A. Ascenzi*,                            A calvarium of late Homo erectus from
I. Biddittu,                            Cepr...

    0                         5

              HOMO ERECTUS        ,                             411

   Figure 2. Compo...
412                                     .  ET AL.

   Late neotectonic faults altered the Pleistocene hydrography,...
             HOMO ERECTUS       ,                            413

414                                      .  ET AL.

   The fifth ensemble (Figure 3; 12–13) has a variable thicknes...
            HOMO ERECTUS       ,                          415

   Figure 4. Chopper art...
416                                     .  ET AL.

   The reconstruction of the calvarium was carried out in two s...
           HOMO ERECTUS       ,                 417

             Figure 5. The Ceprano...
418                                      .  ET AL.

                                  Figure 6. Mid-sagittal cr...
        HOMO ERECTUS    ,                    419

reveals an initial slight slope followed...
420                                         .  ET AL.

            Figure 7. The Ceprano calvarium as viewed in...
           HOMO ERECTUS      ,                        421

   Figure 8. Two right front...
422                                               .  ET AL.

   The species designation Homo heidelbergensis was a...
             HOMO ERECTUS        ,                             423

Biddittu, I. (1972). P...
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A calvarium of late homo erectus from ceprano, italy (ascenzi et al.)


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  1. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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. 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.
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