1. News and Views A re-appraisal of Ceprano calvaria affinities
with Homo erectus, after the new
reconstruction
A. Ascenzi, Italian Institute of Human Paleontology, Piazza Mincio 2, 00198
Rome, Italy. E-mail: bonucci@axrma.uniroma1.it
F. Mallegni,
G. Manzi,
A. G. Segre &
E. Segre Naldini
Keywords: Homo erectus, Journal of Human Evolution (2000) 39, 443–450
cranial morphology, Lower doi:10.1006/jhev.2000.0425
Paleolithic, Ceprano, Italy. Available online at http://www.idealibrary.com on
Introduction Research in the Priverno Basin provides a
Following the reconstruction of the Ceprano chronological outline for the lower part of
calvaria by Clarke (2000), additional minor the stratigraphic series in the central
improvements were made by M. A. de Ceprano Basin, including sediments that
Lumley (with the assistance of C. Lanoux) contain a chopper industry hypothesized
and by one of us (F.M., with the assistance to be broadly contemporaneous with the
of S. Ricci). The present paper provides a hominid. A lagoon deposit, considered to
brief account of these modifications, to- be equivalent to the limnic series at
gether with a reassessment of the mor- Ceprano, includes thin layers of volcani-
phology and morphometrics of the Ceprano clastic augite-biotite sands, identical to
calvaria. This allows an opportunity to those in the Ceprano Basin. During the
critically evaluate some of the Homo erectus Early Pleistocene both basins were con-
characteristics that have been previously re- nected through a common paleohydro-
ported (Ascenzi et al., 1996, 1998; Ascenzi graphic drainage system. This volcaniclastic
& Segre, 1997a,b). We also introduce new layer has an Ar/Ar date, based on biotite, of
data on the geochronology of the site. 1060 110 ka (Rammelzwaal, 1978; Sevink
et al., 1984). The outcrop that contained the
fossil hominid, which was subsequently
Further consideration of the enclosed in a paleocolluvial clay layer
geochronology (Ascenzi et al., 1996; Ascenzi & Segre,
The circumstances of recovery of the speci- 1997a), must be younger than these dates.
men and the geological and archaeological An age of 800–900 ka provides a reasonable
context of the site have been described else- estimate, and one that corresponds closely
where (Ascenzi et al., 1996, 1998; Ascenzi & to the previous correlation of the hominid
Segre, 1997a,b, 2000). These papers report with the lower part of the Cromerian
K/Ar dates obtained from strata overlying complex (Azzaroli, 1951; Turner, 1996).
the horizon from which the calvaria was
recovered. Further geological investigations
Further modifications to the calvaria
in the Priverno Basin, situated about 25 km
from Ceprano, provide independent evi- Having been invited to prepare a mould
dence for estimating the age of the calvaria for casting, M. A. de Lumley removed
and its associated sediments. dental plaster from the calvaria. During the
0047–2484/00/100443+08$35.00/0 2000 Academic Press

2. 444 .  ET AL.
Figure 1. The new reconstruction of the Ceprano calvaria as viewed in right norma lateralis.
Figure 2. The new reconstruction of the Ceprano calvaria as viewed in left norma lateralis.
process, she realized that a large fragment of bone (Figures 1 and 2). Identification of
parietal was incorrectly oriented, and that bregma has allowed us to establish the mid-
it represented the medio-anterior corner of sagittal plane of the cranial vault. In
the right parietal joining the frontal bone addition, de Lumley added two previously
along the first third of the coronal suture. unidentified bone fragments to the recon-
Its correct placement, later refined by F. struction that represent the frontal processes
Mallegni, served to verify the connection of the zygomatic bones (Figures 1 and 2).
between the right parietal and the frontal The modification was carried out by

3.    445
Mallegni, who also recognized that a further maximum breadth at the level of the crista
fragment completed the occipital along supramastoidea.
the sutural margin with the right parietal In superior view, the frontal has a massive
(Figure 1). and continuous supraorbital torus, and
With the current reassembling of the the calvaria appears bursoidal in shape.
calvaria, all available portions of each This latter feature was described by von
bone connect directly to others. In this way, Koenigswald Weidenreich (1939) and
the cranial structure can be confidently Weidenreich (1943) as being characteristic
reconstructed. Evidence on this point of H. erectus skulls from Java, and is con-
has been provided by the final touches of sidered distinctive with respect to the
Mallegni, who was able to show that only a material from Zhoukoudian. In contrast,
few drops of glue are sufficient to join the a frontal trait that distinguishes Ceprano
cranial portions, and only small plaster from the more typical representatives of
reinforcements are necessary to support the H. erectus, both from Asia and Africa, is
assembled calvaria. the rather limited extent of the post-
orbital constriction. The index of minimum
frontal breadth/biorbital breadth is 84·8,
which is relatively high in comparison to
New insights into the morphology and
OH9 (81·3), for example. The enlargement
taxonomy of the Ceprano hominid
of the frontal bone (minimum/maximum
Tables 1 and 2 provide an assessment, both frontal breadth) gives a value of 89·8, which
metrical and descriptive, of the morphologi- is the highest value reported in Table 1.
cal affinities of the Ceprano calvaria. Data By looking at the coronal section of the
reported in the tables can be compared with best preserved right side in posterior view,
those available in the literature, in order to the maximum breadth of the skull occurs
ascertain its affinities to H. erectus. Obvi- at the level of crista supramastoidea, after
ously, additional comparisons will need to which it narrows slightly towards the
be made with other European taxa, particu- angled parietal (nearly 105 ) at the level
larly H. heidelbergensis (Rightmire, 1998) of the superior temporal line. The reduc-
and H. antecessor (Bermudez de Castro et al., tion in width then increases rapidly as the
1997). profile approaches the sagittal suture,
Morphologically and metrically, the where there is no evidence of sagittal
Ceprano calvaria approaches the mor- keeling.
phology in H. erectus, but the degree of The temporal squama rises superiorly,
variation has to be enlarged if the specimen with its parietal margin following a high
is included in the hypodigm. For example, if curve that is somewhat anteriorly elongated.
we consider the features listed in Table 2, The form of the squama is quite differ-
only two-thirds of the character states corre- ent from the Zhoukoudian sample
spond (and not always unequivocally) with (Weidenreich, 1943). It looks more like a
the list of distinctive features generally rec- right-angled triangle, with the sphenoid
ognized as typical of H. erectus (Wood, margin forming the shorter cathetus, the
1991). H. erectus is characterized by a long parietal margin the longer one, and the
and low cranial vault, whereas the Ceprano zygomatic process and its continuation
specimen shows a distinctive profile in along the supramastoid crest the hypo-
which the cranial vault is low (compare data tenuse. The remaining components of the
in Table 1) but relatively short, with a temporal bone are badly damaged. Only
conspicuous medio-lateral expansion and the roof of the auditory canal is intact on

4. 446
Table 1 Metrical comparisons of the Ceprano calvaria (new reconstruction) with Homo erectus specimens from East Asia and Africa
Ceprano Sang.2 Sang.17 Sambung. Zkd.X Zkd.XI Zkd.XII ER 3733 ER 3883 OH 9
Whole vault
M.1 Maximum cranial length 198 206 200 199 192 195 182 182 205
M.11b Biauricular breadth (151) 126 140 137 147 143 151 132 129 135
M.20 Auriculo-bregmatic height (105)
W.12 Supramastoid breadth (161) 141 (?) 161 151 (?) (150) 145 (147) 142 139 146
W.15 Interentoglenoid breadth 109 83 86 89
Breadth/length cranial index (W.12/M.1 100) (81·3) 78·2 75·5 (75·4) 75·5 (75·4) 78 76·4 71·2
Face
M.44 Biorbital breadth (125) 115 (114) (104) 108 (123)
Frontal bone
W.62 Supraorbital vertical thickness (L) 21 [12] 18 16·5 13·2 14·6 8·5 13 18
.  ET AL.
M.9 Minimum frontal breadth (106) 82 99 102 89 84 91 91 88 (100)
M.10 Maximum frontal breadth (118) 102 119 116 (110) 106 108 110 105
M.26 Frontal sagittal arc 118 129 122 124 (119) 118
M.29 Frontal sagittal chord 106 (118) (115) 115 106 113 (104) 101
M.32(5) Frontal angle (138) 139 140
Frontal profile (g–m g–i/op)* 60 55 63 61 56
Frontal breadth index (M.9/M.10 100) (89·8) 80·4 83·2 87·9 (80·9) 79·2 84·3 (82·7) 83·8
Postorbital constriction index (M.9/M.44 100) (84·8) 86·1 (89·5) (87·5) 81·5 (81·3)

5. Table 1 Continued
Ceprano Sang.2 Sang.17 Sambung. Zkd.X Zkd.XI Zkd.XII ER 3733 ER 3883 OH 9
Parietal and temporal bones
M.8 Biparietal maximum breadth 156 137 144 146 137 136 140 127 132 138
M.27 Parietal sagittal arc (98) (103) 109 102 113 92 102 (83) 93
M.30 Parietal sagittal chord 95 (98) 106 96 106 86 91 (79) 86
M.27(3) Lambda–asterion arc (L) 112 (92) 91 93 (99) 92 83 85
Lambda–asterion arc (R) 103 98 82
M.30(3) Lambda–asterion chord (L) 98 (82) 87 (79) 85 (84) 87 78 76
Lambda–asterion chord (R) 92 88 77
W.7 Mastoid height (R) (27) 12 25 20 (23) 30
Occipital bone
M.12 Biasterionic breadth 128 122 142 (111) 113 115 124 121 123
M.31(1) Lambda–inion chord 62 (45) 58 48 52·5 61 49 (54)
Upper scale inclination (l–i g–i/op)* 58 62·5 68 57 61
Inion–endinion distance 22 38 34 35
Occipital upper-scale index (M.31·1/M.12 100) 48·4 (36·9) 40·8 42·5 45·7 49·2 40·5 (43·9)
  
All measurements were made by three different authors on separate occasions on the Clarke’s reconstruction, after the minor additional improvements were
carried out; averages are reported where appropriate (differences less than 10 mm); definitions and numbering follow Martin Saller (1957; M.n) or,
alternatively, Wood (1991; W.n); measurements of length are in mm and those of angles in degrees; measurements involving an evaluation of the exact position
of the reference point/s are enclosed in parentheses; L=left; R=right; Sang.=Sangiran; Sambung.=Sambungmachan; Zkd.=Zoukoudian; ER=East Turkana;
OH=Olduvai; comparative data as reported by Weidenreich (1943), Rightmire (1990) and Wood (1991).
* Denotes the angle formed by the two chords. Inion and opisthocranion are coincidental in Homo erectus.
447

6. Table 2 Character states of the Ceprano calvaria according to the list of distinctive features of Homo erectus by Wood (1991: Table 2.11, p. 37)
448
State Comments
Cranial shape and size
Overall 1a Long cranial vault No See metrical comparisons in Table 1
1b Low cranial vault Yes See metrical comparisons in Table 1
2 Maximum breadth across the angular torus or supramastoid crest Yes Supramastoid crest
3 Thick vault bones Yes e.g., 16 mm at parietal eminence (R)
4 Pronounced postorbital constriction No Slightly expressed
Individual bones
Frontal 6 Frontal keel or ridge (No) Distinct median bump of the frontal squama in spite of a flat,
receding forehead
7 Straight junction of torus and frontal squama No Given the presence of a supratoral sulcus, the plane above the torus
is concave, particularly above glabella
8 Coronal ridge No
Parietal 9 Flattened and rectangular parietal (No) Gently curved; square in shape
Temporal 11 Low temporal squama No Estimated height
12 Small mastoid process No
Occipital 13 Opisthocranion coincident with inion Yes
14 Sharply angulated Yes
16 Discrepancy between inion and endinion Yes
Nasals 17 Broad nasal bones (Yes) Probable, given the massive aspect of the fronto-nasal suture (L)
Specific morphological features
.  ET AL.
Frontal 18 Large, continuous supraorbital ridges with a supratoral sulcus Yes
19 Lateral wing to supraorbital torus (Yes) Not anteriorly projected
Parietal 20 Prominent angular torus at mastoid angle Yes Well expressed, although not particularly prominent
Temporal 21a Marked supramastoid crests Yes
21b Marked mastoid crests No Little relief of the surface of the mastoid process is visible, without a
clear supramastoid sulcus
22 Occipitomastoid ridge (Yes) A ridge medial to the digastric fossa on both sides of the occipito-
mastoid suture is present
23 Juxtamastoid ridge absent Yes
24 Marked suprameatal tegmen (Yes) Present, but not particularly marked
Occipital 26a Occipital torus, with supratoral sulcus Yes
26b Occipital torus continuous with angular torus and supramastoid No Torus confirmed to the occipital
crest
27 Supernumerary bones at lambda No
Endocranial 31 Large posterior branch of middle meningeal artery and vein Yes
Parentheses indicate observation uncertain or equivocal. Note that some traits (1, 21 and 26) have been divided with respect to the original list by Wood (1991), in order
to better describe the features encountered in the Ceprano calvaria, and others (5, 10, 15, 25 and 28–30) have been omitted because of the state of preservation of the
specimen (absence of the region involved). R=right; L=left.

7.    449
the right side, while the tympanic plates of of the braincase. Nevertheless, the geo-
both sides have been lost. The severely dam- graphical provenience has to be taken into
aged petrous portions are short stumps, and consideration when evaluating distinctive
their orientation is difficult to discern. traits, such as the bursoidal shape of the
The general form of the occipital squama skull, the limited extent of the postorbital
is defined by the relationship between its constriction, and the absence of a connec-
maximum (biasterionic) breadth and the tion between torus occipitalis and crista
lambda–inion chord. The index furnished supramastoidea.
by these two dimensions is 49·2, and this
indicates a relatively short length for the
upper scale of the occipital. Furthermore, Acknowledgements
the bone is clearly angled in lateral view (see We are greatly indebted to R. J. Clarke and
Table 1), with a prominent torus occipitalis M. A. de Lumley for their valuable and
and a nuchal plane that faces backward and generous help. We also wish to thank A.
inferiorly. Inion and endinion are not posi- Benvenuti, P. Cassoli, F. Di Baldo, C.
tioned at the same level, with endinion lying Lanoux, S. Ricci, and L. Virgilii for their
as much as 22 mm below inion. skilful technical assistance.
In addition, one important observation
that has not been given sufficient attention is
the thickness of the vault bones in the References
Ceprano specimen. Thickness varies from Ascenzi, A. Segre, A. G. (1997a). Resti di cranio
8·0 mm (measured at lambda on the occipi- umano del Pleistocene medio-inferiore a Ceprano.
tal, following Wood, 1991:n. 113) to 24 mm Rend. Sc, Fis. Nat., Accad. Naz. Lincei S 9, 8, 39–67.
Ascenzi, A. Segre, A. G. (1997b). Discovery of a
(at the external occipital protuberance, Homo erectus calvarium at Ceprano, Central Italy.
Wood n. 115), while the figure obtained in Anthropologie (Brno) 35, 241–246.
the right parietal eminence region (Wood n. Ascenzi, A. Segre, A. G. (2000). The fossil hominid
from Ceprano, Central Italy. In (M. Aloisi, B.
111) is 16 mm. The exceptional thickness of Battaglia, E. Carafoli G. A. Danieli, Eds) The
the bone at the base of the calvaria is noted Origin of Humankind. IVSLA series, vol. II. Venice
by Clarke (2000). Examination of the inter- and Amsterdam: IVSLA IOS.
nal structure of the cranial thickening, as Ascenzi, A., Biddittu, I., Cassoli, P. F., Segre, A. G.
Segre-Naldini, E. (1996). A calvarium of late
observed directly on the broken ones or by Homo erectus from Ceprano, Italy. J. hum. Evol. 31,
X-rays, yielded ambiguous results because 409–423.
fossilization made the boundaries between Ascenzi, A., Biddittu, I., Cassoli, P. F., Segre, A. G.
Segre-Naldini, E. (1998). A calvaria of a Lower
the tables and the diploe difficult to discern.
¨ Pleistocene hominid from Southern Latium, Central
The cranial capacity, measured using the Italy. In (J. Gibert, F. Sanchez, L. Gibert F. Ribot,
millet grain method, is 1057 ml—a value Eds) The Hominids and their Environment
during the Lower and Middle Pleistocene of Eurasia.
within the H. erectus range, according to Proceedings of the International Conference, Orce
Rightmire (1990). 1995. 287–297. Orce: Museo ‘‘J. Gibert’’.
In conclusion, if the Ceprano specimen Azzaroli, A. (1951). The geological age of the Cromer
forest bed. Proc. Prehist. Soc. N.S. 17, 168–170.
is classified as H. erectus it represents the Bermudez de Castro, J. M., Arsuaga, J. L., Carbonell,
earliest specimen of this taxon in Europe. E., Rosas, A., Martinez, I. Mosquera, M. (1997).
The chronology, assuming an age between A hominid from the lower Pleistocene of Atapuerca:
Possible ancestor to Neandertals and modern
800 and 900 ka as the most probable, is humans. Science 276, 1392–1395.
consistent with its archaic morphology, Clarke, R. J. (2000). A corrected reconstruction and
especially the massive continuous frontal interpretation of the Homo erectus calvaria from
Ceprano, Italy. J. hum. Evol. 39, 433.
torus, sharply angled occipital bone, small Martin, R. Saller, K. (1957). Lehrbuch der
cranial capacity, and exceptional thickness Anthropologie, Bd. I.. Stuttgart: G. Fisher.

8. 450 .  ET AL.
Rammelzwaal, A. (1978). Soil genesis and Quaternary Turner, C. (1996). The Early Pleistocene in Europe.
landscape development in the Tyrrhenian coastal Proceed. S.E.Q.S. Cromer Symposium 1–329.
area of South-Central Italy. Publ. Fysisch-Geogr. Norwich, U.K., 1990.
Bodenkundig Lab. Univ. Amsterdam 28, 1–310. Von Koenigswald, G. H. R. Weidenreich, F. (1939).
Rightmire, G. P. (1990). The Evolution of Homo The relationship between Pithecanthropus and
erectus. Cambridge: Cambridge University Press. Sinanthropus. Nature 144, 926–929.
Rightmire, G. P. (1998). Human evolution in the Weidenreich, F. (1943). The skull of Sinanthropus
Middle Pleistocene: the role of Homo heidelbergensis. pekinensis: a comparative study on a primitive
Evol. Anthropol. 6, 218–227. hominid skull. Palaeontologia Sinica, New Series
Sevink, J., Rammelzwaal, A. Spaargaren, O. C. D.10, 1–484.
(1984). The soils of Southern Lazio and adjacent Wood, B. (1991). Koobi Fora Research Project. Hominid
Campania. Publ. Fysisch-Geogr. Bodenkundig. Lab. Remains. Vol. 4. Oxford: Clarendon Press.
Univ. Amsterdam 38, 1–144.