Digital reconstruction of the Ceprano calvarium (Italy), and implications for its interpretation

The Ceprano calvarium was discovered in fragments on March 1994 near the town of Ceprano in southern Latium (Italy), embedded in Middle Pleistocene layers. After reconstruction, its morphological features suggests that the specimen belongs to an archaic variant of H. heidelbergensis, representing a proxy for the last common ancestor of the diverging clades that respectively led to H. neanderthalensis and H. sapiens. Unfortunately, the calvarium was taphonomically damaged. The postero-lateral vault, in particular, appears deformed and this postmortem damage may have infuenced previous interpretations. Specifcally, there is a depression on the fragmented left parietal, while the right cranial wall is warped and angulated. This deformation afected the shape of the occipital squama, producing an inclination of the transverse occipital torus. In this paper, after X-ray microtomography (μCT) of both the calvarium and several additional fragments, we analyze consistency and pattern of the taphonomic deformation that afected the specimen, before the computer-assisted retrodeformation has been performed; this has also provided the opportunity to reappraise early attempts at restoration. As a result, we ofer a revised interpretation for the Ceprano calvarium’s original shape, now free from the previous uncertainties, along with insight for its complex depositional and taphonomic history

Fontana Ranuccio (excavations 2019-2022) and Colle Marino (excavations 2020-2021): Middle Pleistocene chronostratigraphic evidence of the earliest inhabitants in the Anagni basin

Recent fieldwork of the Istituto Italiano di Paleontologia Umana in the Anagni basin (Latin Valley, southern Latium, central Italy) has provided new insights into the dynamics of human settlement and adaptation in this area during the Middle Pleistocene. This paper summarizes the still partially unpublished data on the chronostratigraphy of two important prehistoric sites: Fontana Ranuccio, dating from about 400,000 years ago, and Colle Marino, whose lithic industry has so far been attributed to an early stage of the Middle Pleistocene. At Fontana Ranuccio, stratigraphic studies highlighted the depositional dynamics and the processes involved in the formation of the archeological unit, consisting of partly reworked volcanic material, which can be now divided into distinct sub-units. At Colle Marino, a 35-meter-deep core drilling and an excavation campaign allowed us to verify the nature and provenance of lithic artifacts found on the surface in past decades. Radiometric analyses dated these artifacts from about 700,000 years ago

Modeling the Past: The Paleoethnological Evidence

This chapter considers the earliest Paleolithic, Oldowan (Mode 1), and Acheulean (Mode 2) cultures of the Old Continent and the traces left by the earliest hominids since their departure from Africa. According to the most recent archaeological data, they seem to have followed two main dispersal routes across the Arabian Peninsula toward the Levant, to the north, and the Indian subcontinent, to the east. According to recent discoveries at Dmanisi in the Caucasus, the first Paleolithic settlement of Europe is dated to some 1.75 Myr ago, which indicates that the first “out of Africa” took place at least slightly before this date. The data available for Western Europe show that the first Paleolithic sites can be attributed to the period slightly before 1.0 Myr ago. The first well-defined “structural remains” so far discovered in Europe are those of Isernia La Pineta in Southern Italy, where a semicircular artificial platform made of stone boulders and animal bones has been excavated. The first hand-thrown hunting weapons come from the site of Scho¨ningen in north Germany, where the first occurrence of wooden spears, more than 2 m long, has been recorded from a site attributed to some 0.37 Myr ago. Slightly later began the regular control of fire. Although most of the archaeological finds of these ages consist of chipped stone artifacts, indications of art seem to be already present in the Acheulean of Africa and the Indian subcontinent

The morpho-functional approach as alternative descriptor for the study of the soft bottom malacological communities

Le comunità bentoniche rivestono un ruolo fondamentale nella caratterizzazione e nella funzionalità degli ecosistemi marini costieri, in quanto per le loro caratteristiche di persistenza, costituiscono delle memorie biologiche capaci di integrare nel tempo eventi diversi. Per l’importante ruolo ecologico che il macrobenthos riveste nell’ambito dell’ecosistema marino, la conoscenza dei pattern di diversità che caratterizzano tali popolamenti diventa indispensabile, al fine di identificare aree prioritarie per la loro conservazione e per la regolamentazione delle attività antropiche che insistono in tali zone. Negli ecosistemi marini costieri la diversità funzionale dei popolamenti bentonici è sempre stata tradizionalmente descritta attraverso l’analisi della composizione tassonomica di tali popolamenti. Un certo pattern nella struttura tassonomica può essere interpretato alla luce di cambiamenti in alcune caratteristiche ecologiche proprie dei diversi taxa, cruciali nella descrizione dei presunti ruoli funzionali svolti da tali organismi. Benché l’approccio tassonomico sia in grado di rivelare la risposta agli stress ambientali, può risultare difficile individuare quali caratteristiche ecologiche effettivamente determinino tali reazioni. Organismi che mostrano tratti ecologici simili, non sempre rispondono agli “stressori” allo stesso modo, poiché, pur condividendo alcuni importanti attributi, possono differirne in altri, meno evidenti, ma altrettanto significativi. Le funzioni svolte dalle specie bentoniche sono importanti nella regolazione dei processi ecosistemici e queste funzioni sono determinate dalle caratteristiche biologiche che queste specie mostrano, quali alimentazione, movimento, costruzione di esoscheletri o formazioni accessorie (p.e., tubi), etc. Questo lavoro propone un approccio per lo studio delle comunità macrozoobentoniche di fondo mobile, che nasce dalla comparazione fra la lettura del sistema da parte dei classici descrittori tassonomico-sistematici (specie) e quella dei descrittori alternativi (caratteristiche morfologiche) per il syntaxon a Molluschi. Questo phylum è il gruppo più significativo ed eterogeneo del macrozoobenthos di fondo mobile ed è considerato descrittore efficace delle comunità bentoniche; ciò vuol dire che sono eccellenti indicatori ambientali ed è spesso sufficiente l’analisi della malacofauna marina per ottenere preziose informazioni sullo stato di salute di un ecosistema. L’elemento caratterizzante il syntaxon a Molluschi è la conchiglia, un esoscheletro calcareo che, prodotto almeno nei primi stadi di vita, nella quasi totalità delle specie segue l’accrescimento del corpo dell’animale. In tal senso, la conchiglia è l’interfaccia col quale l’animale si relaziona col suo ambiente esterno e la cui struttura deve essere tale da permetterne le funzioni primarie tipiche (alimentazione, riproduzione, locomozione, ecc.). L’analisi morfologica della conchiglia e la sua variazione fra i taxa può rappresentare quindi un strumento utile per l’identificazione di variazioni nelle condizioni ambientali e la valutazione dello stato ecologico di un ecosistema. Su tale tipo di approccio morfo-funzionale si basa la ricerca oggetto di questa Tesi, con la quale si vuole definire una procedura alternativa per la valutazione dell’assetto delle comunità macrozoobentoniche di fondo mobile e il riconoscimento di tipologie ricorrenti, mediante l’utilizzo di descrittori alternativi a quello tassonomico. È stato utilizzato un data set comprendente 60 campagne di campionamento del macrozoobenthos di fondo mobile effettuate lungo tutta la costa italiana e a diverse profondità. La comparazione della forma della conchiglia di ciascuna specie e l’utilizzo dei modelli geometrici disponibili in letteratura, è stato possibile individuare diverse categorie morfologiche (morphotaxa) trasversali alle tradizionali categorie tassonomiche (specie) con la finalità di rappresentare aspetti diversi delle funzioni ecologiche all’interno delle comunità e quindi dell’ecosistema. Attraverso la comparazione dei risultati ottenuti con l’approccio sinecologico classico (analisi sulle specie o analisi strutturale) e con l’approccio morfo-funzionale (analisi sui morphotaxa), si è valutata la coerenza tra forma ed habitat. Inoltre, l’analisi batimetrica e sedimentologica ha permesso di riconoscere range preferenziali per ciascun morphotaxon, mentre l’analisi biocenotica sugli elementi costitutivi di ciascun morphotaxon e lo studio dei risultati ottenuti tramite analisi multivariata (Analisi delle Corrispondenze, AC; Indicator Species Analysis, ISA) su ciascuna campagna di campionamento ha permesso il riconoscimento di tipologie ricorrenti lungo il gradiente costa-largo. Il lavoro svolto di raccolta e organizzazione dei dati morfologici è stato finalizzato alla creazione di un data set preliminare di dati relativi ai tratti morfologico-funzionali dei Molluschi marini di fondo mobile reperiti nel data set, per valutare e costruire, quindi, strumenti analitici per la messa a punto di una metodologia. Essa si basa sui descrittori alternativi a quello tassonomicosistematico, impiegabile nell’ambito del monitoraggio biologico e quindi nella valutazione dello stato ecologico dell’ecosistema marino.The macrozoobenthic communities have such particular dynamics so as to permit an integrated valuation of the space – temporal alteration of the ecosystem. Soft bottom macrobenthic animals, are in fact relatively sedentary, have long life-spans and are represented by a great number of species with different tolerance to stress and, at last, they play an important role in recycling nutrients. Taking into consideration this heterogeneity and complexity, ecologists have the necessity to identify the emerging properties in order to simplify the structures and the processes of the system. In this framework the “community structure” is much more efficient than the “species” as indicator of the status of the ecosystem as it represents the historical memory of the system. Identical species listed in different time and space are equivalent to equal state of the system and each shift in the list-vector corresponds to a significant change of the state of the system. Although the taxonomical approach has several point of weakness it might be difficult to compare taxonomical identifications made by different experts; the skills and the knowledge of taxonomists may vary due to many different factors such as experience and practice. Moreover the link between the species and their functional role couldn’t be directly evaluated. The contribution of the benthic assemblages to ecosystem functioning will be dependent on the biological characteristics, or traits, exhibited by constituent species, as these determine how the species influence ecological processes. For example, burrowing activities of benthic invertebrates may alter sediment conditions through the resuspension of fine particles into overlying waters, may change the distribution of different-sized particles within sediments and may produce increases in the oxygen and water contents. These activities also transport buried organic matter and nutrients to the sediment–water interface, stimulating benthic microflora and microfauna communities and hence decomposition processes. The present research provides a comparison between the taxonomical and non taxonomical approach in the study of the soft bottom benthic assemblages using a multitraits approach, stressing the role of the Mollusc syntaxon to the ecosystem functioning. This phylum is, among the macrozoobenthic animals, a group of extreme heterogeneity in terms of feeding behaviour, habitus, reproduction and morphology. It is considered as efficient descriptor of benthic communities, meaning that they are excellent environmental indicators and the analysis of the marine malacofauna is often enough to provide valuable information about the state of health of an ecosystem. The characterizing element of the Mollusc syntaxon is the shell, a calcareous exoskeleton produced always in the first life stages and following almost ever the body growth of the animal. For this reason, the shell is the interface through which the animal acquaints itself with its external environment, and its morphology must allow the typical primary functions of the animal (feeding, reproduction, locomotion, etc.). The morphological analysis of the shell and its variation among taxa may represent a useful tool to identify variations in the environmental conditions and to evaluate the ecological status of an ecosystem. The research of this Thesis is based upon this approach, defining an alternative procedure for the evaluation of the structure of the soft bottom benthic communities and the recognition of recurrent typologies through the utilization of alternative descriptors to the taxonomic one. The data set object of the study is made of 60 sample surveys that came from all over the Italian peninsula and collected at several depths. By comparing the shell shape of each species and by using geometrical models available in literature, it is possible to identify morphological categories (morphotaxa) transversal to the traditional taxonomical ones (species) with the aim to represent different aspects of the ecological functions in benthic communities and ecosystems. The comparison between the traditional synecological approach (species or structural analysis)and the morpho-functional one has allowed the valuation of the coherence between shape and habitat. Furthermore, the bathymetric and sedimentological analysis has allowed to recognize preferential ranges for each morphotaxon, while the biocoenotic analysis of the structural elements of each morphotaxon and the study of obtained result of each sample survey obtained by Multivariate Analysis (Correspondence Analysis - CA; Indicator Species Analysis, ISA) has allowed to recognize recurrent typologies along the depth gradient. The collection and organization of morphological information has been useful to realize a preliminary data set relative to the morpho-functional traits of soft bottom marine Molluscs, and therefore, to valuate and create analytical tools based upon alternative descriptors to the taxonomical ones, useful in the biological monitoring and in the valuation of the ecological status of the marine ecosystems.Dottorato di ricerca in Ecologia e gestione delle risorse biologich

Digital reconstruction of the Ceprano calvarium (Italy), and implications for its interpretation

The Ceprano calvarium was discovered in fragments on March 1994 near the town of Ceprano in southern Latium (Italy), embedded in Middle Pleistocene layers. After reconstruction, its morphological features suggests that the specimen belongs to an archaic variant of H. heidelbergensis, representing a proxy for the last common ancestor of the diverging clades that respectively led to H. neanderthalensis and H. sapiens. Unfortunately, the calvarium was taphonomically damaged. The postero-lateral vault, in particular, appears deformed and this postmortem damage may have influenced previous interpretations. Specifically, there is a depression on the fragmented left parietal, while the right cranial wall is warped and angulated. This deformation affected the shape of the occipital squama, producing an inclination of the transverse occipital torus. In this paper, after X-ray microtomography (μCT) of both the calvarium and several additional fragments, we analyze consistency and pattern of the taphonomic deformation that affected the specimen, before the computer-assisted retrodeformation has been performed; this has also provided the opportunity to reappraise early attempts at restoration. As a result, we offer a revised interpretation for the Ceprano calvarium\u27s original shape, now free from the previous uncertainties, along with insight for its complex depositional and taphonomic history