New data on the Middle Pleistocene small mammal fauna from the Homo bearing site of Fontana Ranuccio (Anagni Basin, Central Italy)

The scanty small mammals material of the Middle Pleistocene Fontana Ranuccio site is characterized by the presence of six spe-cies belonging to three order: Eulipotyphla, Lagomorpha, and Rodentia. The small mammal fossils evidence from Fontana Ranuccio, instead the scarce number of specimens, indicates that the site is characterized by a warm and wooded environment with subordinate grassland

The role of climate change in the extinction of the last wild equids of Europe: Palaeoecology of Equus ferus and Equus hydruntinus during the Last Glacial Period

This paper presents as main contribution the combination of membrane osmotic distillation (OD) to dealcoholize red wine with hydrophobic-hydrophilic pervaporation (PV) carried out to add value to the wastewater (extracting water) produced in OD, recycling water and generating bioethanol. Membrane OD with a commercial polypropylene hollow fiber module was applied to partially dealcoholize red wine from 14.0 to 11.0 v/v% ethanol. The OD extracting water, containing only ca. 5.3 wt% ethanol, was treated by sequential PV with both hydrophobic (PDMS or zeolite silicalite-1) and hydrophilic (zeolites mordenite or faujasite) membranes. This hydrophobic-hydrophilic PV produced two main products: bioethanol (recovering 88% of the ethanol removed from the wine) and a 99.4 wt% water-rich product. This water-rich product, with a very low ethanol content, was used as extracting water in the OD, giving rise to an analogous partially dealcoholized wine, in terms of aroma contents (as determined by gas chromatography for 25 compounds), to that achieved when using fresh water

Testing Equid Body Mass Estimate Equations on Modern Zebras-With Implications to Understanding the Relationship of Body Size, Diet, and Habitats of Equus in the Pleistocene of Europe

The monodactyl horses of the genus Equus originated in North America during the Pliocene, and from the beginning of the Pleistocene, they have been an essential part of the large ungulate communities of Europe, North America and Africa. Understanding how body size of Equus species evolved and varied in relation to changes in environments and diet thus forms an important part of understanding the dynamics of ungulate body size variation in relation to Pleistocene paleoenvironmental changes. Here we test previously published body mass estimation equations for the family Equidae by investigating how accurately different skeletal and dental measurements estimate the mean body mass (and body mass range) reported for extant Grevy's zebra (Equus grevyi) and Burchell's zebra (Equus quagga). Based on these tests and information on how frequently skeletal elements occur in the fossil record, we construct a hierarchy of best practices for the selection of body mass estimation equations in Equus. As a case study, we explore body size variation in Pleistocene European Equus paleopopulations in relation to diet and vegetation structure in their paleoenvironments. We show a relationship between diet and body size in Equus: very large-sized species tend to have more browse-dominated diets than small and medium-sized species, and paleovegetation proxies indicate on average more open and grass-rich paleoenvironments for small-sized, grazing species of Equus. When more than one species of Equus co-occur sympatrically, the larger species tend to be less abundant and have more browse-dominated diets than the smaller species. We suggest that body size variation in Pleistocene Equus was driven by a combined effect of resource quality and availability, partitioning of habitats and resources between species, and the effect of environmental openness and group size on the body size of individuals.Peer reviewe

Dietary adaptations of early and middle Pleistocene equids from the Anagni basin (Frosinone, Central Italy)

The intermontane Anagni Basin (Frosinone, central Italy) is an important region for Italian biochronology and paleoecology due to the presence of two rich fossil assemblages dated to the Early (Coste San Giacomo) and Middle Pleistocene (Fontana Ranuccio). Thesesiteshaveyieldedavastcollectionoflargefossilmammalswithawell-documented presence of fossil equids in both localities (represented mostly by isolated teeth). Coste San Giacomo is dated to around 2.1Ma, thereby having recorded the effects of the onset of the Quaternary glacial cycles, which led to a gradual trend toward colder and more arid conditions in the Northern Hemisphere. The fossil equids of this site belong to the first group of grazing stenonid equids of the genus Equus that spread to the Italian Peninsula during the so called "Elephant- Equus" event, which marked the appearance of new large mammals living in herds in open and arid environments. The site of Fontana Ranuccio is dated to around 400 ka, close to the MIS 12-11 succession (the "Mid-Brunhes event"), which marked the end of the Middle Pleistocene Transition. The fossil horses from Fontana Ranuccio represent one of the oldest caballoid (or "true horses") populations of the Italian Peninsula. The Anagni Basin, thus, provides important data to investigate paleoecological adaptations of these groups of equids in response to two critical environmental and climatic shifts of the Pleistocene. We explore their niche occupation by examining long-term dental wear patterns and tooth enamel carbon and oxygen stable isotopic composition. Both taxa appear to have exhibited a narrow dietary niche, displaying a clear abrasive (highly specialized) grass-rich diet. In particular, caballoid equids from Fontana Ranuccio show a more abrasion-dominated mesowear signature. Stenonid equids from Coste San Giacomo exploited broader and more diverse landscapes during the Early Pleistocene, whereas caballoid horses from Fontana Ranuccio appeared to have limited their dietary adaptations to a stricter grazing behavior in more closed environments

Advances in virtual archaeology. Research, preservation, and dissemination

The archaeological and palaeontological record (including human skeletal remains) often bears crack, damage and deformations. The recent rapid development of the diagnostic potentials of “virtual archaeology” has provided innovative tools to manage, study and preserve cultural and natural heritage. These tools include, among others, CT-scans, Laser-scanning, photogrammetry, 3D imaging and rapid prototyping. This approach can contribute to any archaeological context from its discovery to research, preservation, and dissemination. 3D imaging techniques, for instance, substitute physical intervention with a virtual protocol aimed at restoring the original shape of an archaeological item or a fossil specimen. In a similar way, the recovery of digital morphological information can be gathered using data preserved even on a deficient finding through the use of 3D comparative samples. Here we present an extended and updated review about the most innovative protocols applied in virtual archaeology and palaeontology

A REAPPRAISAL OF THE PLEISTOCENE MAMMALS FROM THE KARST INFILLING DEPOSITS OF THE MAGLIE AREA (LECCE, APULIA, SOUTHERN ITALY)

The karst fissures known as “ventarole”, located in the Salentine Peninsula (southernmost part of Apulia, Italy), were first studied by Mirigliano in 1941. These fissures are generally filled with reddish sediments or “terre rosse” in the lower part, and with brownish sediments or “terre brune” in the upper one. Both deposits are particularly rich in vertebrate remains. The mammal assemblages collected within the “terre rosse” from Melpignano and San Sidero are chronologically referred to the early Late Pleistocene (MIS 5), whereas those from the “terre brune” are referred to the Late Pleistocene – Early Holocene (MIS 2/1) transition. These ages were estimated from correlation with the similarly reddish and brownish sediments cropping out in Grotta Romanelli and with other Apulian local faunal assemblages. However, no detailed textural or mineralogical characterization has been carried out on the “ventarole” deposits. Moreover, the presence of several species led to a hypothes of persistence of Middle Pleistocene taxa during the Late Pleistocene in Apulia. In addition, the fauna of Melpignano was proposed as a Faunal Unit of the late Aurelian Land Mammal Age. In the last two decades, a team from the Department of Earth Sciences of Sapienza University of Rome have carried out fieldwork in this fossiliferous area. The rich fossil sample recovered is here presented, coupled with a reassessment of the remains collected since the 1900s. The updated faunal lists of Melpignano and San Sidero include several new species here identified for the first time, in particular Dama clactoniana, Equus mosbachensis and Lynx pardinus. New biochronological and paleoenvironmental data for southern Italy are presented, deriving from the analyses of the mammal remains from Melpignano and San Sidero and their comparison with those from other Middle-Late Pleistocene Apulian sites. Finally, textural and mineralogical analyses performed on several “ventarole” samples allow for a better description of the characteristics of the “terre rosse” and “terre brune”

Evolution of the Family Equidae, Subfamily Equinae, in North, Central and South America, Eurasia and Africa during the Plio-Pleistocene

Studies of horse evolution arose during the middle of the 19th century, and several hypotheses have been proposed for their taxonomy, paleobiogeography, paleoecology and evolution. The present contribution represents a collaboration of 19 multinational experts with the goal of providing an updated summary of Pliocene and Pleistocene North, Central and South American, Eurasian and African horses. At the present time, we recognize 114 valid species across these continents, plus 4 North African species in need of further investigation. Our biochronology and biogeography sections integrate Equinae taxonomic records with their chronologic and geographic ranges recognizing regional biochronologic frameworks. The paleoecology section provides insights into paleobotany and diet utilizing both the mesowear and light microscopic methods, along with calculation of body masses. We provide a temporal sequence of maps that render paleoclimatic conditions across these continents integrated with Equinae occurrences. These records reveal a succession of extinctions of primitive lineages and the rise and diversification of more modern taxa. Two recent morphological-based cladistic analyses are presented here as competing hypotheses, with reference to molecular-based phylogenies. Our contribution represents a state-of-the art understanding of Plio-Pleistocene Equus evolution, their biochronologic and biogeographic background and paleoecological and paleoclimatic contexts

Evolution of the Family Equidae, Subfamily Equinae, in North, Central and South America, Eurasia and Africa during the Plio-Pleistocene

Studies of horse evolution arose during the middle of the 19th century, and several hypotheses have been proposed for their taxonomy, paleobiogeography, paleoecology and evolution. The present contribution represents a collaboration of 19 multinational experts with the goal of providing an updated summary of Pliocene and Pleistocene North, Central and South American, Eurasian and African horses. At the present time, we recognize 114 valid species across these continents, plus 4 North African species in need of further investigation. Our biochronology and biogeography sections integrate Equinae taxonomic records with their chronologic and geographic ranges recognizing regional biochronologic frameworks. The paleoecology section provides insights into paleobotany and diet utilizing both the mesowear and light microscopic methods, along with calculation of body masses. We provide a temporal sequence of maps that render paleoclimatic conditions across these continents integrated with Equinae occurrences. These records reveal a succession of extinctions of primitive lineages and the rise and diversification of more modern taxa. Two recent morphological-based cladistic analyses are presented here as competing hypotheses, with reference to molecular-based phylogenies. Our contribution represents a state-of-the art understanding of Plio-Pleistocene Equus evolution, their biochronologic and biogeographic background and paleoecological and paleoclimatic contexts

Evolution of the Family Equidae, Subfamily Equinae, in North, Central and South America, Eurasia and Africa during the Plio-Pleistocene

Studies of horse evolution arose during the middle of the 19th century, and several hypotheses have been proposed for their taxonomy, paleobiogeography, paleoecology and evolution. The present contribution represents a collaboration of 19 multinational experts with the goal of providing an updated summary of Pliocene and Pleistocene North, Central and South American, Eurasian and African horses. At the present time, we recognize 114 valid species across these continents, plus 4 North African species in need of further investigation. Our biochronology and biogeography sections integrate Equinae taxonomic records with their chronologic and geographic ranges recognizing regional biochronologic frameworks. The paleoecology section provides insights into paleobotany and diet utilizing both the mesowear and light microscopic methods, along with calculation of body masses. We provide a temporal sequence of maps that render paleoclimatic conditions across these continents integrated with Equinae occurrences. These records reveal a succession of extinctions of primitive lineages and the rise and diversification of more modern taxa. Two recent morphological-based cladistic analyses are presented here as competing hypotheses, with reference to molecular-based phylogenies. Our contribution represents a state-of-the art understanding of Plio-Pleistocene Equus evolution, their biochronologic and biogeographic background and paleoecological and paleoclimatic contexts. © 2022 by the authors.Open access journalThis item from the UA Faculty Publications collection is made available by the University of Arizona with support from the University of Arizona Libraries. If you have questions, please contact us at [email protected]

Palaeoecological and palaeoenvironmental reconstruction of the Mediterranean Europe during the Early and Middle Pleistocene based on fossil ungulates dietary adaptations

Il Pleistocene Inferiore e il Pleistocene Medio sono stati caratterizzati da due grandi eventi climatici: l’inizio dei cicli glaciali del Quaternario che hanno instaurato un graduale trend verso climi più freddi e aridi, e la transizione del Pleistocene Medio (Early-Middle Pleistocene Transition) con il passaggio da cicli glaciali regolati da una periodicità di 41 kyr a cicli con periodicità di 100 kyr. Entrambi gli eventi hanno avuto grandi conseguenze sugli ecosistemi terrestri dell’Emisfero boreale, portando in Europa meridionale alla progressiva scomparsa degli ambienti tipicamente subtropicali del Neogene superiore, sostituiti da habitat sempre più aridi e aperti. La risposta degli organismi terrestri a questi cambiamenti climatici è importante per comprendere come variazioni del clima presenti e future, possono influenzare gli habitat della regione del Mediterraneo. Dall'inizio del cenozoico, i mammiferi si sono evoluti e diversificati per occupare una grande varietà di nicchie ecologiche in ambienti che vanno dalla tundra ai deserti, dalle steppe alle foreste tropicali. Tra i grandi mammiferi moderni, gli ungulati erbivori sono particolarmente suscettibili ai cambi di vegetazione in quanto la loro dieta è intrinsecamente legata alla disponibilità di risorse vegetali. Le abitudini alimentari degli ungulati fossili rappresentano quindi un'importante fonte di informazioni riguardo la struttura degli habitat del passato e la loro evoluzione a seguito di variazioni climatiche. In questa tesi, le condizioni paleoambientali di quattro siti del Pleistocene Inferiore e Medio della penisola italiana e della penisola iberica, vengono analizzate e ricostruite mediante l'altezza della corona dei molari (ipsodontia) e i pattern di usura dentaria di artiodattili e perissodattili fossili. Dall'esame delle diete a adattamenti alimentari di questi gruppi di ungulati, è stato possibile acquisire anche nuove conoscenze sulla ripartizione delle nicchie ecologiche e sui meccanismi di sfruttamento delle risorse adottati da questi mammiferi durante il Pleistocene. È stato inoltre sviluppato e testato un nuovo strumento per lo studio delle tracce di microusura presenti sullo smalto dei denti. Dopo il passaggio dal Pliocene al Pleistocene, la regione centrale della penisola italiana è stata caratterizzata da ambienti molto eterogenei con la presenza sia da zone umide/paludose, foreste e praterie aperte. In questo contesto le numerose specie di ungulati simpatrici adottavano un ampio range di abitudini alimentari come riscontrato nelle faune fossili del sito di Coste San Giacomo (2.1 Ma; Anagni, penisola italiana centrale). In questa località taxa di taglia più piccola mostrano morfologie e pattern di usura dentaria tipica di brucatori con una dieta più selettiva, mentre animali di dimensioni maggiori risultano avere uno spettro di adattamenti più ampio con taxa brucatori, pascolatori o a dieta mista. Con il progressivo sviluppo del trend verso climi più freddi e più aridi, si osserva una propagazione degli ambienti aperti nella penisola. A seguito della transizione tra Gelasiano e Calabriano gli ungulati fossili mostrano un range più ristretto di abitudini alimentari come attestato nella località del Pleistocene Inferiore (~ 1.8 Ma) di Olivola (Aulla, penisola italiana centrale). In questo sito la maggior parte dei taxa mostra gradi di usura dentaria riconducibili a una dieta mista con l'assenza di ungulati strettamente brucatori. I cervidi in particolare che durante il Gelasiano consumavano prevalentemente risorse vegetali tenere e poco abrasive in ambienti boschivi, durante il Calabriano modificano le loro abitudini alimentari includendo il consumo di piante più abrasive per trarre vantaggio dalla diffusione di ambienti più aperti. Le praterie aperte erano ancora l'elemento dominante degli ambienti dell'Europa meridionale durante il tardo Pleistocene Inferiore. A seguito del MIS 22, e del così detto "0.9 Ma event", si registra un incremento della stagionalità nella regione del Mediterraneo come attestato dall'alto numero di ungulati a dieta mista stagionale presenti nel livello EVT7 della sezione di Vallparadís (bacino di Vallès-Penedès, penisola iberica NE). Questa intensificazione della stagionalità potrebbe aver portato a considerevoli cambiamenti periodici della qualità di risorse vegetali disponibili. In questo scenario è possibile che i mammiferi erbivori tendessero ad ampliare il proprio dietary breadth includendo più tipologie di piante (più e meno abrasive) nella propria dieta. Una marcata stagionalità ha rappresentato un elemento chiave anche per gli ecosistemi della penisola italiana dopo la fase terminale della transizione del Pleistocene Medio. Questi ambienti che in Italia Centrale (Fontana Ranuccio, bacino di Anagni) hanno agito come habitat-rifugio per popolazioni di ominidi fossili, erano caratterizzati da ambienti eterogenei che spaziavano da foreste a spazi più aperti come comprovato dalla relativa abbondanza di cervidi brucatori e dalla presenza di taxa con una dieta molto o fortemente ricca di piante erbacee. La discrepanza osservata tra i risultati ottenuti dei pattern di usura dentari a lungo e breve termine nelle popolazioni di uro fossile (Bos primigenius) suggeriscono che una forte stagionalità con rigidi periodi avversi potrebbe aver spinto questi grandi animali a nutrirsi periodicamente anche di risorse vegetali sub-ottimali rispetto alla loro dieta abituale. Utilizzando alcuni dei dati raccolti durante il progetto di ricerca, è stato inoltre sviluppato un programma open access (MicroWeaR) per migliorare la metodologia corrente di analisi dei pattern di microusura dentaria. Il software, scritto in linguaggio di programmazione R, è un tool liberamente scaricabile che permette di quantificare e classificare in modo semi-automatico le tracce microscopiche di usura. Il programma è stato testato su foto di superfici dentarie appartenenti a due mammiferi erbivori fossili (un cervide e un primate) ottenute con uno stereomicroscopio e un microscopio elettronico a scansione. I risultati ottenuti dall'analisi effettuata con MicroWeaR sono consistenti con quelli acquisiti tramite utilizzo di altri software di elaborazione di immagini. MicroWeaR rappresenta quindi un valido strumento per uno studio rapido e preciso delle abitudini alimentari di taxa fossili.The Early and Middle Pleistocene were characterised by two major climatic events: the onset of the Quaternary glacial cycles which led to a gradual trend towards cooler and more arid conditions and the Early-Middle Pleistocene Transition (EMPT) with cycles changing from a 41 kyr to 100 kyr periodicity. Both events had great consequences on Northern Hemisphere terrestrial ecosystems, and in Southern Europe, led to the progressive disappearance of the late Neogene subtropical environments and to the spread of more arid open habitats. The response of past biota to these climatic changes is important to understand how present and future climate shifts can affect the structure of Mediterranean habitats. Since the beginning of the Cenozoic era, mammals evolved and diversified to occupy a variety of ecological niches in different environments spanning from the cold tundra to arid deserts, from steppes to rainforests. In modern terrestrial large mammals, herbivorous ungulates are particularly susceptible to vegetation change as their feeding behaviours are intrinsically linked to plant resource availability. Dietary adaptations of fossil ungulates thus represent an important source of information about past habitats and their evolution following climatic variations. In this dissertation, palaeoenvironmental conditions of four Early and Middle Pleistocene sites of the Italian and Iberian Peninsula are reconstructed by means of molar crown height (hypsodonty) and dental wear patterns of fossil artiodactyls and perissodactyls. By examining ungulate dietary proclivities, new insights on the niche partitioning and resource exploitation mechanisms of this group during the Pleistocene are also provided. Moreover, a new tool to investigate dental microwear patterns in fossil taxa is here tested and described. After the Plio/Pleistocene transition, heterogeneous environments characterised the Central Italian Peninsula, with the occurrence of wetlands, closed canopies and open grasslands. In this context, numerous ungulates could live in sympatric conditions by adopting a wide range of feeding behaviours as reported from the Early Pleistocene fossil assemblage of Coste San Giacomo (2.1 Ma; Anagni, Central Italian Peninsula). Smaller taxa were generally selective feeders while larger animals had more diverse dietary adaptations spanning from browsers to mixed feeders to grazers. As the trend towards cooler and more arid climates continued, open landscapes spread further in the peninsula. Following the Gelasian/Calabrian transition, fossil ungulates display a narrower range of feeding behaviours as attested in the Early Pleistocene (~ 1.8 Ma) locality of Olivola (Aulla, Central Italian Peninsula). In this site, most taxa display a mixed diet with no occurrence of strict browsing signals. Cervids in particular during the Gelasian consumed predominantly soft plant resources in wooded environments, but during the Calabrian shifted their diet to include more abrasive items taking advantage of the diffusion of more open habitats. Open grasslands were still a dominant element of the Southern European environments during the late Early Pleistocene. After MIS 22, the so-called "0.9 Ma event", an increase in seasonality is recorded in Mediterranean Europe as attested by the high number of seasonal mixed feeders in layer EVT7 from the Vallparadís Section (Vallès-Penedès Basin, NE Iberian Peninsula). This increment of seasonality may have led to considerable periodic changes in the quality of food items available to herbivores which in adverse seasons widened their dietary breadth feeding on both soft and more abrasive plant resources. Seasonality also played a key role in shaping the ecosystems of the Italian Peninsula after the end of the EMPT, which acted as refugium habitats for archaic human populations. In the Middle Pleistocene site of Fontana Ranuccio (Anagni basin, Central Italian Peninsula), the relative abundance of browsing cervids and the presence of taxa with a grass-rich mixed or grazing diet point to heterogeneous environments spanning from forests to more open landscapes. The pronounced discrepancy between long-term and short-term dental wear patterns of the large aurochs Bos primigenius suggests the occurrence of marked seasonality with harsh adverse periods in which this species was forced to feed even on sub-optimal food items. Using the collected dental microwear data, an open access program to improve current 2D microwear analysis has been designed. The software written in R environment is a free tool, MicroWeaR, which allows for a semi-automatic quantification and classification of microwear features. The program has been tested on pictures of two fossil herbivorous mammals (a cervid and a primate) taken with a stereomicroscope and a scanning electron microscope. Results are consistent with those obtained with other image processing software, thus MicroWeaR represents a robust program for precise and cheaper dental microwear analysis, and a valid tool to investigate dietary behaviours of fossil taxa