1. A. Ascenzi*, A calvarium of late Homo erectus from
I. Biddittu, Ceprano, Italy
P. F. Cassoli†,
A. G. Segre & On 13 March 1994, a fragmented, incomplete and highly fossilized, human
calvarium was discovered in situ by one of the authors (I.B.) during excavations
E. Segre-Naldini for the construction of a highway near Ceprano, a town in southern Latium,
Italian Institute of Human situated about 55 miles S.E. from Rome. The remains come from a clay lying
Palaeontology, *Section of Morbid below sandy volcaniclastic gravels whose age is estimated by K–Ar to be
Anatomy, ‘‘La Sapienza’’ University & 700 ka. The shape and capacity (ca. 1185 ml) of the calvarium show that
†Soprintendenza Museo the hominid from Ceprano possesses several, but not all, of the features of
Preistorico-Etnografico ‘‘Pigorini’’, Homo erectus. 1996 Academic Press Limited
Rome, Italy
Received 1 November 1995
Revision received 22 February
1996 and accepted 24 April 1996
Keywords: calvarium, Homo erectus,
Lower Paleolithic, Italy, Ceprano,
earlier Pleistocene. Journal of Human Evolution (1996) 31, 409–423
Introduction
Ceprano is a town in central Italy, about 55 miles south of Rome. It is situated in the Ceprano
basin, one of the suite of the seven pleistocene basins that form the middle Sacco–Liri river
valley between Anagni and Cassino (Figure 1).
Over the past 140 years the Sacco–Liri river valley has been the object of geological
(Ponzi, 1857–1858; Branco, 1877; Viola, 1902; Devoto, 1965; Bergomi Nappi, 1973;
Angelucci et al., 1974; Civetta et al., 1981) and prehistoric exploration (Nicolucci, 1868,
1873; De Lorenzo d’Erasmo, 1932; Blanc, 1956). In addition, at regular intervals over
the last 35 years the Italian Institute of Human Palaeontology has carried out excavations at
Ranuccio–Anagni site (Biddittu et al., 1979; Segre Ascenzi, 1984), at Pofi (Blanc
Taschini, 1958–1961; Fedele, 1962; Biddittu Segre, 1978), at Arce (Biddittu, 1972), in the
Liri–Sora basin (Segre et al., 1984), in the Pontecorvo basin (Biddittu Cassoli, 1969) and
other sites situated further south (Biddittu Cassoli, 1969; Segre Biddittu, 1981). The
material excavated and collected comprised stone artefacts from the lower Paleolithic
(Biddittu, 1972, 1974b; Biddittu Segre, 1978, 1982a), Acheulean tools (Biddittu Cassoli,
1969; Biddittu, 1974a; Biddittu et al., 1979; Biddittu Segre, 1984), bone artefacts (Biddittu
et al., 1979; Biddittu Segre, 1982b; Bruni, 1987), vertebrate fossils (Biddittu et al., 1979;
Segre Ascenzi, 1984; Cassoli Segre-Naldini, 1984, 1995) and fresh water mollusca
(Settepassi Verdel, 1985).
On 13 March 1994, during digging for the construction of a highway through the Campo
Grande area near the town of Ceprano, fragments of a human calvarium damaged by a
bulldozer were found in situ by one of the authors (I.B.) in a clay layer. A methodical
exploration of the site lasting several weeks enabled all the remaining bone fragments to be
recovered. These were used to reconstitute an incomplete calvaria.
The present paper (1) reports a geological survey of the discovery site whose aim has been
to ascertain its geochronological age; (2) provides an initial presentation of the calvarium which
attempts to indicate its hominid type.
0047–2484/96/110409+15 $25.00/0 1996 Academic Press Limited

2. 410
0 5
km
A
M
er
in
C
og
R.
P
CP
Li
r
92
iR
.
.  ET AL.
Sacco R.
Rome
791
CV
0 50 Naples
km 1 2 3 4 5 6 7 8 9
Figure 1. Ceprano basin location map showing general geomorphologic features. 1, Oligo-Miocene sandstone and Cretaceous reef carbonatic facies; 2,
Villafranchian and lower Pleistocene fan-shaped terraced gravel and sands; 3, Ceccano’s older volcanic group; 4, later volcanics; 5, hilly morphology:
middle Pleistocene gravels, volcaniclastic sands and clays, all covered by later quaternary colluvial; 6, position of human calvarium at ‘‘Campo Grande’’;
7, fossil vertebrates sites; 8, lower Paleolithic artefacts ‘‘chopper facies’’; 9, K–Ar dated volcanics. Towns: C, Ceccano; P, Pofi; CV, Castro dei Volsci;
CP, Ceprano; A, Arce.

3.     HOMO ERECTUS  ,  411
Figure 2. Composite section through the Ceprano basin. N, Neotectonic raised countryside area. L, Low
basin area. 1, Oligocene. 2, Upper Pleistocene, Mousterian (Figure 3: 1 and M). 3, Middle Pleistocene,
Acheulean (Figure 3: 2–8 and A2, A1). Lower middle Pleistocene: 4, colluvial-fluvial (Figure 3: 9–13 and H
calvaria position, C chopper artefacts). 5, silt-clay limnic series; 6, basal coarse gravels. 7, Villafranchian(?):
residues of highly fragmented fan cone pebble remnants. 8, Miocene: Messinian, Tortonian shallow marine
sandstone; 9, Mesozoic: Cretaceous carbonatic rock series. F, fault. The same section is reported below on
scale d=1; h= 2.
Geology and evolution of the Ceprano Basin
Before attributing a geochronological age to the calvarium, some connected geological
problems and features must be considered.
The prequaternary Sacco–Liri Rivers valley substratum displays a succession of buried
paleomorphological basins of previllafranchian tectonic origin caused, and further depressed,
by fault systems. Tectonic movements kept place between Messinian and middle Pleistocene.
The last tectonic phases were accompanied by volcanic activity between 700 and 100 ka. The
valley is delimited at S.W. by the overthrusted Lepini–Ausoni mesozoic limestone mountain
system (Figure 2).
After the last Miocene Messinian regressive marine facies, with the microfauna of very
shallow water in stratified marls, the oldest Quaternary sediments are rare Villafranchian red
clay and breccia remains containing Stephanorhinus etruscus, preserved in deep paleokarst
morphological cavities within the lower Miocene and Cenomanian limestone near the town of
Ceccano, in the northern part of the basin.
The lower part of a fan-shaped series of consolidated boulders and sandy gravels, forming
a band along the foothills on the right side of the basin, is the next younger formation (Figure
1). The pebbles of these coarse-grained deposits originate from the disintegration of late
Miocene sandstone and hardened conglomerate issuing from a phase of tectogenic activity.
Superimposed sand and gravels of the formation can be ascribed to a final Villafranchian age.
In the later Pleistocene, the fan-shaped formations were terraced, dissected and then deeply
eroded. The area covered by these deposits once extended far into the basin, and the oldest
lithic industry of chopper facies mostly on quartzite and quartzsandstone is found in many of
the remains of the formations (Biddittu, 1974a,b), in the Castro dei Volsci countryside.
Chopper facies of the same kind are also found at the Campo Grande site (Figure 3, no. 13-C).
No Pliocene deposits have been found in the basin.

4. 412 .  ET AL.
Late neotectonic faults altered the Pleistocene hydrography, and fragmented the oldest fan
cones along the basin south side. Two sectors may be distinguished between Castro dei Volsci
and Falvaterra towns where these faults are more or less recognizable.
Two neotectonic faulted and folded substratum ridges rise from the bottom of the basin.
This, as well as their shaped and covered paleomorphology, account for the differences within
the series of Pleistocene sedimentary lithofacies corresponding to the depression deep below
the surface.
Some of the Ernican little fissure volcanoes that lie along the faults in the longitudinal
neotectonic system are included in the Ceprano basin. Eleven K–Ar dated sites using
leucit-tefrit lava (Viola, 1902; Civetta et al., 1981) established that Ceccano is the oldest
volcanic unit ranging between 700 20 and 680 20 ka (Basilone Civetta, 1975; Fornaseri,
1985), other volcanic units are included between 370 40 and 120 60 ka (Figure 1). In the
area surveyed at Ceprano itself, augite and leucite fluvial sands are found throughout the
higher layers of the middle Pleistocene series, ending above the unconformity situated over the
layer containing the hominid calvarium. All the seven available K–Ar data come from lava
samples. Neotectonics, paleomorphology, and this later volcanic activity repeatedly modified
the Middle Pleistocene paleohydrography of the basin. As a result, the thickness of the
sedimentary quaternary series varies from about 15 m on the S and S.W. sides of the basin to
over 50 m at some points of the central area. In the general stratigraphic column, six
depositional ensembles are recognizable. They are distinguished by continuous, and in some
places marked unconformities (Figure 3).
At the top of the series, a variable ‘‘terra rossa’’ layer with artefacts of archaic-Mousterian
facies (Figure 3, layer 1) it is underlined by generalized gravel, convoluted silt and augite sand
layers (2) and fluvial current and cross-bedded facies (3). Upper Acheulean artefacts and fauna
(A2) (Biddittu Cassoli, 1969; Biddittu, 1974a) are preceded by an irregular buried
morphology, the paleovalleys are filled by leucite–tephra and tufitic boulders (4), that are
fragments of tuffaceous strata interbedded in the alluvio–colluvial deposits.
The third ensemble (Figure 3; 5–7) is of fluvial facies, yellow silt and clay layer. Its lower
part, which at some sites in the basin (e.g. the Meringo river, Figure 1), has a much greater
thickness than in others, is of fluvial gravels and augite cross-bedded sands containing
scattered Unio (Auricolaria) sinuata freshwater shells (7) of very large size; this formation
contains the older Acheulean artefact facies (A1), with the presence of bone tools and fauna
comprising Elephas Palaeloxodon antiquus, Stephanorhinus hemitoechus, Hippopotamus sp., Megaloceros
verticornis, Dama dama clactoniana, Castor sp. and Emys orbicularis. This level may be correlated
with the Anagni basin Ranuccio facies (Biddittu et al., 1979; Segre Ascenzi, 1984; Biddittu
Segre, 1982b, 1984) dated K–Ar 458 6 ka, located about 22 miles from the Ceprano
site.
The next layers (8) include a ferruginous crust, silt with oxidated marks of swamp flora,
analcimized leucite sand, limestone freshwater concretions and distinctive, cylindrically shaped
travertine rods of Carex sp. and, below these, a ferromanganiferous crust denoting an evident
unconformity.
The fourth ensemble (Figure 3; 9–11) constitutes marshy and paleocolluvial facies (9): it is
the outcrop site of the human calvaria (H). This locality is part of a hard clay layer about 3 m
thick (9), exhibiting concoidal cracks and transverse fissures and including small scattered limy
nodules. In this barren layer, which is macro- and even microfossil sterile, pollen analysis was
totally negative. All this reported evidence indicates that the site was a colluvial–alluvial
paleosoil originating from a slope that ended in a low marshy pool. In this basin paleosols in

5.     HOMO ERECTUS  ,  413
m110
m0 H a b c U
1
M
U
14
2
A2
3
15
4
U 30
16
5
6
10
7 17
A1
8
U
H 9 18
40
10
19
11
U
12
C
13 20
S
U
Figure 3. Generalized composite stratigraphic column for the central Ceprano basin. Mark signs: H, human
calvarium; a, artefacts; b, fossil vertebrates; c, freshwater shells; U, unconformities. Explanations of numbers
and letters are given in the main text.
colluvial deposits are developed on mesozoic limestone slopes and on tectonized oligomiocene
clays and shales.
This bed is underlying by travertine (10) and by yellow sands and concretions rising on an
unconformity surface containing clay pebbles (11) left by the erosion of still lower layers (14),
so at least two phases of erosion and colluviation, (4) and (11), are identified in connection with
the latest tectonic.

6. 414 .  ET AL.
The fifth ensemble (Figure 3; 12–13) has a variable thickness, up to maximum of 4–5 m. It
comprises above cross-bedded fluvial sands containing only Unio sp. shells that are mostly
fragmented (12). The underlying group of layers (13) begins with a 20 cm layer containing
pebbles and quartzite chopper artefact facies, probably derived from the extreme outward
limit of fan-shaped formations (no. 2 in Figure 1) on the side of Castro dei Volsci. The series
of grey and red sands forming a differentiated, unsorted suite (13) is interrupted by a thin hard
oxidized layer consisting of lime and ferromanganese compounds and by a freshwater silt layer
with Valvata sp. and Pisidium amnicum mollusc fauna. All of this sand group contains a
fragmentary fauna with Elephas throgontherii that is ascribable to the earlier part (i.e. Galerian)
of the Cromer complex. It is still under study. These lowest sand layers (Figure 3, no. 13)
contain only worn augite crystals that come from much more distant volcanics or from an
older, buried volcanic phase that does not outcrop locally; leucite crystals (useful for K–Ar
dating) have not been found.
Below this there is an unconformity that marks the limit between the whole of the upper
fluvial–continental series (Figure 3; 1–13) and continuous limnic facies (14–19) of the lower
Pleistocene. This lower part of the succession has become known only through an industrial
drilling operation (Figure 3, right side). It comprises: clay with freshwater shells, mostly
showing a Valvata–Pisidium facies (14); a peaty silt with lignite (15); lime and sandstone
concretions (16); clay and silt with scattered concretions (17); a peaty clay and lignite layer (18);
freshwater shelly clay (19); coarse basal gravel with pebbles, which is the oldest basal filling of
the basin (20) and a late Miocene series, mostly consisting of sandstones which constitute the
basement of the basin (S).
Field and laboratory studies are continuing as a development of this initial report.
Age of the calvarium
Our current conclusions on the chronological situation of the Ceprano hominid can now be
offered. Indications on its age are provided by the following geological data related to the basin
in filling.
(1) Because the calvarium layer contains no volcanic deposits, it is not directly datable. It is
also doubtful how it can be directly related to the different chronologies of the two volcanic
units (Figure 1: 3 and 4). hence, the only reference point to absolute age in the quaternary
series reported here (Figure 3) is that provided by the volcanics of the Ceccano and Pofi
eruptive district. The stratigraphic evidence shows that the clay (9) containing the calvarium is
older than the arrival of the fluvial volcanic sands (7): but, the leucite crystals, which constitute
a high proportion of those sands, may have ages that differ from the eruptive cycles, and thus
are probably mixed. The oldest have been dated 700 ka (Basilone Civetta, 1975; Fornaseri,
1985).
(2) The superimposed lower Acheulean layer (Figure 3; A1), which is correlated with the
K–Ar age of 458 5·7 ka of the Ranuccio–Anagni site (Biddittu et al., 1979; Segre Ascenzi,
1984) in the same Sacco river valley, is much higher in position than the calvarium, and is
separated by an unconformity from the aforementioned layers.
(3) The highly fossilized nature of the calvarium is in disagreement with the clay that
contains it. This finding is supported by its complete isolation, together with the total absence
of any accompanying organic remains. In accounting for the isolation of the calvarium on the
basis of its provenance from a partly destroyed layer, it should be noted that pebbles once
belonging to the underlying limnical series (Figure 3: 14–10) were eroded, and became

7.     HOMO ERECTUS  ,  415
Figure 4. Chopper artefact facies of Castro dei Volsci and Campo Grande sites; scale bar cm. 1, 6, bifacial
choppers; 2, subdiscoid nucleus and 3, nucleus; 4, convex scraper fragment; 5, denticolated scraper; 7, used
splint.
inclusions in the later layer (11). A similar event took place in a higher layer (4), where large
fragments of consolidated tufite are included in the sand-filled buried paleovalley (Figure 3: U).
The above examples, together with the former considerations, justify the conclusion that the
calvarium was most probably removed long ago from a much older deposit that is no longer
extant.
(4) The very evident overlying unconformity between 8–9 denotes an environmental change
that produced a swampy travertine facies.
(5) The underlying layer (13) with chopper industry and the faunal assembly age with E.
throgontherii show a greater age than the calvaria-bearing clay series (Figure 3: 9–11). Thus, the
minimum age of the calvarium must be estimated as older than 700 ka; the most probable age
can be estimated at rather even more than 800 ka, near or within the lower Cromer complex
limit. Lastly, it may be supposed that the calvaria was originally connected with the underlying
layer (Figure 3: 13) which has a chopper flake artefact assemblage Castro dei Volsci facies
(Figure 4).
Thus our conclusion is that this human fossil occurrence has a very substantial and evidently
Lower Pleistocene age.
Calvarium state of preservation and reconstruction
The bone fragments were recovered within loose soil; they were greyish, and were all highly
mineralized. Some of them presented distinctive morphological features, such as a browridge
or an occipital torus, which indicated that they belonged to a calvarium.
Many of the bone pieces fitted together rather well, especially those that had been
fragmented by the action of the bulldozer. It was impossible to determine the exact location of
many small pieces within the calvarium, but they did seem to pertain to the facial bones.

8. 416 .  ET AL.
The reconstruction of the calvarium was carried out in two stages. In the first the pieces
separated by recent fractures were assembled, and some bones (or bone fragments) isolated at
the level of sutures were put together.
The assembly of single fragments was, however, insufficient because fairly large gaps were
left at some sites. Various groups of bone fragments were provisionally fixed using small
plexiglas strips. These last also proved useful in fixing isolated bone fragments that had no
direct connection with the other pieces, but showed specific features indicating their likely
position within the gaps.
Special care was taken to place the frontal bone in the correct position, because it had no
direct connection with the remainder of the calvarium, due to the incompleteness of the
anterior portion of the two parietal bones, which consist of many separate small bone
fragments. To achieve this, the small pieces of the parietal bones were carefully examined on
their two surfaces (endo- and extracranial) so as to determine their exact degree of curvature.
Considering this feature, each piece was placed in what appeared to be the best possible
anatomical position. At this point, it was ascertained that the bone pieces were not completely
separated, but few contact points were present between the bone pieces, and between some of
these last and the frontal bone specially at level of internal or intermediate bone layers. In
consequence, the frontal bone was positioned in such a way as to be consistent with the
parietal bone pieces, and also to have an orientation corresponding to that of the occipital
bone. To achieve the last requisite a method resembling that previously developed for the
reconstruction of Arago 21 skull was applied (Ascenzi et al., 1986). It consisted of recording a
series of parallel lines using a pencil on the external surface of both the frontal and occipital
squama of the skull, the most central one corresponding to the median line of each bone. The
distance between any two successive lines was 5 mm. Using iron wires, it was verified that in
each of the two bones the corresponding lines ran in the same direction. The availability of the
final result of all the above operations was indirectly verified by comparing the length of the
glabella–inion axis of the Ceprano calvarium with that of other skulls from the same epoch.
The results indicate a high degree of resemblance. In particular, the glabella–inion distance of
the Ceprano calvarium was only 2 mm shorter than that of the Petralona skull.
The second stage in the reconstruction of the Ceprano calvarium consisted of the
progressive removal of the plexiglas strips, and the filling of gaps with plaster. Before the strips
were removed, a systematic CT-scan was carried out to carefully test the degree of correctness
of the reconstruction. In this way some mistakes were corrected.
Considering the restored Ceprano calvarium as a whole, its appearance is that of an
incomplete specimen. It comprises the frontal bone, which includes gaps, especially on the left
side, the incomplete parietal and temporal bones, the occipital bone, mainly consisting of the
squama, and the greater sphenoidal wings. The great care taken in reconstructing the
calvarium leaves no doubt that it displays a substantial distortion related to a congenital
malformation (see the later discussion about the orientation of the occipital torus), and,
possibly, partly due to a post-mortem deformation. As indicated in Figure 6, therefore, the
appearance of the calvarium is not strictly symmetrical.
Description of the calvarium
The two sides of the skull will be described jointly, although a preference is given to the
features observed on the right side, where the bones have been assembled in more compact
form than on the left side.

9.     HOMO ERECTUS  ,  417
Figure 5. The Ceprano calvarium as viewed in lateral (a) and in frontal norma (b).
In Figure 5(a) corresponding to the norma lateralis, the calvarium appears low. At the front
the torus frontalis and at the back the torus occipitalis are very prominent in the antero–
posterior direction. Thus the opisthocranion and inion coincide, and the maximum length,
which amounts to 208 mm, coincides with the glabella–inion axis. Behind the torus frontalis
there is almost no sulcus, and the outline of the calvarium rises toward the bregma. The exact
position of this last point is uncertain because of the erosion of the frontal squama along the
coronal suture. The rounded contour of the vault continues backward as far as the occipital
torus. A sulcus supratoralis is hardly appreciable on the occipital. Beyond the torus, the
contour bends sharply, forming a 115 angle between the occipital squama and the nuchal
plane.
The mid-sagittal craniogram shown in Figure 6 further establishes the lowness of the vault.
The bregma and the vertex are 64 mm and 66 mm, respectively, from the glabella–inion axis.

10. 418 .  ET AL.
Figure 6. Mid-sagittal craniogram.
In addition, Schwalbe’s angle, giving the inclination of the frontal bone, is 50 ; this turns out
to be very similar to the inclination angle of the occipital squama with respect to the
glabella–inion axis, measuring 51 .
It is impossible to exactly determine the dimensions of the coronal and temporal margins of
the parietal bones, so no certain description of the shape of these bones can be given. The
impression is that their height may exceed their length. In the posterior sector the temporal
line which emerges from an appreciable torus angularis hardly raises any real ridge. Fine radial
impressions of varying length are distributed near the temporal margin, indicating the origins
of small bundles of temporal muscle.
The temporal bone shows a low position within the calvarium. The squama appears
rounded and elongated with respect to its height. The mastoid is large. Its lateral surface lies
below the level of the supramastoid crest, extending prominently backward. In addition, it
clearly bends inward. On the medial side, the incisura mastoidea is wide along its whole
length. The supramastoid crest is connected to the root of the zygomatic arch. This arch is
missing, but on the right side the rather wide, upper surface of its root suggests that the arch
spanned a considerable portion of the fossa temporalis. The glenoid fossa is deep. Its anterior
part extends gradually into the preglenoid plane, while its posterior part is separated from the
incomplete external auditory meatus by the crista post-glenoidalis. The tympanic part of the
temporal bone is missing. The distance between both the glenoidal fossae amounts to 152 mm.
In norma frontalis [Figure 5(b)] the frontal torus constitutes a continuous, almost rectilinear,
bulky ridge that occupies not only the supraorbital margins, but also the glabella region, where
it bends downwards and somewhat backwards. The maximum height of the two supraorbital
portions is 20 mm, while the height of the glabella portion on the mid-line is 18 mm. The left
chamber of the frontal sinus occupies the entire breadth of the medial portion of the
corresponding supraorbital torus, while the right chamber constitutes only part of the breadth
of the medial portion of the corresponding torus. Behind the frontal torus the squama, which
is flattened on both sides, shows a slight median tuberosity. There is no trace of any keeling nor
of a metopic suture.
In the norma occipitalis [Figure 7(a)] the calvarium is low with respect to its breadth. On the
right side (where the superior preservation of the skull has permitted a more complete
reconstruction), starting from the middle line of the vault, the outline of the parietal bone

11.     HOMO ERECTUS  ,  419
reveals an initial slight slope followed laterally by an almost orthogonal bending, so that an
abrupt vertical orientation occurs. At the boundary between the parietal and the temporal
bone the vertical outline ends in a swelling that protrudes beyond the supramastoid crest. As
shown in Figure 8, this last feature can be demonstrated by examining the two superimposed
frontal half-craniograms recorded from the right side of the calvarium. The superior and
external half-craniogram (a) has been taken through the top of the swelling at the boundary
with the temporal squama, while the internal half-craniogram (b) skims the apex of the
supramastoid crest. It appears obvious that the swelling of the temporal squama, rather than
the supramastoid crest, is the highest point of the protrusion. The distance between the two
points is just over 2·5 mm. Below the supramastoid crest the calvaria outline bends
medianwards, closely following the re-entering contour of the mastoid. In contrast with the
right side, there is some doubt about the true point of maximum lateral prominence on the left
side, because of the distortion of the calvarium.
The occipital bone is broad and low. The asterion–asterion breadth is 133 mm and the
height of the squama 64 mm, so that the squama index is 48·1. The torus occipitalis appears
as a rather flat bulge with a smooth surface; it crosses the bone in an oblique direction. The
left portion is larger than the right, and definitely runs at a higher level, indicating a congenital
malformation. There is no direct connection between the torus and the supramastoid crest. An
occipital crest is located at the inion. The posterior portion of the nucal area is slightly concave.
The inion and endinion are not at the same level; the endinion lies as much as 22 mm under
the inion.
In the norma verticalis the strong bursoidal shape of the parieto–occipital outline, which is
reminiscent of the Javan Homo erectus calvaria (Weidenreich, 1943), contrasts with the well
developed, but not exceptionally large postorbital constriction [Figure 7(b)]. Besides this, the
calvarium displays a marked asymmetry due to distortion. As a result, the point of maximum
lateral protrusion corresponds to the middle portion of the skull outline on the right side, and
to the posterior portion on the left.
In the norma basalis the greater sphenoidal wings are thick and robust. The lateral end of
each wing occupies the angular space delimited by the frontal bone and the temporal squama.
The cavity corresponding to the facies cerebralis is restricted to a small but deep niche, with
little evidence of impressions and juga. A large lateral recess of a sphenoidal sinus extends far
into the left greater wing. The petrous portions of the temporal bones are badly damaged on
both sides. No surface details have survived. The long axis of the pyramid is hardly
recognizable, so its orientation appears hard to determine.
The cranial bones are unusually thick, particularly in the basal sectors. The average
thickness of the right parietal bone drops from 11·5 mm at the base to 8 mm at the vault. The
capacity of the calvarium, as measured by applying the millet seed method after meticulous
reconstruction of the cranial base is 1185 ml. The massive size of the bones suggests that the
calvarium can be attributed to a male. The open-ended sutures reveal an age that is probably
above 20 and below 40.
One paleopathologically acquired lesion is an oblique furrow in the right supraorbital torus.
This appears to be due to a healed depressed fracture.
Taxonomic affinities of the calvarium
In examining the position of the Ceprano calvarium in the context of human evolution, it
should be pointed out that the main features of this specimen are broadly comparable with

12. 420 .  ET AL.
Figure 7. The Ceprano calvarium as viewed in occipital (a) and in vertical norma (b).
those commonly present in Asian H. erectus. The cranial vault is low, with a flattened, retreating
forehead. The supraorbital ridges are massive and extremely prominent. They are continu-
ously connected to the glabellar torus, which is equally robust in structure. The opisthocranion
coincides with the inion, so that the glabella–inion distance corresponds to the maximum
sagittal length of the skull. There is a considerable angle between the occipital squama and the
nuchal plane, and the torus occipitalis lies at the vertex of this angle. The occipital squama is
very large when the asterion–asterion distance is compared with its height. The inion and
endinion are not at the same level; there is a large distance between them.
Apart from these features, which are to be considered among the most prominent of
H. erectus, other features are rather different from those of that hominid. There is no distinct

13.     HOMO ERECTUS  ,  421
Figure 8. Two right frontal half-craniograms touching the prominence of the temporal squama (a) and the
apex of the supramastoid crest (b).
sagittal keel or parasagittal depression in the frontal squama where, in contrast to the parietal
bones, the vault preserves its continuity. The endocranial capacity of the calvarium (1185 ml)
is greater than that of H. erectus, which has been defined as not much above 1000 ml
(Rightmire, 1990). This last change is a consequence of the reduction of the mastoid
protuberance as an outward bulge, the displacement of the utmost lateral salience of the skull
from the supramastoid crest to the temporal squama and an increased bending of the parietal
bone, with an attendant increase in its prominence. Other features that differentiate the
Ceprano calvarium from that of H. erectus sensu strictiori are the lessened post-orbital
constriction, the relative reduction in the massivity of the cranial bones of the vault with
respect to those of the base, and the development of the frontal sinuses.
At this point, the data provided by the European human fossils of the earlier Pleistocene
should be examined to determine their taxonomic affinities with the Ceprano calvarium. Until
recently, the oldest known inhabitants of Europe, some of whom were possibly as old as
500 ka, are represented by the fossils from Arago, Bilzingsleben, Mauer, Montmaurin,
Petralona and Vertesszöllös. They are similar to H. erectus in their angulated occipital bone
with transverse torus, robust supraorbital torus, heavy alveolar regions and robust mandibular
shape. These features have led several workers to place these specimens within H. erectus,
although the same hominids exhibit features, such as expansion of the cranial vault, decreased
postorbital constriction and less occipital angulation, which point towards more recent
European specimens and away from H. erectus. On this basis, the term late H. erectus has been
considered by us to be the most suitable for the Ceprano calvarium.

14. 422 .  ET AL.
The species designation Homo heidelbergensis was attributed to this group of fossils on the basis
of the site where the oldest sample was discovered.
The following group of more recent European hominids, despite their archaic morphology,
exhibit features which anticipate the later early Neandertals; the most representative
specimens are the calvaria from Steinheim and Swanscombe, and, to a lesser degree, those of
Atapuerca (Arsuaga et al., 1993).
Lastly, the group of early Neandertals from the penultimate glacial and last interglacial
period reveals a steady decrease in characteristics reminiscent of H. erectus and an increasing
prevalence of Neandertal features.
The reported sequence of hominids ranging from the H. erectus to the Neandertals has, very
recently been enriched by the discovery of fragmented human remains dating back more than
780 ka and discovered in the TD6 level of the Pleistocene cave site at Gran Dolina, Sierra de
Atapuerca (Aguirre et al., 1990; Carbonell et al., 1995). These fragments belong to at least four
individuals represented by cranial (a large piece of frontal squama), mandibular and dental
remains and are considered to belong to a primitive form of H. heidelbergensis, although,
according to Carbonell et al. (1995) this taxonomic name could be changed in the future as a
result of the substantial increase in the number of specimens. These new human fossil remains
demonstrate that Western Europe has been settled at least since the late early Pleistocene.
A thorough examination of the features of the Ceprano calvarium, as compared with those
of the series of hominids from the lower and middle Pleistocene in Europe appears to justify
the conclusion that the Ceprano calvarium belonged to a late H. erectus. There is much less
justification for considering the same remain as pertaining to the species called H. heidelbergensis,
because the Mauer mandible shows absolutely no correspondence to the Ceprano calvaria.
Instead, a surprisingly good fit associates the Ceprano calvarium with mandible 3 from
Ternifine (Morocco). This occurrence could be fortuitous, but it could reveal a relationship
between the Ceprano hominid and the early Pleistocene hominids from North Africa, an issue
that now deserves attention in view of further discoveries and investigations.
Acknowledgements
The authors wish to thank Dr L. Sadori of the Section of Paleobotany, ‘‘La Sapienza’’
University, Rome, for the pollen analysis; Mr G. Bretzel for the photographs of the calvaria;
Mr A. Benvenuti and Mr L. Virgilii for technical assistance.
References
Aguirre, E., Arsuaga, J. L., Bermúdez de Castro, J. M., Carbonell, E., Ceballos, M., Díez, C., Enamorado, J.,
Fernández-Jalvo, Y., Gil, E., Garcia, A., Martín-Nájera, A., Martínez, I., Morales, J., Ortega, A. I., Rosas, A.,
Sánchez, B., Sesé, C., Soto, E. Torres, T. J. (1990). The Atapuerca Sites and the Ibeas Hominids. Hum. Evol. 5,
55–73.
Angelucci, A., Brotzu, P., Civitelli, G., Morbidelli, L. Traversa, G. (1974). The pleistocene volcanism of the middle
Latine valley, petrographic features of the chief lava crops (In Italian). Geol. Romana 13, 83–123.
Arsuaga, J.-L., Martinez, I., Garcia, A., Carretero, J.-M. Carbonell, E. (1993). Three new human skulls from the
Sima de los Huesos Middle Pleistocene site in Sierra de Atapuerca, Spain. Nature 362, 534–537.
Ascenzi, A., Marchetti, A. M. Micheli, M. (1986). Comparison between the Tautavel hominid, the italian
anteneandertals and the Saccopastore hominid (In French). L’Anthropologie (Paris) 90, 515–537.
Basilone, P. Civetta, L. (1975). The volcanic activity of the Monti Ernici dated by the K–Ar method (In Italian). Bull.
Soc. It. Mineral Petrol. 31, 175–179.
Bergomi, G. Nappi, G. (1973). About some volcanites of the high and middle valley of the Sacco river, Southern
Latium (In Italian). Bull. Serv. Geol. It. 94, 47–71.

15.     HOMO ERECTUS  ,  423
Biddittu, I. (1972). Pleistocene and pre-acheulean artefacts at Arce and Fontana Liri. Quaternaria 16, 35–52.
Biddittu, I. (1974a). Pleistocene layer with acheulean bifacials in the district of Ceprano, Frosinone (In Italian). Mem.
Ist. It. Paleont. Umana 2, 61–67.
Biddittu, I. (1974b). Pre-acheulean layers at Castro dei Volsci, Frosinone (In Italian). Mem. Ist. It. Paleont. Umana 2,
51–60.
Biddittu, I. Cassoli, P. F. (1969). A lower paleolithic site at Pontecorvo in the province of Frosinone (In Italian).
Quaternaria 10, 167–197.
Biddittu, I. Segre, A. G. (1978). A lower pleistocene site at Pompi quarry near Pofi, Frosinone (In Italian). Archeol.
Etrusco-Laziale 1, 78–79.
Biddittu, I. Segre, A. G. (1982a). A middle-lower pleistocene site with archaic artefacts from pebbles in the Anagni
basin, Latium (In Italian). Atti Riun. Sc. Ist. It. Preist. Protost. Firenze 23, 567–576.
Biddittu, I. Segre, A. G. (1982b). Bone utilization during the italian lower paleolithic era (In Italian). Atti Riun. Sc.
Ist. Preist. Protost. Firenze 23, 89–105.
Biddittu, I. Segre, A. G. (1984). Splint artefacts and bifacials: new findings of the lower Paleolithic at
Anagni-Fontana Ranuccio, Frosinone (In Italian). Atti Riun. Sc. Ist. It. Preist. Protost. Firenze 24, 105–112.
Biddittu, I., Cassoli, P. F., Radicati, F., Segre, A. G., Segre-Naldini, E. Villa, I. (1979). Anagni, a K–Ar dated lower
and middle Pleistocne Site, Central Italy, preliminary report. Quaternaria 21, 53–71.
Blanc, A. C. (1956). The pleistocene layer of Pofi, Frosinone (In Italian). Quaternaria 2, 261–262.
Blanc, A. C. Taschini, M. (1958–1961) Excursion and excavation campaign in the Pofi district, Frosinone (In
Italian). Quaternaria 5, 335–336.
Branco, W. (1877). The Ernici volcanoes in the Sacco valley (In Italian). Atti R. Accad. Lincei Serie 3, 1, 811–817.
Bruni, L. (1987). Discovery of a bone bifacial in the lower Paleolithic at Fontana Ranuccio, Anagni (In Italian). Latium
4, 3–9.
Carbonell, E., Bermúdez de Castro, J. M., Arsuaga, J. L., Díez, J. C., Rosas, A., Cuenca-Bescós, G., Sala, R.,
Mosquera, M. Rodríguez, X. P. (1995). Lower Pleistocene, Hominids and Artefacts from Atapuerca-TD6
(Spain). Science 269, 826–830.
Cassoli, P. F. Segre-Naldini, E. (1984). New contribution to the acquaintance of the Villafranchian and Pleistocene
fauna in the Anagni basin, Frosinone (In Italian). Atti Riun. Sc. Ist. It. Preist. Protost. Firenze 24, 115–118.
Cassoli, P. F. Segre-Naldini, E. (1995). The Villafranchian in the Latium region (In Italian). Atti Convegno sul
Villafranchiano—AIQUA (In press).
Civetta, L., Innocenti, F., Manetti, P., Peccerillo, A. Poli, G. (1981). Geochemical characteristics of potassic
volcanoes from Mounts Ernici, Southern Latium, Italy. Contrib. Mineral Petrol. 78, 37–47.
De Lorenzo, G. D’Erasmo, G. (1932). The Paleolithic man and the Elephas antiquus in the Southern Italy (In Italian).
Atti R. Accad. Sc. Fis. Mat. Napoli S.2, 19, 1–107.
Devoto, G. (1965). Lacustrine Pleistocene in the lower Liri Valley. Geol. Romana 4, 291–368.
Fedele, P. (1962). Prospection in Pofi district from June 1956 to December 1958 (In Italian). Quaternaria 5, 321–322.
Fornaseri, M. (1985). Geochronology of volcanic rocks from Latium, Italy. Rend. Soc. It. Mineral Petrolog. 40, 74–106.
Nicolucci, G. (1868). Human antiquity in Central Italy (In Italian). Rend. R. Accad. Sc. Fis. Mat. Napoli 7, 130–135.
Nicolucci, G. (1873). The stone age in the Neapolitan provinces (In Italian). Rend. R. Accad. Sc. Fis. Mat. Napoli 12,
19–22.
Ponzi, G. (1857–1858). On the discovery of the extinct Ernici volcanoes in Latine valley (In Italian) Atti Pontif. Accad.
Nuovi Lincei 11, 51–62.
Rightmire, G. P. (1990). The Evolution of Homo erectus. Cambridge: Cambridge University Press.
Segre, A. G. Ascenzi, A. (1984). Fontana Ranuccio: Italy’s earliest middle Pleistocene hominid site. Current
Anthropology 24, 230–233.
Segre, A. G. Biddittu, I. (1981). Lower Paleolithic: the oldest marks of the human activity in the Sacco valley (In
Italian). Archeol. Etrusco-ital. 4, 26–34.
Segre, A. G., Biddittu, I. Cassoli, P. F. (1984). The Sora paleolacustrine basin (Frosinone district) and its musterian
layers (In Italian). Atti Riun. Sc. It. Preist. Protost. Firenze 24, 149–154.
Settepassi, F. Verdel, U. (1985). Continental quaternary mollusca of lower Liri valley, Southern Latium. Geologica
Romana 4, 369–451.
Viola, C. (1902). The chief types of lava in the Ernici volcanoes (In Italian). Boll. R. Com. Geol. Ital. 32, 104–148.
Weidenreich, F. (1943). The skull of Sinanthropus pekinensis: a comparative study of a primitive hominid skull.
Palaeontologia Sinica, New Series D 10, 1–484.