Bollettino SPI Vol. 64 (1) - OPEN ACCESS!
The palaeontological legacy of Augusto Azzaroli and Danilo Torre
Published in June 2025
Guest Editors:
Saverio Bartolini-Lucenti,
Luca Pandolfi & Lorenzo Rook
Index
- Bartolini-Lucenti S., Pandolfi S. & Rook L. (2025)
Introduction to “The palaeontological legacy of Augusto Azzaroli and Danilo Torre”
p. i
- Kotsakis T. (2025)
Foreward – Memories of a “Roman” on the group of vertebrate palaeontologists of Florence
pp. iii-v
- Rook L. & Ferretti P.M. (2025)
Dedication to Giovanni Ficcarelli (1937-2024) (Rignano sull’Arno, May 2nd 1937-Firenze, April 21st 2024)
pp. vii-xi
- Rook L., Bartolini-Lucenti S. & Pandolfi L. (2025)
The University of Florence Vertebrate Palaeontology legacy
pp. 1-9
doi:10.4435/BSPI.2025.01
- Zoboli D., Zedda M. & Palombo M.R. (2025)
On the rediscovered fossil remains of Macaca majori Azzaroli, 1946 (Primates, Cercopithecidae) from the Capo Figari type locality (Sardinia, Italy) housed in the Muséum national d’Histoire naturelle (Paris, France)
pp. 11-17
doi:10.4435/BSPI.2025.02
In this report, we present and briefly describe a small sample of the endemic Sardinian macaque Macaca majori Azzaroli, 1946 from the Pleistocene type locality of Capo Figari (Golfo Aranci, northeastern Sardinia). The famous Swiss palaeontologist Charles Immanuel Forsyth Major (1843-1923) donated the fossils to the Muséum national d’Histoire naturelle of Paris (France) in 1914. This still unpublished material represents one of the first M. majori samples ever collected and is of great scientific and historical importance. The history of palaeontological research is comprised within such collections, rendering their preservation and that of the inherent information they contain essential for palaeontological studies.
- Faggi A. (2025)
Revising the carnivores of Baccinello V3 faunal assemblage: palaeobiogeographic implications in the Messinian Italian carnivore record
pp. 19-46
doi: 10.4435/BSPI.2025.03
The Late Miocene and, in particular, the Messinian age is a period characterised by profound palaeoclimatic and palaeogeographic changes. However, Late Miocene fossil remains of terrestrial vertebrates are rare in Italy compared to other European countries, thus limiting investigation of such changes on the Peninsula. The present study deals with new fossil material of carnivorans from the Turolian (Messinian, Late Miocene) belonging to the so called Baccinello V3 faunal assemblage from the Baccinello-Cinigiano basin and adds new considerations regarding the dispersal of carnivores on the Italian Peninsula at the end of the Miocene. Revision of the material housed at the Naturhistorisches Museum Basel and the Museo di Geologia e Paleontologia dell’Università di Firenze has allowed the carnivoran fauna of this site to be updated to now include seven taxa belonging to four different subfamilies (Felidae, Hyaenidae, Viverridae, Mustelidae). The identified forms include: Paramachaerodus orientalis, Amphimachairodus cf. giganteus, Pristifelis sp., cf. Hyaenictitherium, Viverra howelli, Plesiogulo crassa, Guloninae indet., Mustelidae indet. Palaeoecological analyses of the Baccinello V3 carnivora guild highlight the presence of a forested habitat with few open grass areas. In addition, revision of the carnivores from other Messinian Italian sites was carried out, paying particular attention to the no longer existing locality of Gravitelli.
- Bartolini-Lucenti S., Madurell-Malapeira J., Garrido G., Rook L. & Arribas A. (2025)
Early dispersal and niche partitioning in Canidae from an Early Pleistocene site of Spain (Fonelas P-1, Guadix-Baza basin, Granada)
pp. 47-67
doi: 10.4435/BSPI.2025.04
The Fonelas P-1 site in southern Spain (~2.0 Ma) offers an exceptional window into the Early Pleistocene canid diversity and ecology in Europe. This study identifies two medium-sized canid morphotypes from the site, corresponding to the species Canis arnensis Del Campana, 1913 and Canis etruscus Forsyth Major, 1877. Morphological and ecomorphological analyses, integrating cranial, dental, and mandibular features, confirm their taxonomic identities and ecological differentiation. Canis arnensis at Fonelas P-1 expands the geographic and temporal range of this species, providing robust evidence of its presence in southern Europe before 2 Ma. The co-occurrence of C. arnensis and C. etruscus at this well-dated site underscores their ecological divergence, with C. arnensis exhibiting a mesocarnivorous dietary niche and C. etruscus showing adaptations closer to hypercarnivory. These findings illuminate the complexity of Early Pleistocene canid guilds, offering critical insights into their dispersal patterns, adaptive strategies, and interactions within taphocenoses.
- Werdelin L. & Fourvel J.-B. (2025)
A review of fossil Ictonychinae (Mustelidae) from the Plio-Pleistocene of Africa
pp. 69-76
doi: 10.4435/BSPI.2025.05
We here describe one new specimen of Prepoecilogale from South Africa whilst reviewing previously published material. African representatives of the subfamily are known from Morocco, Algeria, Kenya, Tanzania, and South Africa. The South African specimens are assigned to Prepoecilogale bolti (Petter, 1987). North and East African specimens are referred to the extant genera Ictonyx, Poecilictis, and Poecilogale. The specimens from Laetoli, previously assigned to Prepoecilogale, are here reassigned to the new species Ictonyx harrisoni and to an unnamed n. gen. and n. sp.
- Loddi C., Madurell-Malapeira J. & Bartolini-Lucenti S. (2025)
Plioviverrops faventinus reloaded: the last survivor of a successful genus
pp. 77-105
doi: 10.4435/BSPI.2025.06
The taxonomic and systematic conundrum of Middle to Late Miocene small-sized hyaenids includes several taxa generally ascribed to the genera Protictitherium and Plioviverrops. Especially the latter was often related to the extant aardwolf, Proteles cristatus, although a recent study suggests the new genus Gansuyaena as being the plausible ancestor of the African Proteles. In general terms, few studies have concentrated specifically on these small hyaenids, and little is known with regard to their systematics and phylogenetic
relationships. Rather, they were ecomorphologically divided in civet-like and mongoose-like hyaenids based on their morphological and metric features. Often overlooked in literature, Plioviverrops faventinus was originally described by Danilo Torre, who was the first to recognise the peculiarity of the most recently established species of the genus Plioviverrops. In the present paper, we revise the sample from the type locality of Cava Monticino (Brisighella, Ravenna, Italy) clarifying the affinities and differences with other species of Plioviverrops as well as of Protictitherium and Gansuyaena. Moreover, we focus on dentognathic features to test the ecological preferences of P. faventinus and other small-sized hyaenids, compared to a sample of extant Herpestidae and Viverridae. Our ecological results suggest the marked hypocarnivorous/ invertebrivorous specialisation of P. faventinus similar to some extant herpestids.
- Martínez-Navarro B., Madurell-Malapeira J., Bartolini-Lucenti S., Lordkipanidze D. & Rook L. (2025)
The giant short-faced hyaena from Dmanisi: taxonomy and palaeobiology
pp. 107-131
doi: 10.4435/BSPI.2025.07
Abundant, complete and well-preserved specimens attributed to several carnivorans were recovered from the Calabrian palaeoanthropological site of Dmanisi, Georgia (ca. 1.8 Ma). Herein, we describe the hyaenid remains unearthed from this locality, comparing them with other similar forms from coeval sites from Eurasia. The main objective of this contribution is to test the possible co-occurrence of two hyaena species in Dmanisi, as has been reported by previous authors. Our analyses show that the whole Dmanisi sample corresponds to a single taxon of large and stout size, the giant short-faced hyaena Pachycrocuta brevirostris (Gervais, 1850). Furthermore, we discuss the taphonomic implications of this species in the accumulation of fossil assemblages during the Early Pleistocene in Eurasia and, based on the dental anatomy, the palaeobiology of this hyaenid, suggesting different feeding behaviours for juvenile and adult individuals in the exploitation of fat and meat.
- Madurell-Malapeira J. (2025)
A critical review of the Pliocene and Pleistocene European Felidae fossil record
pp. 133-163
doi: 10.4435/BSPI.2025.08
The fossil record of European Pliocene and Pleistocene felids reveals a rich diversity of taxa and provides key insights into their evolutionary, ecological, and biogeographic dynamics. This review synthesises current knowledge on felid species across Europe, highlighting the effects of climatic oscillations and human interactions on their survival and extinction. The study integrates fossil, morphological, isotopic, and genetic data to trace evolutionary trajectories, shedding light on adaptation strategies such as dietary specialisation, morphological shifts, and resilience to environmental stressors. Additionally, unresolved questions about lineage origins, intercontinental dispersals, and evolutionary responses to fluctuating climates highlight avenues for future research. By combining palaeontological and archaeological evidence, this review underscores the ecological significance of felids in shaping Pleistocene ecosystems and provides a comprehensive framework for understanding their evolutionary history. It emphasises the need for further studies to clarify taxonomy, improve chronological resolution and explore biogeographic patterns across Europe’s fossil record.
- Ferretti M.P., Bellucci L. & Rustioni M. (2025)
Cranial morphology and variation in the iconic Villafranchian proboscidean Mammuthus meridionalis (Nesti, 1825) from the type locality of Upper Valdarno (Tuscany, Italy)
pp. 165-187
doi: 10.4435/BSPI.2025.09
The aim of this article is to provide a detailed description of the cranial anatomy of Mammuthus meridionalis (Nesti, 1825) based on the type material from Upper Valdarno (UV), Italy. The UV sample includes both male and female individuals and a fragmented skull of a new- born, allowing us to highlight variation in cranial features linked to both sexual dimorphism and ontogeny. The southern mammoth occupies a central role in elucidating the evolutionary pattern of mammoths during the Early Pleistocene (Villafranchian Land Mammal Age), when the first cold adapted mammoth species, Mammuthus trogontherii (Pohlig, 1885), the steppe mammoth, originated. A detailed knowledge of the morphology and variation of the skull of this iconic Villafranchian species is crucial to clarify the complex systematic and evolutionary pattern of mammoth evolution during this time interval. The study supports the conclusion proposed by Augusto Azzaroli more than forty years ago, that the Upper Valdarno mammoth population was not directly ancestral to the steppe mammoth, which dispersed to Western Europe at the beginning of the Middle Pleistocene.
- Ros-Montoya S., Martínez-Navarro B., Bukhsianize M., Martín-Serra A., Espigares M.P. & Lordkipanidze D. (2025)
A morphological and morphometric study of the elephant remains from Dmanisi (Early Pleistocene, Georgia)
pp. 189-205
doi: 10.4435/BSPI.2025.10
We describe the morphological and morphometric characteristics of the complete collection of proboscidean remains from the site of Dmanisi, ascribed to the species Mammuthus meridionalis (Nesti, 1825). The upper and lower third molars have been compared with the more primitive species Mammuthus rumanus (Ștefănescu, 1924) and Mammuthus meridionalis from the most important sites of Eurasia. Bivariate analysis, principal component analysis and morphological comparison with more than fifty sites have been carried out. The results revealed that the Dmanisi remains do correspond to the species Mammuthus meridionalis and are similar to the Upper Valdarno specimens, in Italy, where this species was originally described.
- Cirilli O. & Bernor R.L. (2025)
One step beyond: a revision of Middle Miocene North American Pliohippus mirabilis provides new clues about equid evolutionary history
pp. 207-226
doi: 10.4435/BSPI.2025.11
The evolution of North American equids has always been used as a paradigmatic case of macroevolutionary patterns through time and space. Nevertheless, despite many phylogenetic, palaeoecological and macroevolutionary studies, their taxonomy at the genus and species level remains obscure, especially for the tribe Equini. Herein, we describe the cranial and postcranial elements of Middle Miocene Pliohippus mirabilis, at different ontogenetic stages, and we compare these with a suite of North American Miocene and Pliocene Equini. Our results clarify the morphology of this species, as well as provide new insights on its position in the evolutionary history of the Equinae that ultimately led to the evolution of the genus Equus. Our results support the hypothesis that Pliohippus mirabilis represents the basal species of Pliohippus, as well as the ancestral form for the radiation of the Equini during the Neogene of North America. Moreover, we provide a comparison of the postcranial morphology of its metapodials, discussing previous hypotheses on the origin of monodactyly. Finally, we present the most up-to-date biochronological framework of the Middle Miocene to Pliocene Equini.
- Pandolfi L., Cerdeño E. & Antoine P.-O. (2025)
Cranial variability in Stephanorhinus hemitoechus (Mammalia, Rhinocerotidae): A revision of Azzaroli’s subspecies and the systematics of Middle-Late Pleistocene European rhinoceroses
pp. 227-244
doi: 10.4435/BSPI.2025.12
Western European rhinoceroses have been the subject of several investigations in recent decades, but only a few studies have taken into account the broad morphological variability in extant rhinoceroses. The latter has given rise to doubts concerning the validity and variation of fossil species. Among the Pleistocene fossil rhinoceroses, Stephanorhinus hemitoechus represents one of the most recorded and described species in Western Europe, being documented by well-preserved cranial remains. Moreover, in addition to the nominotypical subspecies, three further subspecies of S. hemitoechus were erected based on differences in cranial features, namely Rhinoceros hemitoechus falconeri Azzaroli, 1962, Rhinoceros hemitoechus aretinus Azzaroli, 1962 and Dicerorhinus hemitoechus intermedius Azzaroli in Anfossi & Cantaluppi, 1987. The validity of these subspecies and the cranial variability of S. hemitoechus are herein revised after consideration of the available morphological traits of the lectotype of the species, comparison with fossil cranial remains assigned to S. hemitoechus and the morphological variability in extant rhinoceroses. In both extant and fossil rhinoceroses, morphological features such as the shape and size of the occipital face and the position of the nasal notch and of the orbit are affected by ontogenetic patterns, revealing marked changes during the life cycle of an individual. Some other features of the cranium instead, such as the development of the nuchal crest, may have been affected by changes in feeding habitus or sexual dimorphism. Accordingly, the occurrences of subspecies of S. hemitoechus are herein rejected, both from a chronostratigraphic and geographic perspective. Finally, we suggest herein that an estimation of the age at death of the studied material and a more comprehensive comparison with the extant species be provided in future studies in order to arrive at a better understanding of fossil rhinoceroses and their taxonomic assignment.
- Croitor R. (2025)
A revision of Cervus ruscinensis Depéret, 1890 (Cervidae, Mammalia) from the Early Pliocene of Roussillon (France)
pp. 245-259
doi: 10.4435/BSPI.2025.13
This paper presents a detailed morphological redescription of a well-preserved skull of Cervus (Capreolus) ruscinensis Depéret, 1890, from the Early Pliocene of Roussillon. The study includes a Principal Component Analysis of craniometric data, along with a re-evaluation of morphological features, that provide compelling arguments for the establishment of the new genus Xenocervulus. The unique ecomorphological cranial and antler characteristics of the Ruscinian deer suggest specific adaptations for a distinctive form of intraspecific head-to-head male combat. This behaviour, resembling that observed in saber-horned antelopes, is unusual among cervids. The small, lowcrowned upper cheek teeth suggest that this deer followed a typical folivorous or browsing diet, consistent with other small cervids.
- Ghinassi M., Fidolini F., Papini M., Rook L. & Sagri M. (2025)
The Upper Valdarno Basin (Northern Apennines, Italy): from “Azzaroli & Lazzeri (1977)” to the present understanding of a pivotal continental succession in the Pliocene-Pleistocene of Europe
pp. 261-287
doi: 10.4435/BSPI.2025.14
This paper reviews studies of the Upper Valdarno Basin, in order to highlight the unique stratigraphic and palaeontological characteristics of the area. Over recent decades, the stratigraphy of the basin has been meticulously refined, establishing it as one of the most well-documented basins in the Northern Apennines. The sedimentary fill of the basin consists of three unconformity-bounded units: Castelnuovo dei Sabbioni (CSB), Montevarchi (VRC), and Torrente Ciuffenna (UFF) Synthems, with the Palazzolo sub-basin comprising the Fosso Salceto (OLC) and Torrente Ciuffenna (UFF) Synthems. Formed during the Late Pliocene, the basin underwent significant geological changes, transitioning from deposition of fluvial gravels to lacustrine conditions around 3.1 Ma, followed by deltaic sand accumulation and subsequent tectonic uplift events. The Early Pleistocene saw the formation of floodplain lakes and swamps, leading to the accumulation of organic-rich sediments. The vertebrate fossil record of the Upper Valdarno Basin is distinguished by exceptional preservation of the material and is of historical significance, with documentation dating back to the Renaissance. The efforts and finds both by fossil hunters and palaeontologists have contributed to the importance of the collection in defining numerous Pliocene and Pleistocene species and genera, as well as in confirming biochronological subdivisions of the continental timescale. The palaeontological studies initiated by Augusto Azzaroli in the post-war period further enriched the documentation of the fossil collection of the basin, underlining its status as a crucial site in the understanding of Pliocene and Pleistocene vertebrate faunas.
- Bertini A., Marino M., Combourieu-Nebout N., Girone A., Maiorano P.,
Niccolini G. & Suc J.-P. (2025)
Unravelling land-sea correlations during the Pliocene-Pleistocene in the Mediterranean area
pp. 289-312
doi: 10.4435/BSPI.2025.15
In this paper, we synthetised pollen, foraminiferal, and calcareous nannofossil data from marine and continental successions in Italy to investigate the complex dynamics of climate-induced shifts in terrestrial and marine ecosystems during the Late Pliocene and Early to Middle Pleistocene. Additionally, by focusing on three key case studies derived from interdisciplinary research of marine sedimentary successions (Semaforo, Vrica-Santa Lucia and Montalbano Jonico) we present concurrent evidence from both marine and continental
proxies, offering a refined perspective on the environmental history of the Mediterranean. This integrated approach enhances the precision of reconstructing climate system interactions during critical intervals of the Quaternary, mitigating potential discrepancies arising from independent chronologies and methods of study. Overall, the synthesis and case studies contribute to a more comprehensive understanding the role of the Mediterranean in global climate variability, emphasising the sensitivity of the region to both continental and marine climatic drivers throughout the Quaternary.
- Savorelli A., Mazza P.P.A., Borrani A., Delfino M., Pavia M. & Masini F. (2025)
The palaeontological legacy of the Late Miocene Gargano’s Terre Rosse (southeastern Italy): evolutionary patterns, adaptive radiations, and unresolved origins
pp. 313-335
doi: 10.4435/BSPI.2025.16
The Gargano region in the Apulian domain offers an exceptional fossil record that is rich in diversity and continuity, making it a key site for studying evolutionary processes on Mediterranean islands and palaeo-islands. This study provides an updated synthesis of the palaeontological data for the Gargano fauna, integrating the well-known mammalian records with those of the avifauna and herpetofauna. The fossil evidence highlights several modes of evolutionary change, including size variation in small mammals consistent with the “Island Rule”, adaptive radiations, and the emergence of new taxa through dispersal. Notably, the fauna of the Gargano islands reveals adaptive strategies that mirror the evolutionary processes of continental taxa, with the evolution of new morphological and ecological adaptations such as fossoriality in Mikrotia and malacophagy in Apulogalerix. Recent discoveries of the oldest fissure deposits in the region have significantly enhanced our understanding of the biochronological framework and shed new light on the origins of the Gargano fauna, contributing to ongoing debates about the genesis of these Miocene insular communities. Despite the remarkable richness of the fossil record, many aspects of the evolutionary history of this unique fauna remain unresolved, underscoring the fragmentary nature of the evidence and the challenges of reconstructing phylogenetic relationships based on incomplete records. This study aims to offer a comprehensive overview of the evolutionary history of the Gargano insular fauna, addressing complex and unresolved issues in the hopes of stimulating future research on this intricate palaeontological puzzle.proxies, offering a refined perspective on the environmental history of the Mediterranean. This integrated approach enhances the precision of reconstructing climate system interactions during critical intervals of the Quaternary, mitigating potential discrepancies arising from independent chronologies and methods of study. Overall, the synthesis and case studies contribute to a more comprehensive understanding the role of the Mediterranean in global climate variability, emphasising the sensitivity of the region to both continental and marine climatic drivers throughout the Quaternary.
- Lyras G.A., van der Geer B.G.M., Kafousias M.S. & van der Geer A.A.E. (2025)
On the cranial anatomy of the smallest insular hippopotamus (Cyprus, Pleistocene)
pp. 337-348
doi: 10.4435/BSPI.2025.17
The Pleistocene Cyprus dwarf hippopotamus (c. 132 kg) is the smallest insular hippopotamus known thus far. This species was part of an extremely depauperate fauna, in which the only other large herbivore was represented by a dwarf elephant, in agreement with a high degree of geographic isolation. Here we describe for the first time its cranial anatomy and reconstruct a virtual model of its skull. Unique characters of this taxon are the lack of upper fourth premolar in adult specimens and lophodont teeth. All other cranial phylogenetically relevant characters firmly place the Cyprus dwarf hippo within the genus Hippopotamus. The increased frontation of its orbits and the elevated neutral position of its head are derived features in respect to its hypothetical ancestor, H. antiquus. These features are here explained as correlated to a higher degree of terrestriality and deviation from a grazing lifestyle. The peculiarities of its dentition indicate a wider dietary niche than in other hippopotamids, probably containing a higher percentage of browse intake compared to mainland Hippopotamus. Wear facets on the incisors point to an increased lateral component during chewing, which results in more efficient chewing, probably to cope with potential (seasonal) resource limitations on the island. The mandible is either anteriorly broad with large canines, or narrower with small canines, probably due to sexual dimorphism which was retained from its ancestor. The combination of extreme body size shift, a higher degree of terrestriality, and a change in dietary niche is here hypothesised as an evolutionary adaptation to this island environment, facilitated by ecological release from competition and predation that allowed the species to survive for hundreds of thousands of years.
- Raia P., Castiglione S., De Durante A., Esposito A., Girardi G., Melchionna M., Mondanaro A. & Serio C. (2025)
From linear measurements in multivariate analysis to computational palaeontology
pp. 349-358
doi: 10.4435/BSPI.2025.18
Palaeontology, traditionally rooted in fieldwork and direct observation of fossil remains, is undergoing a transformative shift thanks to technological and mathematical innovations. These advances have expanded the scope and depth of palaeontological research, improving our understanding of evolutionary processes, shape evolution, phylogenetic relationships, and taxonomic diversification. Statistical tools, particularly phylogenetic comparative methods, have become essential for evaluating evolutionary rates and patterns across species. Geometric morphometrics has revolutionised the study of biological form, enabling more accurate reconstructions of fossilised organisms and detailed analyses of evolutionary trends. Additionally, new imaging technologies, such as scanning electron microscopy (SEM) and synchrotron radiation, have enhanced the study of fossils and opened new avenues for detailed analysis. In the last few years, Artificial Intelligence (AI) and machine learning, though still in their early stages, are showing promise for automating fossil classification, identifying patterns in large datasets, and even advancing image-based systematic taxonomy. While AI tools are not yet a replacement for expert palaeontologists, they offer significant support, particularly in curating large collections and facilitating rapid classification processes. However, the integration of all these statistical and technological tools into palaeontological practice presents challenges, particularly in terms of interpreting results accurately. This underscores the need for palaeontologists to develop a foundational understanding of the algorithms and statistical methods they employ, ensuring proper application and reducing the risk of erroneous inferences. As these mathematical and computational tools continue to evolve, they are set to revolutionise palaeontology, enabling more efficient, accurate, and innovative research. Furthermore, these advancements are contributing to the growing field of conservation palaeobiology, with potential applications in understanding climate change, extinction events, and species adaptation, offering critical insights into contemporary conservation efforts.
- Bellucci L., Dominici S., Mazza P.P.A. & Savorelli A. (2025)
The Pliocene and Pleistocene land mammal collections of the Museo di Storia Naturale dell’Università di Firenze: a centuries-old heritage into the third millennium
pp. 359-370
doi: 10.4435/BSPI.2025.19