supraacetabular sulcus, the ilio-
sciatic buttress, the sacropelvic region, the posterior wall of
acetabulum and the lesser sciatic notch, and the superior
pubis ramus (Table 11).

§Anterior Margin of the Ilium

Since Krapina 255.6 is an isolated fragment of the
anterior iliac border, it is not possible to orient it properly
relative to any other element of the hip bone. Therefore,
Krapina Cx.1 and 255.8 are the only elements in which
the morphology and orientation of the entire ventral
margin of the ilium can be described (Figure 1). In the
genus Homo (including Krapina specimens), when the
anterior border of the ilium is oriented in medial view,
with the ventral border of the sciatic notch in vertical
position and the gluteal surface against the observer, a
pronounced dorsally concave notch (interspinal groove)
can be seen between the anterior inferior iliac spine
(AIIS) and the anterior superior iliac spine (ASIS) (Fig-
ure 1b, 18b,c). The depth and width of the interspinal
groove is a very variable trait within both the fossil speci-
mens and extant humans. In contrast, specimens as-
signed to Australopithecus and Paranthropus (AL 288-1,
Stw 431, SK 50, SK 3155, TM 1605) tend to have a wider
and more asymmetrical interspinal groove (Figure 18a).

Regarding the anterior inferior iliac spine (AIIS) mor-
phology, three specimens from Krapina (Cx.1, Cx.3+Cx.6
and 255.8) show a deeply excavated pelvic surface of the
AIIS (sulcus iliacus) for the passage of the fibres of the
iliopsoas muscle and its sinovial burse (Figure 1c). When
the spine is observed in ventral view, it seems to be
»twisted« caudally and flattened mediolaterally, showing
an apparently gracile morphology (Figure 1b). From this
same perspective, the AIIS forms an obtuse angle with
the ventral margin of the iliac blade (Figure 1c). This
morphology is commonly found in the fossils from the
Middle Pleistocene of Europe (Arago, Sima de los Hue-
sos) and in their Neandertal descendents (Neandertal 1,
Kebara 2, Amud 1, La Chapelle-aux-Saints 1). On the
other hand, some specimens of Australopithecus (AL 288-1,
Sts 14) and Paranthropus (SK 3155) show a flat pelvic
surface of the AIIS (Figure 19a), while others (SK 50,
Stw 431) show a concave morphology of this area. In
addition, the Australopithecus AIIS is straight and aligned
with the anterior margin of the iliac blade (Figure 19a),
whereas in Paranthropus the AIIS forms an obtuse angle
with the iliac blade. In Homo ergaster (KNM ER 3228,
OH 28, Figure 19b) and fossil (Qafzeh 9, Skhul 4) and
recent modern humans (Figure 19c), the pelvic surface of
the AIIS also shows a concave shape, although it is not
excavated, and therefore, the AIIS is not twisted caudally.

Period biol, Vol 109, No 4, 2007. 353

Krapina Innominate Bones A. Bonmatí



354 Period biol, Vol 109, No 4, 2007.

A. Bonmatí Krapina Innominate Bones
T
A

B
L
E

9

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th
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va
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ab
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de
fin

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on

s
in

pa
re

nt
he

se
s.



Period biol, Vol 109, No 4, 2007. 355

Krapina Innominate Bones A. Bonmatí
T
A

B
L
E

1
0

Z
-s

c
o
re

s
v
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1.
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(3
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1,
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1,
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1,
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3.
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ax
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id
th

(3
5)

–
–

–
–

–
–

–
–

–
–

–
1,

08
–

–
–

–
–1

,4
2

4.
Pr

oj
ec

tio
n

of
th

e
A

SI
S

(3
5)

–1
,8

4
–

–
–

–
–

–
–

–
–

–
1,

31
2,

76
1,

15
–

–
–

5.
Su

pr
aa

ce
ta

bu
la

r-
Il

io
au

ri
cu

la
r

di
am

et
er

(3
5)

0,
89

–
1,

90
–

–
–

–
–

–
–

–
2,

57
3,

36
1,

11
–

–
–0

,3
4

6.
M

in
im

um
w

id
th

of
th

e
ili

um
(3

5)
–0

,8
1

–0
,7

2
–0

,5
1

–0
,7

2
–

0,
78

–
–

–
–

–
1,

24
1,

56
–0

,2
8

–
–

–

7.
A

II
S-

Il
io

au
ri

cu
la

r
po

in
td

ia
m

et
er

(3
5)

–
–

1,
56

–
–

–
–

–
–

–
–

1,
74

2,
77

–0
,1

7
–

–
–

8.
A

II
S-

G
re

at
er

sc
ia

tic
no

tc
h

di
am

et
er

(7
4)

–
–

–0
,6

5
–1

,3
3

–
0,

39
–

–
0,

13
0,

51
–

–
1,

16
–0

,7
4

–
–

–

9.
H

ei
gh

to
ft

he
sc

ia
tic

no
tc

h
(5

7)
–1

,7
7

–
–0

,4
8

–
–

–
–

–
–0

,5
2

–
–

–0
,5

2
–

–1
,8

4
–

–
–

10
.L

en
gt

h
of

th
e

au
ri

cu
la

r
su

rf
ac

e
(3

5)
–

–
0,

00
–

–
–

–
–

–
–

–
–0

,5
9

–
0,

06
–

–
–

11
.C

ot
ilo

sc
ia

tic
w

id
th

(5
7)

–1
,0

6
–

–1
,8

6
–

–
–

–
–0

,3
6

–
2,

67
–

–
1,

46
0,

73
–1

,4
6

–0
,6

5
–

–1
,5

2

12
.P

ub
oa

ce
ta

bu
la

r
w

id
th

(7
4)

–
1,

34
0,

56
–

–
–

–
–

1,
00

–
–

–
0,

04
1,

78
–

–
–

13
.I

sc
hi

al
le

ng
th

(3
5)

–
0,

63
–

–
–

–
–

–
–

–
–

–0
,0

2
–

–0
,4

3
–

–
–

14
.N

on
-a

rt
ic

ul
ar

Is
ch

ia
ll

en
gt

h
(3

0)
–

2,
07

–
–

–
–

–
–

–
–

–
2,

24
–0

,1
5

–
–0

,6
9

–
–

–

15
.P

ub
ic

le
ng

th
(3

5)
–

–
–

–
–

–
–

–
–

–
–

–
–

2,
04

–
–

–

16
.N

on
-a

rt
ic

ul
ar

pu
bi

c
le

ng
th

(3
5)

–
–

–
–

–
–

–
–

–
–

7,
13

–
–

4,
29

5,
57

3,
78

2,
77

17
.D

ep
th

of
th

e
su

pe
ri

or
pu

bi
s

ra
m

us
(6

2)
–

–
2,

39
–

2,
69

–
–

–
–

2,
69

–
–

–
–

–
–1

,9
3

–1
,5

5
–

3,
19

–
–

3,
29

18
.M

ax
im

um
ho

ri
zo

nt
al

ac
et

ab
ul

ar
di

am
et

er
(3

5)
–

1,
26

–0
,3

0
–

–
–

–
–

–
–

–
–

–
–0

,1
9

–
–

–

19
.M

ax
im

um
ve

rt
ic

al
ac

et
ab

ul
ar

di
am

et
er

(3
5)

0,
41

0,
73

0,
57

–
–

–
–

–
1,

65
3,

22
2,

04
2,

41
1,

15
1,

65
–

–
–

20
.M

ax
im

um
de

pt
h

of
th

e
ac

et
ab

ul
um

(3
5)

2,
11

1,
81

–0
,4

2
–

–
–

–
–

–
–

–
3,

20
1,

58
2,

26
–

–
–

A
va

lu
e

gr
ea

te
r

th
an

or
eq

ua
lt

o
±

2
(b

ol
d)

is
ta

ke
n

to
in

di
ca

te
a

si
gn

ifi
ca

nt
di

ff
er

en
ce

fr
om

th
e

m
od

er
n

hu
m

an
m

ea
n

va
lu

e.
So

ur
ce

of
th

e
va

ri
ab

le
de

fin
iti

on
s

in
pa

re
nt

he
se

s.



However, the AIIS of H. erectus and H. sapiens forms an
obtuse angle with the iliac blade (Figure 19c,d).

The vertical iliac buttress is split in two at the level of
the AIIS in Krapina Cx.1 (Figure 1d), with one branch
running anteriorly to the ASIS (acetabulospinal but-
tress) and the other superiorly to the iliac tubercle (aceta-
bulocristal buttress), with a smooth depression between
both buttresses. Although clearly noticeable, these struc-
tures are topographically continuous with the adjacent
margins of the external surface of the iliac blade. The hip
bones attributed to Australopithecus show a single ventral
iliac buttress (acetabulospinal buttress) close to the ante-
rior margin (Figure 18). Regarding the genus Homo, two
buttresses are found in specimens attributed to Homo
ergaster (KNM ER 3228, OH 28, Figure 18b), Homo
heidelbergensis [Arago 44, Sima de los Huesos sample
(77)] and Homo neanderthalensis (Neandertal 1, Kebara
2, Amud 1, La Ferrassie 1, La Chapelle-aux-Saints 1)
resembling those observed in the Krapina specimens.
Among fossil modern humans, Qafzeh 9 is distorted but
it is possible to distinguish a ventral buttress (acetabulo-
spinal buttress) close to the anterior border, whereas
Skhul 4 shows the acetabulocristal buttress. Modern Ho-
mo sapiens commonly show a single acetabulocristal but-
tress (Figure 18c), however, some individuals do posses
an acetabulospinal buttress together with a depressed
area between them.

§Supraacetabular sulcus

In fossils and extant modern humans, the region of
the ilium just above the cranial portion of the acetabu-
lum, where the reflected head of the rectus femoris muscle
is attached, shows a diverse morphology from convex or
flat shape (i.e absent of supraacetabular sulcus) to slightly
or more rarely well-depressed areas (Figure 18c). Aus-
tralopithecus also shows a large degree of variation (AL
228-1 and Stw 431, slightly depressed; Sts 14, well de
pressed, Figure 18a), whereas Paranthropus (SK50, SK3155)

and early representatives of the genus Homo (KNM ER
3228 -Figure 18b-, KNM WT 15000) have a well-de-
pressed triangular area that runs between the acetabulo-
spinal buttress and the cranial portion of the acetabular
margin. This configuration is also developed in the later
Lower and Middle Pleistocene African specimens OH 28
and Broken Hill E719 as well as the European Ne-
andertal lineage specimens. Krapina Cx.1, Cx.2, 255.1
and 255.8 resemble other Neandertals and archaic mem-
bers of the genus Homo in showing a wide and deep
sulcus (Figures 1b, 3, 8, 12).

Interestingly, in Krapina Cx. 1 and 255.8, the superior
margin of the acetabulum protrudes greatly in lateral
direction, whereas other remains (Krapina Cx.2 and 255.1)
show however, in all the Krapina specimens, the supra-
acetabular sulcus is clearly distinguishable, suggesting
that this feature is not only the result of the protrusion of
the acetabular margin, but, rather represents a clear de-
pression on the subchondral bone of the ilium.

§Iliosciatic buttress

In Krapina Cx.1, Cx.3+Cx.6, Cx.5 and 255.8, the
surface spreading from the arcuate line to the deepest
portion of the greater sciatic notch (iliosciatic buttress)
tends to show a narrow and pillar-shaped morphology
(Figure 20a,b). Within the fossil record, the iliosciatic
morphology is greatly variable, ranging from narrow and
pillar-shaped morphologies (SK 3155, SK 50, Arago 44,
AT-Pelvis 1, AT-800) to wider and flatter morphologies
(AL 288-1), as well as intermediate shapes (Sts 14, KNMER
3228, OH 28, AT-3807+AT-3808+AT-3809+AT-3300,
AT-1004). However, when Neandertals, including the
Krapina sample are compared with fossil and modern
humans, the former tend to show a narrower and more
robust morphology than do the latter, which commonly
show a gracile, wider, flatter and in some cases slightly
depressed morphology (Figure 20c).

356 Period biol, Vol 109, No 4, 2007.

A. Bonmatí Krapina Innominate Bones

Figure 18. Lateral view of A. africanus (a, cast), H. ergaster (b, cast) and a modern human male (c, original), showing differences and similarities in
the anterior portion of the iliac blade. The rectangular shaded regions represent the acetabulospinal (anterior) and acetabulocristal (posterior)
buttresses and the triangular shaped regions, the supra-acetabular sulcus. Arrows point to the interspinous notch. Scale bar = 2 cm.



Period biol, Vol 109, No 4, 2007. 357

Krapina Innominate Bones A. Bonmatí
T
A

B
L
E

1
1

S
u
m

m
a
ry

o
f
th

e
m

o
rp

h
o
lo

g
ic

a
l
fe

a
tu

re
s

o
f
th

e
in

n
o
m

in
a
te

b
o
n
e

in
fo

s
s
il

a
n
d

liv
in

g
h
u
m

a
n
s
.

A
u

st
ra

lo
pi

th
ec

u
s

Pa
ra

n
th

ro
pu

s
H

om
o

er
ga

st
er

/
er

ec
tu

s
E

ur
op

ea
n

M
id

dl
e

Pl
ei

st
oc

en
e

H
om

o

1.
In

te
rs

pi
na

lg
ro

ov
e

W
id

er
an

d
as

ym
m

et
ri

c
gr

oo
ve

W
id

er
an

d
as

ym
m

et
ri

c
gr

oo
ve

Va
ri

ab
le

de
pt

h
an

d
te

nd
ec

y
to

sy
m

et
ry

Va
ri

ab
le

de
pt

h
an

d
te

nd
ec

y
to

sy
m

et
ry

2.
Su

lc
us

ili
ac

us
C

om
m

on
ly

fla
ts

ha
pe

Va
ri

ab
le

D
er

pr
es

se
d,

no
n

ex
ca

va
te

d
A

II
S

pe
lv

ic
su

rf
ac

e
D

ep
re

ss
ed

,e
xc

av
at

ed
A

II
S

pe
lv

ic
su

rf
ac

e

3.
O

ri
en

ta
tio

n
of

th
e

A
II

S
in

ve
nt

ra
lv

ie
w

O
ri

en
te

d
al

on
g

th
e

ve
nt

ra
l

m
ar

gi
n

O
bt

us
e

an
gl

e
w

ith
th

e
ve

nt
ra

l
m

ar
gi

n
O

bt
us

e
an

gl
e

w
ith

th
e

ve
nt

ra
l

m
ar

gi
n

O
bt

us
e

an
gl

e
w

ith
th

e
ve

nt
ra

l
m

ar
gi

n

4.
M

or
ph

ol
og

y
of

th
e

A
II

S
St

ra
ig

ht
St

ra
ig

ht
St

ra
ig

ht
Tw

is
te

d

4.
Il

ia
c

B
ut

tr
es

s
Si

ng
le

ve
nt

ra
lb

ut
tr

es
s

Si
ng

le
ve

nt
ra

lb
ut

tr
es

s
D

ou
bl

e
bu

tt
re

ss
(v

en
tr

al
an

d
la

te
ra

l)
D

ou
bl

e
bu

ttr
es

s(
ve

nt
ra

la
nd

la
te

ra
l)

5.
Su

pr
aa

ce
ta

bu
la

r
su

lc
us

Va
ri

ab
le

W
el

l-
de

pr
es

se
d

W
el

l-
de

pr
es

se
d

W
el

l-
de

pr
es

se
d

7.
Il

io
sc

ia
tic

bu
tt

re
ss

sh
ap

e
Va

ri
ab

le
N

ar
ro

w
,p

ill
ar

sh
ap

e
W

id
e,

ro
bu

st
sh

ap
e

Va
ri

ab
le

8.
A

nt
er

io
r

fo
ss

a
of

th
e

po
st

au
ri

cu
la

r
su

lc
us

Va
ri

ab
le

A
bs

en
t

Va
ri

ab
le

P
re

se
nt

ed

9.
Po

st
er

io
r

w
al

lo
ft

he
ac

et
ab

ul
um

F
la

ts
ur

fa
ce

F
la

ts
ur

fa
ce

F
la

ts
ur

fa
ce

F
la

ts
ur

fa
ce

(e
xc

ep
tA

ra
go

44
-c

on
ve

x
su

rf
ac

e-
)

10
.O

bt
ur

at
or

in
te

rn
us

su
lc

us
po

si
tio

n
C

ra
ni

al
–

Va
ri

ab
le

Va
ri

ab
le

11
.P

ec
tin

ea
ls

ur
fa

ce
m

or
ph

ol
og

y
R

ou
nd

ed
–

–
F

la
tt

o
do

rs
al

ly
de

pr
es

se
d

12
.

D
ev

el
op

m
en

to
ft

he
ob

tu
ra

to
r

ne
rv

e
su

lc
us

Po
or

ly
–

–
W

el
l-

de
ve

lo
pe

d

13
.S

ec
tio

n
at

th
e

ob
tu

ra
to

r
ne

rv
e

su
lc

us
R

ou
nd

ed
m

or
ph

ol
og

y
–

–
S-

IF
la

tte
ri

ng
,p

la
te

-l
ik

e
m

or
ph

ol
og

y

Ta
bl

e
11

(c
on

tin
ua

tio
n)

N
ea

nd
er

ta
ls

Fo
ss

il
H

om
o

sa
pi

en
s

E
xt

an
th

um
an

s

1.
In

te
rs

pi
na

lg
ro

ov
e

Va
ri

ab
le

de
pt

h
an

d
te

nd
ec

y
to

sy
m

m
et

ry
Va

ri
ab

le
de

pt
h

an
d

te
nd

ec
y

to
sy

m
m

et
ry

Va
ri

ab
le

de
pt

h
an

d
te

nd
ec

y
to

sy
m

m
et

ry

2.
Su

lc
us

ili
ac

us
D

ep
re

ss
ed

,e
xc

av
at

ed
A

II
S

pe
lv

ic
su

rf
ac

e
D

ep
re

ss
ed

,n
on

ex
ca

va
te

d
A

II
S

pe
lv

ic
su

rf
ac

e
Va

ri
ab

le

3.
O

ri
en

ta
tio

n
of

th
e

A
II

S
in

ve
nt

ra
lv

ie
w

O
bt

us
e

an
gl

e
w

ith
th

e
ve

nt
ra

lm
ar

gi
n

O
bt

us
e

an
gl

e
w

ith
th

e
ve

nt
ra

lm
ar

gi
n

O
bt

us
e

an
gl

e
w

ith
th

e
ve

nt
ra

lm
ar

gi
n

4.
M

or
ph

ol
og

y
of

th
e

A
II

S
Tw

is
te

d
St

ra
ig

ht
St

ra
ig

ht

4.
Il

ia
c

B
ut

tr
es

s
D

ou
bl

e
bu

tt
re

ss
(v

en
tr

al
an

d
la

te
ra

l)
Va

ri
ab

le
Va

ri
ab

le

5.
Su

pr
aa

ce
ta

bu
la

r
su

lc
us

W
el

l-
de

pr
es

se
d

Va
ri

ab
le

C
om

m
on

ly
fla

to
r

sl
ig

ht
ly

de
pr

es
se

d

7.
Il

io
sc

ia
tic

bu
tt

re
ss

sh
ap

e
N

ar
ro

w
,p

ill
ar

sh
ap

e
W

id
e,

F
la

ts
ha

pe
Va

ri
ab

le
,t

en
de

cy
to

w
id

er
an

d
fla

tt
er

sh
ap

e

8.
A

nt
er

io
r

fo
ss

a
of

th
e

po
st

au
ri

cu
la

r
su

lc
us

A
bs

en
to

r
sl

ig
ht

de
pr

es
si

on
(K

eb
ar

a
2)

A
bs

en
t

9.
Po

st
er

io
r

w
al

lo
ft

he
ac

et
ab

ul
um

F
la

ts
ur

fa
ce

C
on

ca
ve

-c
on

ve
x

m
ar

gi
ns

,c
on

ve
x

su
rf

ac
e

C
on

ve
x

su
rf

ac
e

10
.O

bt
ur

at
or

in
te

rn
us

su
lc

us
po

si
tio

n
Va

ri
ab

le
C

au
da

l
Va

ri
ab

le

11
.P

ec
tin

ea
ls

ur
fa

ce
m

or
ph

ol
og

y
F

la
tt

o
do

rs
al

ly
de

pr
es

se
d

Ir
re

gu
la

r
to

ve
nt

ra
lly

cu
rv

ed
Ir

re
gu

la
r

to
ve

nt
ra

lly
cu

rv
ed

12
.

D
ev

el
op

m
en

to
ft

he
ob

tu
ra

to
r

ne
rv

e
su

lc
us

W
el

l-
de

ve
lo

pe
d

W
el

l-
de

ve
lo

pe
d

W
el

l-
de

ve
lo

pe
d

13
.S

ec
tio

n
at

th
e

ob
tu

ra
to

r
ne

rv
e

su
lc

us
E

xt
re

m
e

S-
I

F
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§Sacropelvic region

The sacropelvic region of the innominate bone shows
three easily distinguishable areas, from ventral to dorsal
direction, the auricular surface that directly articulates
the ilium with the sacrum, the postauricular sulcus that
runs parallel to the posterior margin of the auricular
surface, and the iliac tuberosity, the very rough and pro-
minent area for the attachment of the sacroiliac liga-
ments. In the Krapina sample, the sacropelvic region is
only partially preserved in Cx.3+Cx.6 and Cx.5. Both
specimens preserve the auricular surface and the postau-
ricular sulcus. The latter shows a smooth and polished
appearance of the subchondral bone, a regular width and
well defined margins all along its extension. However,
none of the remains in the Krapina collection preserve
the iliac tuberosity.

The size, shape and appearance of each of the three
sacropelvic areas in both fossil and extant humans are
highly variable. However, Lower and Middle Pleistocene
members of the genus Homo which preserve the iliac
tuberosity (KNM ER 3228, OH 28, Arago 44, AT-Pelvis
1, AT-800, AT-1004) show a very protruding and wide
morphology of this area, remarkably elevated from the
rest of the sacropelvic region. In addition, the outline of
this tuberosity resembles a rhomboid shape in medial
view. These features are not seen in the rest of the fossil
sample studied and are rarely found in the same degree
in Homo sapiens.

Interestingly, Krapina Cx.3+Cx.6 and Cx.5 show a
remarkable groove (anterior fossa of the postauricular
sulcus) in the superior ventral portion of the sulcus, just
at the border with the iliac fossa (Figure 7c). This region
is only preserved in a few fossil specimens. The inno-
minate bones from Sterkfontein (Sts 14), and particular-
ly, from Olduvai (OH 28), Sima de los Huesos (AT-1004,
AT-800 and AT-3807+AT-3808+AT-3809+AT-3300),
and Arago show this fossa well developed. Relying on
associations between hip bones and sacra in the Sima de
los Huesos collection, this fossa articulates with the la-

tero-dorsal portion of the sacrum alae. In contrast, the
remaining fossil specimens show no development of this
fossa and the living human sample posseses it in very low
frequencies.

§Posterior wall of the acetabulum and
lesser sciatic notch

The posterior wall of the acetabulum is preserved in
four specimens of the Krapina sample; Cx.1, Cx.2, Cx.3+
Cx.6 and 255.7. In lateral view, perpendicular to the
acetabular plane, the surface remains flat and is not
adapted to the convex posterior acetabular margin. This
morphology appears to be the normal condition in both
Australopithecus and archaic members of the genus Ho-
mo, including Neandertals (Kebara 2), and is also seen in
Skhul 4. However, Arago 44, and especially modern
humans, show a slightly bulky morphology, adapted to
the convex posterior acetabular margin.

All of the innominates bones that preserve relevant
portions of the ischial tuberosity and the adjacent lesser
sciatic notch for the passage of the obturator internus
muscle (Cx.1, Cx.2 and 255.9) show a continuous margin
of the ischial tuberosity. Therefore, none exhibit a con-
cave dorsal margin of the tuberosity, as other Neandertals
do (Kebara 2 and Neandertal 1). To analyze the frequen-
cy and taxonomic utility of this trait, Trinkaus (78) pro-
posed classifying its expression into three categories ac-
cording to the position of the obturator internus sulcus
relative to ischial tuberosity. Based on these categories, he
reported an »intermediate« configuration for Krapina
Cx.1, and a »cranial« configuration for Cx.2 and 255.9.
However, in our opinion, all three specimens fall within
the intermediate configuration of his classification. Within
the fossil record, Sts 14, AL 288-1, OH 28 and AT-1004
show a cranial configuration, AT-Pelvis 1, AT-800, AT-
3807+AT-3808+AT-3809+AT-3300, Broken Hill E719
and Neandertal 1 exhibit an intermediate one, whereas
KNM ER 3228, Arago 44, Kebara 2 (right ischium),
Skhul 4 (left ischium) and Skhul 5 show a caudal

358 Period biol, Vol 109, No 4, 2007.

A. Bonmatí Krapina Innominate Bones

Figure 19. Ventral and medial views of the AIIS in three different specimens corresponding to A. africanus (a), H. ergaster (b) and modern humans
(c). Shaded regions show distinctive morphological traits (see text). Not pictured to the same scale.



position of the obturator internus sulcus relative to the
ischial tuberosity. The large degree of variation in the
expression of this trait does not appear to closely follow
taxonomic categories (78), making the significance of an
intermediate configuration in the Krapina specimens
difficult to interpret.

§The superior pubic ramus

As mentioned previously, the length and cranio-cau-
dal thickness of the pubic bone is one of the most cha-
racteristic and well-known features of the Neandertal
pelvis, including those of the Krapina sample, and it
clearly separates them from Homo sapiens. In addition to
the metric differences on the variables (Table 10), the
pubic bone in Neandertals, including Krapina, also shows
a distinctive morphology in the proximal half of the
superior ramus. This area includes the pectineal surface
cranially and the obturator nerve sulcus, limited by the
obturator crests, caudally. Antero-posteriorly, it is limited
by a rounded ventral border and a usually flatter dorsal
border with the pectineal crest along its cranial margin of
the latter.

In the great apes, the pubic ramus is mostly flattened
in the A-P direction. Accordingly, the proximal half of its
superior pubis of shows cranially a thin and acute pecti-
neal crest. The obturator nerve sulcus is formed by ven-
tral and dorsal crests derived from the obturator crest that
are poorly developed and run very close to each other. As
a result, they exhibit a short and narrow obturator sulcus.

The innominate bones attributed to the genus Aus-
tralopithecus (AL 288-1, Sts 14, Sts 65, Stw 431) show an
intermediate morphology between apes and the genus
Homo. Cranially, the proximal half of the superior pubis
ramus is more developed in the A-P direction than apes,
and therefore, it exhibits a rounded bar-like morphology,
although no specimen shows a well-developed pectineal
crest. However, the obturator nerve sulcus is similar in
size and shape to that of the great apes. No superior pubis
ramus is sufficiently preserved to observe this trait in

Homo ergaster / erectus. Several Middle Pleistocene Homo
specimens from the European site of the Sima de los
Huesos (AT-1006, AT-3497+AT-3813+AT-3814, AT-
1693+AT-1709, AT-2502+AT-2508, among others) ex-
hibit a marked S-I flattening of the superior ramus,
showing a wide and mostly flat pectineal surface which is
depressed parallel to a well-defined pectineal crest (77).

Neandertal specimens (Kebara 2, Tabun C1, Shani-
dar 1, Shanidar 3, La Ferrassie 1, Amud 1), including
those from Krapina (Cx.2, Cx.3+Cx.6 and 255.10) have
exaggerated this morphology, and the proximal half of
the superior pubis ramus takes on an extreme plate-like
morphology. In contrast, modern Homo sapiens show a
rounded bar-like shape of the superior ramus, with a
curved pectineal surface. In addition, the pectineal crest
shows a pitted and rugose appearance and is rarely pro-
jected as a bony sheet. However, both Neandertal lineage
specimens and modern Homo sapiens show a long, deep
and wide obturator sulcus, probably because of a larger
space between the ventral and dorsal portions of the
obturator crest.

CONCLUSIONS

The innominate bones from the Krapina site are one
of the most important collections to study the evolution
of the hip bone. This sample is composed of 14 elements
representing a minimum of seven individuals: two adults
(Krapina Cx. 2 and Cx.3+ Cx.6), two late adolescents
(Krapina Cx. 1 and 255.8), and three children (Krapina
255.3, 255.4, 255.5). Among the adult individuals, one is
probably a male (Krapina Cx.2), whereas the other is
most probably a female (Krapina Cx.3+Cx.6). The two
sub-adult individuals most likely represent males, whe-
reas the younger immature individuals are of unknown
sex. The body mass calculated for three individuals (Kra-
pina Cx.1, Cx.2 and Cx.3+Cx.6) ranges from 64.9-66.2 kg.

The Krapina specimens show some metric differences
compared with other Neandertals, particularly in their

Period biol, Vol 109, No 4, 2007. 359

Krapina Innominate Bones A. Bonmatí

Figure 20. The iliosciatic buttress (shaded regions) in two specimens from Krapina (a, b) and a modern human male (c), showing morphological
differences in extension and robusticity. Not to scale.



small values for the »maximum vertical acetabular dia-
meter«, and one specimen (Cx. 2) shows a remarkably
high value for the »non-articular ischial length«. Inter-
estingly, studies of the dentition (18) and the temporal
bone (84) have also identified a few metric aspects which
appear to be particular to the Krapina sample. Despite
these slight differences, when the entire Neandertal sample
(including Krapina) is considered, the »pubic length«,
and probably also the »depth of the superior pubic ra-
mus«, are significantly different from modern humans,
and these two traits could be considered unique Nean-
dertal characters.

Regarding the morphological features, none of the
Krapina specimens show any derived trait that distin-
guishes them from the rest of the Neandertal sample.
However, Neandertals can be distinguished from all ot-
her hominid taxa (Table 11), by the pillar-shaped and
narrow space between the cranial border of the greater
sciatic notch and the arcuate line and the extreme S-I
flattening and plate-like morphology of the superior pu-
bic ramus. This distinctive pelvic morphology can be
seen in a less-developed state in their European Middle
Pleistocene precursors.

Acknowledgements: We thank J. Radov~i} and the staff
of the Croatian Museum of Natural History for kindly
permitting us to study the Krapina Hominid Collection. To
D. Frayer, A. Mann, J. Monge for inviting us to contribute to
this monograph on Krapina material. Also, to M. de Morais
and the direction of the instituto de Antropología of the
Universidade de Coimbr, for granting us access to the mo-
dern human collection. Thanks to Y. Rak for access to the
fossils of Kebara, Amud, Qafzeh and Skhul, as well as
several high quality cast, housed at the Institute of Anatomy
of the Sackler Medical School of the University of Tel-Aviv.
We thank the Anthropological Institute and Museum of
Zurich for the access to its cast collection. To T. Holliday for
kindly providing us with his modern human dataset. We
appreciate the constructive comments and help with the
preparation of the English manuscript provided by R. Quam
and F. Gracia, also to L. Dalen and A. Gómez for their
assistance with the statistic analysis and J. M. Carretero, A.
Esquivel, N. García, A. Gracia, C. Lorenzo and I. Martínez,
for their help and work in the field. A. Bonmatí has a grant
from the Cátedra Fundación Duques de Soria/Fundación
Atapuerca. This work is supported by the Ministerio de
Educación y Ciencia of the Government of Spain, Project
No. BOS2003-08938-C03-01.

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