Papers by Clément Zanolli
Keywords: X-ray microtomography, dental virtual histology, human dental enamel, evolution of huma... more Keywords: X-ray microtomography, dental virtual histology, human dental enamel, evolution of human life history Summary: A sample of human deciduous teeth from the Roman Imperial necropolis of Velia (I-II cent. CE) was measured by high-resolution, phase-contrast synchrotron X-ray microtomography at the SYRMEP beamline of the Elettra laboratory. Results show the suitability and potential of the setup available at SYRMEP in paleo-ontogenetic studies on tooth enamel, through visualization of the growth markers in the mineralized tissue.
The characterization and quantification of human dental enamel microstructure, in both permanent ... more The characterization and quantification of human dental enamel microstructure, in both permanent and deciduous teeth, allows us to document crucial growth parameters and to identify stressful events, thus contributing to the reconstruction of the past life history of an individual. Most studies to date have focused on the more accessible post-natal portion of the deciduous dental enamel, even though the analysis of prenatal enamel is pivotal in understanding fetal growth, and reveals information about the mother's health status during pregnancy. This contribution reports new data describing the prenatal enamel development of 18 central deciduous incisors from the Imperial Roman necropolis of Velia (I-II century CE, Salerno, Italy). Histomorphometrical analysis was performed to collect data on prenatal crown formation times, daily secretion rates and enamel extension rates. Results for the Velia sample allowed us to derive a new regression formula, using a robust statistical approach, that describes the average rates of deciduous enamel formation. This can now be used as a reference for pre-industrial populations. The same regression formula, even when daily incremental markings are difficult to visualize, may provide a clue to predicting the proportion of infants born full term and pre-term in an archaeological series.
Objectives: The aim of this study is to compare the degree and patterning of inter-and intra-indi... more Objectives: The aim of this study is to compare the degree and patterning of inter-and intra-individual metameric variation in South African australopiths, early Homo and modern humans. Metameric variation likely reflects developmental and taxonomical issues, and could also be used to infer ecological and functional adaptations. However, its patterning along the early hominin postcanine dentition, particularly among South African fossil hominins, remains unexplored. Materials and Methods: Using microfocus X-ray computed tomography (mXCT) and geometric morphometric tools, we studied the enamel-dentine junction (EDJ) morphology and we investigated the intra-and inter-individual EDJ metameric variation among eight australopiths and two early Homo specimens from South Africa, as well as 32 modern humans. Results: Along post-canine dentition, shape changes between metameres represented by relative positions and height of dentine horns, outlines of the EDJ occlusal table are reported in modern and fossil taxa. Comparisons of EDJ mean shapes and multivariate analyses reveal substantial variation in the direction and magnitude of metameric shape changes among taxa, but some common trends can be found. In modern humans, both the direction and magnitude of metameric shape change show increased variability in M 2-M 3 compared to M 1-M 2. Fossil specimens are clustered together showing similar magnitudes of shape change. Along M 2-M 3 , the lengths of their meta-meric vectors are not as variable as those of modern humans, but they display considerable variability in the direction of shape change. Conclusion: The distalward increase of metameric variation along the modern human molar row is consistent with the odontogenetic models of molar row structure (inhibitory cascade model). Though much remains to be tested, the variable trends and magnitudes in metamerism in fossil hominins reported here, together with differences in the scale of shape change between modern humans and fossil hominins may provide valuable information regarding functional morphology and developmental processes in fossil species.
MicroCT imaging is increasingly used in paleoanthropological and zooarchaeological research to an... more MicroCT imaging is increasingly used in paleoanthropological and zooarchaeological research to analyse the internal microstructure of bone, replacing comparatively invasive and destructive methods. Consequently the analytical potential of this relatively new 3D imaging technology can be enhanced by developing discipline specific protocols for archaeological analysis. Here we examine how the microstructure of mammal bone changes after burning and explore if X-ray computed microtomography (microCT) can be used to obtain reliable information from burned specimens. We subjected domestic pig, roe deer, and red fox bones to burning at different temperatures and for different periods using an oven and an open fire. We observed significant changes in the three-dimensional microstructure of trabecular bone, suggesting that biomechanical studies or other analyses (for instance, determination of age-at-death) can be compromised by burning. In addition, bone subjected to very high temperatures (600°C or more) became cracked, posing challenges for quantifying characteristics of bone microstructure. Specimens burned at 600°C or greater temperatures, exhibit a characteristic criss-cross cracking pattern concentrated in the cortical region of the epiphyses. This feature, which can be readily observed on the surface of whole bone, could help the identification of heavily burned specimens that are small fragments, where color and surface texture are altered by diagenesis or weathering.
MicroCT analysis of archaeozoological remains: a non-destructive approach to the age estimation
Journal of Human Evolution, 2014
Fieldwork performed during the last 15 years in various Early Pleistocene East African sites has ... more Fieldwork performed during the last 15 years in various Early Pleistocene East African sites has significantly enlarged the fossil record of Homo erectus sensu lato (s.l.). Additional evidence comes from the Danakil Depression of Eritrea, where over 200 late Early to early Middle Pleistocene sites have been identified within a w1000 m-thick sedimentary succession outcropping in the Dandiero Rift Basin, near Buia. Along with an adult cranium (UA 31), which displays a blend of H. erectus-like and derived morphoarchitectural features and three pelvic remains, two isolated permanent incisors (UA 222 and UA 369) have also been recovered from the 1 Ma (millions of years ago) Homo-bearing outcrop of Uadi Aalad. Since 2010, our surveys have expanded to the nearby (4.7 km) site of Mulhuli-Amo (MA). This is a fossiliferous area that has been preliminarily surveyed because of its exceptional concentration of Acheulean stone tools. So far, the site has yielded 10 human remains, including the unworn crown of a lower permanent molar (MA 93). Using diverse analytical tools (including high resolution mCT and mMRI), we analysed the external and internal macromorphology and microstructure of the three specimens, and whenever possible compared the results with similar evidence from early Homo, H. erectus s.l., H. antecessor, H. heidelbergensis (from North Africa), Neanderthals and modern humans. We also assessed the UA 369 lower incisor from Uadi Aalad for root completion timing and showed that it compares well with data for root apex closure in modern human populations.
New dental remains of Anoiapithecus and the first appearance datum of hominoids in the Iberian Peninsula
Newdental remains of the fossil great ape Anoiapithecus brevirostris are described from the Middl... more Newdental remains of the fossil great ape Anoiapithecus brevirostris are described from the Middle Miocene local stratigraphic series of Abocador de Can Mata (ACM) in els Hostalets de Pierola (Vallès-Penedès Basin, NE Iberian Peninsula). These specimens correspond to maxillary fragments with upper teeth from two female individuals from two different localities: left P3eM1 (IPS41712) from ACM/C3-Aj (type locality; 11.9 Ma [millions of years ago]); and right M1eM2 and left P4eM2 (IPS35027) from ACM/C1-E* (12.3 e12.2 Ma). Relative enamel thickness is also computed in the latter individual and re-evaluated in other Middle Miocene hominoids from ACM, in order to better assess their taxonomic affinities. With regard to
maxillary sinus development, occlusal morphology, molar proportions and enamel thickness, the new specimens show greater resemblances with the (male) holotype specimen of A. brevirostris. They differ from Pierolapithecus catalaunicus in displaying less inflated crests, a more lingually-located hypocone, and relatively lower-crowned molars; from Dryopithecus fontani, in the relatively thicker enamel and lowercrowned molars; from Hispanopithecus spp., in the more inflated crown bases, less peripheral cusps and more restricted maxillary sinus; and from Hispanopithecus laietanus also in the thicker crests, more restricted occlusal foveae, and relatively lower-crowned molars. The new specimens of A. brevirostris show some slight differences compared with the holotype of this species: smaller size (presumably due to sexual size dimorphism), and less distally-tapering M2 occlusal contour (which is highly variable in both extant and extinct hominoids). The reported remains provide valuable new evidence on dental intraspecific variation and sexual dimorphism in Anoiapithecus. From a taxonomic viewpoint, they support the distinction of this taxon from both Dryopithecus and Pierolapithecus. From a chronostratigraphic perspective, IPS35027 from ACM/C1-E* enlarges the known temporal distribution of Anoiapithecus, further representing the oldest record (first appearance datum) of hominoids in the Iberian Peninsula.
The early Pleistocene deciduous hominid molar FS-72 from the Sangiran dome of java, Indonesia: A taxonomic reappraisal based on its comparative endostructural characterization
American journal of physical anthropology, Jan 6, 2015
Among the ten fossil hominid deciduous teeth reported so far from the Pleistocene sediments of th... more Among the ten fossil hominid deciduous teeth reported so far from the Pleistocene sediments of the Sangiran Dome of Java are two isolated lower second molars: specimens PCG.2 from the Kabuh Formation and FS-72 from the Pucangan Formation. While PCG.2 appears to be certainly attributable to Homo erectus, FS-72 is somewhat more problematic, even though it is commonly listed within the Indonesian H. erectus hypodigm. Largely because of its large size, it was originally attributed to Meganthropus paleojavanicus. Subsequent study highlighted a set of metric and nonmetric crown features also found in Australopith and African early Homo (notably H. habilis) homologues. An additional problem with the taxonomic assignment of isolated teeth from the Pleistocene of Java is the presence of Pongo in these same deposits. To assess the taxonomic affinity of FS-72, we investigated its inner structure (tissue proportions and enamel-dentine junction morphology) by using techniques of 2-3D virtual ima...
Journal of Human Evolution, 2014
Fieldwork performed during the last 15 years in various Early Pleistocene East African sites has ... more Fieldwork performed during the last 15 years in various Early Pleistocene East African sites has significantly enlarged the fossil record of Homo erectus sensu lato (s.l.). Additional evidence comes from the Danakil Depression of Eritrea, where over 200 late Early to early Middle Pleistocene sites have been identified within a w1000 m-thick sedimentary succession outcropping in the Dandiero Rift Basin, near Buia. Along with an adult cranium (UA 31), which displays a blend of H. erectus-like and derived morphoarchitectural features and three pelvic remains, two isolated permanent incisors (UA 222 and UA 369) have also been recovered from the 1 Ma (millions of years ago) Homo-bearing outcrop of Uadi Aalad. Since 2010, our surveys have expanded to the nearby (4.7 km) site of Mulhuli-Amo (MA). This is a fossiliferous area that has been preliminarily surveyed because of its exceptional concentration of Acheulean stone tools. So far, the site has yielded 10 human remains, including the unworn crown of a lower permanent molar (MA 93). Using diverse analytical tools (including high resolution mCT and mMRI), we analysed the external and internal macromorphology and microstructure of the three specimens, and whenever possible compared the results with similar evidence from early Homo, H. erectus s.l., H. antecessor, H. heidelbergensis (from North Africa), Neanderthals and modern humans. We also assessed the UA 369 lower incisor from Uadi Aalad for root completion timing and showed that it compares well with data for root apex closure in modern human populations.
This contribution investigates the inner organizational pattern (tooth tissue proportions and ena... more This contribution investigates the inner organizational pattern (tooth tissue proportions and enamel–dentine junction morphology) of seven Homo erectus permanent molar crowns from the late Lower-early Middle Pleistocene Kabuh Formation of the Sangiran Dome (Central Java, Indonesia). The previous study of their external characteristics confirmed the degree of time-related structural reduction occurred in Javanese H. erectus, and also revealed a combination of nonmetric features which are rare in the Lower and early Middle Pleistocene dental record, but more frequently found in recent humans. In accordance with their outer occlusal morphology, the specimens exhibit a set of derived internal features, such as thick to hyperthick enamel, an incomplete expression of the crest patterns at the enamel–dentine junction (EDJ) level, a sharp EDJ topography. As a whole, these features differ from those expressed in some penecontemporaneous specimens/samples representing African H. erectus/ ergaster and H. heidelbergensis, as well as in Neanderthals, but occur in recent human populations. Further research in virtual dental paleoanthropology to be developed at macroregional scale would clarify the polarity and intensity of the intermittent exchanges between continental and insular Southeast Asia around the Lower to Middle Pleistocene boundary, as well as should shed light on the still poorly understood longitudinal evolutionary dynamics across continental Asia.
Fieldwork performed during the last 15 years in various Early Pleistocene East African sites has ... more Fieldwork performed during the last 15 years in various Early Pleistocene East African sites has significantly enlarged the fossil record of Homo erectus sensu lato (s.l.). Additional evidence comes from the Danakil Depression of Eritrea, where over 200 late Early to early Middle Pleistocene sites have been identified within a w1000 m-thick sedimentary succession outcropping in the Dandiero Rift Basin, near Buia. Along with an adult cranium (UA 31), which displays a blend of H. erectus-like and derived morphoarchitectural features and three pelvic remains, two isolated permanent incisors (UA 222 and UA 369) have also been recovered from the 1 Ma (millions of years ago) Homo-bearing outcrop of Uadi Aalad. Since 2010, our surveys have expanded to the nearby (4.7 km) site of Mulhuli-Amo (MA). This is a fossiliferous area that has been preliminarily surveyed because of its exceptional concentration of Acheulean stone tools. So far, the site has yielded 10 human remains, including the unworn crown of a lower permanent molar (MA 93). Using diverse analytical tools (including high resolution mCT and mMRI), we analysed the external and internal macromorphology and microstructure of the three specimens, and whenever possible compared the results with similar evidence from early Homo, H. erectus s.l., H. antecessor, H. heidelbergensis (from North Africa), Neanderthals and modern humans. We also assessed the UA 369 lower incisor from Uadi Aalad for root completion timing and showed that it compares well with data for root apex closure in modern human populations.
Journal of human evolution, 2013
New dental remains of the fossil great ape Anoiapithecus brevirostris are described from the Midd... more New dental remains of the fossil great ape Anoiapithecus brevirostris are described from the Middle Miocene local stratigraphic series of Abocador de Can Mata (ACM) in els Hostalets de Pierola (Vallès-Penedès Basin, NE Iberian Peninsula). These specimens correspond to maxillary fragments with upper teeth from two female individuals from two different localities: left P 3 eM 1 (IPS41712) from ACM/C3-Aj (type locality; 11.9 Ma [millions of years ago]); and right M 1 eM 2 and left P 4 eM 2 (IPS35027) from ACM/C1-E* (12.3 e12.2 Ma). Relative enamel thickness is also computed in the latter individual and re-evaluated in other Middle Miocene hominoids from ACM, in order to better assess their taxonomic affinities. With regard to maxillary sinus development, occlusal morphology, molar proportions and enamel thickness, the new specimens show greater resemblances with the (male) holotype specimen of A. brevirostris. They differ from Pierolapithecus catalaunicus in displaying less inflated crests, a more lingually-located hypocone, and relatively lower-crowned molars; from Dryopithecus fontani, in the relatively thicker enamel and lowercrowned molars; from Hispanopithecus spp., in the more inflated crown bases, less peripheral cusps and more restricted maxillary sinus; and from Hispanopithecus laietanus also in the thicker crests, more restricted occlusal foveae, and relatively lower-crowned molars. The new specimens of A. brevirostris show some slight differences compared with the holotype of this species: smaller size (presumably due to sexual size dimorphism), and less distally-tapering M 2 occlusal contour (which is highly variable in both extant and extinct hominoids). The reported remains provide valuable new evidence on dental intraspecific variation and sexual dimorphism in Anoiapithecus. From a taxonomic viewpoint, they support the distinction of this taxon from both Dryopithecus and Pierolapithecus. From a chronostratigraphic perspective, IPS35027 from ACM/C1-E* enlarges the known temporal distribution of Anoiapithecus, further representing the oldest record (first appearance datum) of hominoids in the Iberian Peninsula.
Journal of human evolution, 2013
The Buia Homo site, also known as Wadi Aalad, is an East African paleoanthropological site near t... more The Buia Homo site, also known as Wadi Aalad, is an East African paleoanthropological site near the village of Buia that, due to its very rich yield from the late Early Pleistocene, has been intensively investigated since 1994. In this paper, which reports on the finds of the 2010e2011 excavations, we include new fossil evidence on previously identified taxa (i.e., reptiles), as well as the very first description of the small mammal, fish and bird remains discovered. In particular, this study documents the discovery of the first African fossil of the genus Burhinus (Aves, Charadriiformes) and of the first rodent from the site. This latter is identified as a thryonomyid rodent (cane rat), a relatively common taxon in African paleoanthropological faunal assemblages. On the whole, the new occurrences documented within the Buia vertebrate assemblage confirm the occurrence of taxa characterized by strong water dependence. The paleoenvironmental characteristics of the fauna are confirmed as fully compatible with the evidence obtained through sedimentology and facies analysis, documenting the sedimentary evolution of fluvio-deltaic and lacustrine systems.
Nuclear Instruments and Methods in Physics Research Section A, 2013
A set of portable/transportable X-ray analytical instruments based on radiography, microtomograph... more A set of portable/transportable X-ray analytical instruments based on radiography, microtomography, fluorescence and diffraction have been built and are being operated at the Multidisciplinary Laboratory (MLAB) of the 'Abdus Salam' International Centre for Theoretical Physics (ICTP) in collaboration with Elettra-Sincrotrone Trieste. This is part of a project funded by the Region Friuli Venezia Giulia (Italy) and the ICTP, which aims to develop innovative X-ray analytical tools for noninvasive studies of cultural heritage objects and palaeontological remains. The X-ray instruments at MLAB are also used for hands-on training activities involving students and scientists from developing countries. The MLAB analytical tools complement the microtomography instruments available at Elettra-Sincrotrone Trieste. Examples of our first studies in archaeological and palaeontological applications are presented here.
PloS one, 2013
This contribution reports fifteen human fossil dental remains found during the last two decades i... more This contribution reports fifteen human fossil dental remains found during the last two decades in the Sangiran Dome area, in Central Java, Indonesia. Among this sample, only one of the specimens had already been briefly described, with the other fourteen remaining unreported. Seven of the fifteen isolated teeth were found in a secured stratigraphic context in the late Lower-early Middle Pleistocene Kabuh Formation. The remaining elements were surface finds which, based on coincidental sources of information, were inferred as coming from the Kabuh Formation. Mainly constituted of permanent molars, but also including one upper incisor and one upper premolar, this dental sample brings additional evidence for a marked degree of size variation and time-related structural reduction in Javanese H. erectus. This is notably expressed by a significant decrease of the mesiodistal diameter, frequently associated to the reduction or even loss of the lower molar distal cusp (hypoconulid) and to a more square occlusal outline. In addition to the hypoconulid reduction or loss, this new sample also exhibits a low frequency of the occlusal Y-groove pattern, with a dominance of the X and, to a lesser extent, of the+patterns. This combination is rare in the Lower and early Middle Pleistocene paleoanthropological record, including in the early Javanese dental assemblage from the Sangiran Dome. On the other hand, similar dental features are found in Chinese H. erectus and in H. heidelbergensis. As a whole, this new record confirms the complex nature of the intermittent exchanges that occurred between continental and insular Southeast Asia through the Pleistocene.
Comptes Rendus Palevol, 2013
American journal of physical anthropology, 2012
Currently, the human deciduous dental record from the Pleistocene deposits of the Sangiran Dome, ... more Currently, the human deciduous dental record from the Pleistocene deposits of the Sangiran Dome, Java, consists of only eight specimens. Here we report two deciduous crowns collected near the village of Pucung. While their precise geo-chronological context remains unknown, a provenance from the Early–Middle Pleistocene Kabuh Formation, or from the Early Pleistocene “Grenzbank Zone,” is very likely. These isolated specimens consist of an upper first molar (PCG.1) and a lower second molar (PCG.2). Taxonomic discrimination of the Indonesian tooth record is difficult because of the convergence in crown size and appearance between Pongo and Homo. Accordingly, as PCG.2 still bears a concretion masking most of its features, we coupled the outer analysis of the two specimens with an investigation of their inner morphology. In addition to external characteristics, virtual imaging and quantitative assessment of inner morphology and tissue proportions support an attribution to the taxon Homo, and we preliminary allocate both specimens toH. erectus. Am J Phys Anthropol 2012. © 2012 Wiley Periodicals, Inc.
American Journal of Physical Anthropology, 2010
Qualitative and quantitative characterization through functional imaging of mineralized tissues i... more Qualitative and quantitative characterization through functional imaging of mineralized tissues is of potential value in the study of the odontoskeletal remains. This technique, widely developed in the medical field, allows the bi-dimensional, planar representation of some local morphometric properties, i.e., topographic thickness variation, of a three-dimensional object, such as a long bone shaft. Nonetheless, the use of morphometric maps is still limited in (paleo)anthropology, and their feasibility has not been adequately tested on fossil specimens. Using high-resolution microtomographic images, here we apply bi-dimensional virtual ''unrolling'' and synthetic thickness mapping techniques to compare cortical bone topographic variation across the shaft in a modern and a fossil human adult femur (the Magdalenian from Chancelade). We also test,
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Papers by Clément Zanolli
maxillary sinus development, occlusal morphology, molar proportions and enamel thickness, the new specimens show greater resemblances with the (male) holotype specimen of A. brevirostris. They differ from Pierolapithecus catalaunicus in displaying less inflated crests, a more lingually-located hypocone, and relatively lower-crowned molars; from Dryopithecus fontani, in the relatively thicker enamel and lowercrowned molars; from Hispanopithecus spp., in the more inflated crown bases, less peripheral cusps and more restricted maxillary sinus; and from Hispanopithecus laietanus also in the thicker crests, more restricted occlusal foveae, and relatively lower-crowned molars. The new specimens of A. brevirostris show some slight differences compared with the holotype of this species: smaller size (presumably due to sexual size dimorphism), and less distally-tapering M2 occlusal contour (which is highly variable in both extant and extinct hominoids). The reported remains provide valuable new evidence on dental intraspecific variation and sexual dimorphism in Anoiapithecus. From a taxonomic viewpoint, they support the distinction of this taxon from both Dryopithecus and Pierolapithecus. From a chronostratigraphic perspective, IPS35027 from ACM/C1-E* enlarges the known temporal distribution of Anoiapithecus, further representing the oldest record (first appearance datum) of hominoids in the Iberian Peninsula.