dinosaur tracks and bones are widely distributed across Brazil, reflecting a variety ofgeological, paleogeographic, paleoclimatic andpaleoenvironmental contexts. This review paper provides a brief summary of these finds. Skeletal remains are predominantly referred to sauropod species, with fewer specimens belonging to theropods, while ornithischian body fossils are notably wanting. The Cretaceous dinosaur tracks found in the Brazilian syneclises and rift basins are also discussed herein. These tracks are primarily associated with theropods, but also indicate the presence of sauropods, ornithopods and ankylosaurs.
The soft anatomy andskull of the extant Equus (Equidae, Mammalia) help in the understanding of the preorbital fossa (POF) of extinct Equidae. The preorbital fossae which are prevalent in the extinct species are reduced but not absent in Equus. None of the facial muscles of Equus coincide with their facial fossae. The nasolabialis muscle is situated over the shallow preorbital fossa not the levator labii superioris. The levator labii superioris is closer to the orbit; its tendon is supported by a special fascial sling of the nasolabialis. Although the preorbital fossa in the modern horse is externally shallow, it is present as folded bone within that bony fossa. It is also present internally and therefore is not lost. The tholos, the murus reticulatus and the pontis are internal structures of the paranasal sinuses of Equus and may represent parts of the original preorbital fossa. The preorbital fossae in extinct equids are probably independent of the nasolabialis and levator labii superioris muscles. They simply coincide in space and may be functionally unrelated structures. These muscles were most likely attaching on taut membranes that covered the preorbital fossae in extinct taxa. Micro CTs of Merychippus reveal that the floor of the fossa is an extremely thin bone (0.3 mm) and unlikely suitable for muscle support. The nasolacrimal duct disappears in the fossa area. Embryology of mammals supports that the fossa is the duct wide open. Rostral to the fossa, the duct is reconstituted. In other words, it is reassembled. The fossae could contain glands or a sound-producing structure. Fossae most likely were derived from embryonic placodes. Membranes may have covered the fossae. The extinct equids were grazing less than Equus; the fossae are reduced in Equus probably because of its extreme grazing. Apparently, the maxillae are better in supporting the high force and grazing functions than the nasal capsule. In Equus, parts of the fossae are relocated in the paranasal sinuses (e.g., tholos, murus reticulatus). The internal nasal cavity of Merychippus reveals structures homologous to those observed in Equus.
recent (2014-2019) controlled excavations carried out in the Kostanay Region of northern Kazakhstan have provided new data that contribute to the knowledge of the Eocene marine fauna from the Turgay Strait. Herein, we present a report of the cartilaginous and bony fish remains collected during campaigns carried out at two western Asian Eocene localities. The host sediments were deposited in a large epicontinental marine basin throughout the Early Paleogene, when the Turgay Strait connected the Arctic waters to the North and the Peri-Tethys to the South. The bony fish material examined is poorly preserved and is referred in this study to a variety of teleost groups, including eels (Anguilliformes), codfishes (Gadiformes), mackerels (Scombriformes) and indeterminate percomorphs, whereas the chondrichthyans are represented by lamniform sharks. This report documents one of the easternmost occurrences of these groups of fishes in the northern hemisphere during the Paleogene, providing evidence of their presence in the Turgay Strait before its closure.
The unique mass occurrence of tiny heteromorph ammonites found in a single layer in the Mt. Cipollara locality of Cerreto d'Esi (Maiolica Formation, Umbria-Marche Basin, Italy) provides critical insights into the depositional environments of the upper portion of the Cretaceous Maiolica Formation. The mechanisms behind the formation and preservation of this ammonite assemblage within black shales remain poorly understood. To address this knowledge gap, we used biostratigraphy and stable carbon isotopes to analyze the entire section containing the layer rich in ammonites. We then conducted a high-resolution paleoecological analysis on this specific layer. Samples were systematically collected across multiple stratigraphic layers to ensure comprehensive coverage. The focus was on documenting the morphology, abundance, and associated sedimentary structures of the ammonite assemblage. Additionally, sedimentological petrographic examinations were carried out to clarify depositional processes. Our results reveal a diverse ammonite assemblage dominated by the Leptoceratoididae family of early Barremian age, showing relatively good preservation in a predominantly dysoxic, low-energy environment at the seafloor. Calcareous nannofossil data suggest the presence of a well-stratified water column, with a low salinity lid. The multidisciplinary analyses indicate that these black shales not only served as a repository for ammonite remains but also reflected localized paleoecological conditions characterized by reduced turbulence and increased organic deposition. This unique sedimentary and paleontological context suggests that deposition may have been influenced by regional fluctuations in climatic and oceanographic conditions. Consistent with recent literature on Leptoceratoididae, the presence of both normal coiled (oligotrophic) and heteromorphic (mesotrophic) ammonites in the assemblage suggests that the stratification of water masses favored the coexistence of both types, leading to ecological segregation and explaining the paleontological record.
describe a left mandible ofDelphinidae (Cetacea, Odontoceti) from the Lower Pliocene of theAsilah Basin, northwestern Morocco. Despite its fragmentary nature, the specimen reveals key morphological features that support affinities with extant Globicephalinae, i.e., a short mandibular symphysis; a reduced alveolar portion; and a posteroalveolar portion characterized, laterally, by a nearly straight dorsal margin and a ventral margin that sharply curves posteroventrally. However, the Asilah mandible differs from all known extant and fossil globicephalines in its unique character combination, and therefore is here referred to Globicephalinae gen. and sp. indet. pending the discovery of more complete materials. The overall morphology of the reconstructedAsilah mandibles implies a feeding strategy intermediate between suction and grasping. Therefore, this fossil seemingly provides further evidence of a Pliocene generalist delphinid lacking specialized features towards either suction or macropredation, which in turn would have evolved independently multiple times across the family. More generally, this record further supports a high disparity within Delphinidae as early as in Pliocene times, which may correlate with high dietary niche partitioning.
fossil flatfish specimens discovered in laminated diatomites of the Mediterranean Basin are herein re-examined and described. They belong to the single species Arnoglossus sauvagei (Capellini, 1878) (Bothidae, Pleuronectiformes). The latter s' affinities with extinct and extant species of the genus Arnoglossus living in the Mediterranean Sea are discussed.
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 ofMiddle 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.
Lower Kimmeridgian Silenum, Strombecki, and Herbichi (pars) Mediterranean ammonite zones are recognised from the Upper Jurassic succession at Zeng & odblac;v & aacute;rkony (MecsekMountains, Hungary) based on 189 specimens that comprise newly collected material and revision of historical collections. The Silenum Zone is recognised for the frst time from the region. The poorly preserved and fragmented ammonite fauna is typical of the Ammonitico Rosso lithofacies and is of bathypelagic origin: beside corroded, dissolved internal moulds of ammonites, rare lamellaptychi and belemnites are recognised. The ammonite fauna consists of 16 genera and 19 species. The following taxa are recorded for the first time from Hungary: Aspidoceras sesquinodosum, Idoceras sautieri, Lytoceras orsinii, Metastreblites sp. ind., Orthosphinctes (Orthosphinctes) evolutus, O. (O.) polygratus, O. (O.) pseudoachilles, P. progeron. The microfacies are a typical Kimmeridgian wackestone-packstone Saccocoma-microfacies with stray individuals of Stomiosphaera moluccana, as well as rare benthic (Lenticulina) and planktic (protoglobigerinoid) foraminifers. Submediterranean faunal elements (O. [A.] cf. schaireri) are mixed with typical Mediterranean ammonites (Sowerbyceras, Metastreblites, Trenerites) indicating the palaeobiogeographical position of the Tisza microcontinent close to the southern stable European shelf margin in the northern Tethys. The Early Kimmeridgian ammonite assemblage of the Mecsek Mountains is similar to the Eastern Carpathians, UmbriaMarche Apennines, Venetian Alps, SE France (Mt. Crussol), and B & eacute;tic Cordilleras faunas, indicating a northern Tethyan outer shelf environment situated close to the stable European continental margin but with less ratio of phylloand lytoceratid ammonites.
parvulus Daniltshenko, 1968 is a poorly known fish described based on a few scarcely preserved articulated skeletal remains from the basalEocene deposits of the Danata Formation, Kopet-Dag foothills, southwestern Turkmenistan. Despite its peculiar morphology, tProtorhamphosus Daniltshenko, 1968 has been traditionally aligned with the snipefish family Macroramphosidae. However, the relationships of this fish have never been properly discussed and its relevance for the understanding of the syngnathiform fishes remains elusive. Herein a new description of tProtorhamphosus parvulus is provided. The syngnathoid affinities of tProtorhamphosus are confirmed, but the attribution to the Macroramphosidae is questioned. Unfortunately, due to the inadequate preservation of the available material it is not possible to determine the phylogenetic position of this Eocene fish within the syngnathoids.
Few studies have been conducted on Carboniferous marine bivalves in the British Isles and Ireland since pioneering studies performed during the 19thcentury. Herein, we examine from a systematic, taphonomic and palaeoecological point of view, the bivalve fauna occurring as a minor component of brachiopod-dominated fossil assemblages in upper Visean (upper Brigantian) mud mounds in the southern Peak District, Derbyshire (UK). Our results show moderate bivalve diversity, with the fauna being composed of eight genera, representing eight families in four orders (Nuculida, Arcida, Ostreida, Pectinida) and one superorder (Anomalodesmata). Sulcatopinna flabelliformis (Martin, 1809), Aviculopecten planoradiatus M'Coy, 1851, and Cosmomya variabilis (M'Coy, 1851) are herein re-described. Taphonomic bias related to the early dissolution of the aragonitic shell of seminfaunal and infaunal taxa cannot account alone for the scarcity of bimineralic epifaunal pectinids. Indeed, the scarcity of bivalves is also likely due to the presence of the diverse brachiopod community, acting as competitive dominant and ecosystem engineer, well-adapted to the low-turbidity and mesotrophic environment of the mud mound.
Uqua Section of the western Carnic Alps (Italy) represents the reference succession where conodont studies in the Ordovician of southern Europe were initated. The pioneer paper by Serpagli (1967) described the fauna there recovered as morphospecies, in use at that time, with the introduction of several new taxa that were carefully designated but illustrated only with drawings. A biostratigraphically diagnostic conodont fauna of more than 1500 specimens, preliminarily reported by Vai & Spalletta (1980) but never described, is herein analyzed andiguredfor the irst time. The Amorphognathus ordovicicus Biozone is represented by a well-preserved conodont assemblage of 15 species belonging to 15 genera. Common taxa are representatives of Hamarodus brevirameus, Scabbardella altipes andAmorphognathus ordovicicus. They co-occur with less common species including Dapsilodus mutatus, Eocarniodus gracilis andPlectodina alpina. This fauna, documenting the Hamarodus brevirameus-Dapsilodus mutatus-Scabbardella altipes biofacies, is strikingly similar to other occurrences elsewhere in the Carnic Alps. The uppermost calcareous blocks, associated with an Hirnantian brachiopod fauna, do not vary signiicantly in conodont contentfrom the lower levels. The Uqua Section does not reveal the typical Hirnantian conodont signature, indicated by the genera Sagittodontina and Istorinus, reported in the Carnic Alps only from the Cellon Section.
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.
Abundant complete and well-preserved specimens attributed to several carnivorans were recovered from the Calabrian palaeoanthropological site ofDmanisi, Georgia (ca. 1.8Ma). Herein, we describe the hyaenid remains unearthedfrom 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.
cetacean carcasses that sink to the bottom of the deep sea are usually colonised by a wide variety of organisms, forming the so-called whale fall communities. We studied a fragmentary long bone of an unidentified cetacean from the palaeontological collection of the Universitd di Modena e Reggio Emilia collected near Montegibbio (Modena, Italy). The dating of the associated sediments, by means of calcareous nannofossils, indicates a early-middle Tortonian age for the specimen, allowing us to assign it to the Termina Formation. The bone, which shows signs of bioerosion on the outer surface, was analysed using 3D photogrammetry and micro-CT scanning to characterise the external and internal shape of the fossil traces and to understand which organism produced them. Externally, the bone surface shows rounded to irregular pits, a few millimetres to four centimetres wide, where the compact bone has been removed exposing the underlying cancellous bone. Smaller pits are isolated, while larger ones merge. These surface traces are similar to those made by the bone-eating worm Osedax on modern marine vertebrates, a polychaete of the family Siboglinidae that uses a specialised root tissue to penetrate the bones from which it obtains nutrients via endosymbiotic, heterotrophic bacteria. However, only small cavities with a single hole reflect the shape of an individual animal's root system and are of diagnostic value. Using digital segmentation of the scan slices, we isolated, measured and reconstructed the 3D morphology of single boreholes. We distinguished three distinct morphotypes andfound similarities in size and shape with boreholes produced by modern Osedax species. Our findings, which represent the oldest fossil record of Osedax traces from the Mediterranean area, and the firstMiocene record worldwide assessed via micro-CT scan images, are consistent with the presence of a high diversity of cetaceans in the Late Miocene proto-Mediterranean Sea.
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 (Stef & abreve;nescu, 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.
This paper reviews studies of the Upper Valdarno Basin, in order to highlight the unique stratigraphic andpalaeontological 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.
The herein described proboscidean fossils were recovered from two Middle Miocene localities, Dhok Bun Ameer Khatoon (DBAK) and Kohtehra, located in Punjab, Pakistan. The deposits refer to the lower and upper Chinji Formation of the Lower Siwalik Subgroup of the Indian subcontinent. The material includes one deciduous premolar, two premolars, two tusk fragments, a left mandibular fragment with a tusk in the alveolus and an M1. Taxonomically, the material is allocated to three genera Prodeinotherium (Deinotheriidae), Gomphotherium (Gomphotheriidae) and Protanancus (Amebelodontidae). The reported Middle Miocene genera, especially Prodeinotherium and Gomphotherium, are rare and these findings contribute to the substantial morphological and anatomical knowledge of the Siwalik proboscideans. Moreover, the DP3 and lower tusk of Prodeinotherium, and the lower tusk of the adult individual of Protanancus are reported for the first time from the Siwalik Group.
This study delves into the rich collections of Pliocene and Pleistocene fossil vertebrate remains housed in the Museo di Storia Naturale dell'Universitd di Firenze, tracing their origins back to the 17th Century with notable acquisitions by Grand Duke Ferdinando II and Cardinal Leopoldo. Comprising approximately 15,000 fossil remains primarily sourced from the Valdarno region, these collections offer invaluable insights into palaeontological history and the evolution of past ecosystems. The Museum's commitment to excellent curation of these specimens, coupled with its promotion of research initiatives, has significantly enhanced our understanding of Pliocene and Pleistocene vertebrate palaeontology. Through ongoing research efforts, the Museum continues to deepen our knowledge of past climates, environments, and the evolution of life on Earth. This study aims to underscore the significance of these collections, emphasising their pivotal role in advancing scientific knowledge. Furthermore, it pays homage to the visionary leadership of past directors Augusto Azzaroli and Danilo Torre, whose contributions have been instrumental in expanding the Museum's collections and fostering groundbreaking research. The "Valdarno Pliocene and Pleistocene Mammals" collection stand as a testament to the enduring legacy of scientific inquiry and discovery, enriching our understanding of the natural world and has just appointed by the International Commission on Geoheritage as Geoheritage Collection of global importance because of its particularly high scientific, historical and educational relevance for geological sciences.
The Fonelas P-1 site in southern Spain (similar to 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 etruscusForsyth Major, 1877. Morphological and ecomorphological analyses, integrating cranial, dental, and mandibular features, confirm their taxonomic identities and ecological differentiation. Canis arnensis atFonelasP-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.
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 CavaMonticino (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-sizedhyaenids, 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.
