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The Messinian salinity crisis and its precursor events have been the greatest environmental perturbation of the Mediterranean Sea to date, offering an opportunity to study the response of marine ecosystems to extreme hydrological change and a large-scale biological invasion. The restriction of the marine connection between the Mediterranean and the Atlantic Ocean resulted in stratification of the water column and high-amplitude variations in seawater temperature and salinity already from the early Messinian. Here, we present a unified and revised marine fossil record of the Mediterranean (https://doi.org/10.5281/zenodo.13358435, Agiadi et al., 2024) that covers the Tortonian stage, the pre-evaporitic Messinian stage, and the Zanclean stage and encompasses 23 032 occurrences of calcareous nannoplankton, dinoflagellates, foraminifera, corals, ostracods, bryozoans, echinoids, mollusks, fishes, and marine mammals. This record adheres to the FAIR principles, is updated in terms of taxonomy, and follows the currently accepted stratigraphic framework. Based on this record, knowledge gaps are identified, which are due to spatiotemporal inconsistencies in sampling effort and the distribution of sedimentary facies, as well as the inherent differences in the preservation potential between the groups. Additionally, sampling bias in old records may have distorted the record in favor of larger, more impressive taxa within groups. This record is now ready to be used to answer both geological and biological questions about the Mediterranean Sea and beyond and is amendable when new fossil data are brought to light.

The Natural History Museum of the University of Pisa hosts the most important osteological collection of extant cetaceans in Italy as well as one of the most relevant all over Europe. Furthermore, it also preserves a significant palaeontological collection that includes several holotypes and otherwise unique specimens of Archaeoceti (archaic cetaceans), Mysticeti (baleen whales) and Odontoceti (toothed whales). Here, we provide a historical overview of these collections and the corresponding displays, with special attention paid to the origin, development and design of the ‘Archaeocete Hall’ and ‘Cetacean Gallery’. These comprise what may be the largest exhibition worldwide among those dedicated exclusively to cetaceans—one that includes 28 complete skeletons and one skull belonging to 27 extant species as well as fossils of nine extinct species. Our review also reveals that the museum exhibitions feature the oldest known specimen of Mesoplodon bowdoini and the type specimen of Ziphius savi, the latter being a validly described species that is currently regarded as a junior synonym of Ziphius cavirostris. Also significant is the display of several holotype specimens of fossil species such as the protocetid archaeocete Aegyptocetus tarfa, the balaenid baleen whales Balaena montalionis and Balaenula astensis, and the monodontid Casatia thermophila. The Archaeocete Hall and Cetacean Gallery are highly appreciated by visitors as well as perused by the museum’s educational team. The online archiving of 3D models of many of the MSNUP specimens on the open-access digital repository Sketchfab and their subsequent dissemination through the Wikimedia platforms has led to the creation of a major osteological resource—one that is broadly accessible to internet users worldwide.

Abstract Body size is of fundamental importance to our understanding of extinct organisms. Physiology, ecology and life history are all strongly influenced by body size and shape, which ultimately determine how a species interacts with its environment. Reconstruction of body size and form in extinct animals provides insight into the dynamics underlying community composition and faunal turnover in past ecosystems and broad macroevolutionary trends. Many extinct animals are known only from incomplete remains, necessitating the use of anatomical proxies to reconstruct body size and form. Numerous limitations affecting the appropriateness of these proxies are often overlooked, leading to controversy and downstream inaccuracies in studies for which reconstructions represent key input data. In this perspective, we discuss four prominent case studies (Dunkleosteus, Helicoprion, Megalodon and Perucetus) in which proxy taxa have been used to estimate body size and shape from fragmentary remains. We synthesise the results of these and other studies to discuss nuances affecting the validity of taxon selection when reconstructing extinct organisms, as well as mitigation measures that can ensure the selection of the most appropriate proxy. We argue that these precautionary measures are necessary to maximise the robustness of reconstructions in extinct taxa for better evolutionary and ecological inferences.

The lower Messinian Calcare di Rosignano Formation (Tuscany, Italy, 43° N) preserves one of the youngest and northernmost examples of coral reefs in the Mediterranean. The outcropping succession of the Acquabona quarry consists of four main facies, namely, in ascending stratigraphic order: (1) coral boundstone, (2) coralline algal rudstone, (3) serpulid floatstone to packstone, and (4) peloidal packstone to grainstone. The succession displays a trend toward increasingly more shallow conditions and progressively more restricted water circulation. The coral reef displays a limited coral biodiversity and a remarkable abundance of heterotrophs, similar to modern coral reefs developed at the edges of the ecological niche of symbiont-bearing colonial corals. The widespread presence of coral colonies pervasively encrusted by coralline algae and benthic foraminifera suggests that short-term environmental perturbations caused temporary shutdowns of the coral-dominated carbonate factory. Moving upwards, there are fewer corals and more highly adaptable carbonate producers like coralline algae and serpulids. This suggests that the decline of corals had been caused by the conditions in the basin becoming more stressful, up to the collapse of the coral community. The overall succession indicates that coral-dominated ecosystems located at the edges of the coral zone are very sensitive; they can be affected even by minor perturbations and easily collapse if negative conditions persist.

We present a technique for the detection of vertebrate skeletons buried at shallow depths through the use of a ground-penetrating radar (GPR). The technique is based on the acquisition of high-resolution data by medium-to-high frequency GPR antennas and the analysis of the radar profiles by a new forward modelling method that is applied on a set of representative traces. This approach allows us to obtain synthetic traces that can be used to build detailed reflectivity diagrams that plot spikes with a distinct amplitude and polarity for each reflector in the ground. The method was tested in a controlled experiment performed at the top of Cerro Los Quesos, one of the most fossiliferous localities in the Ica Desert of Peru. We acquired GPR data at the location of a partially buried fossil skeleton of a large whale and analyzed the reflections associated with the bones using the new technique, determining the possible signature of vertebrae, ribs, the cranium (including the rostrum), and mandibles. Our results show that the technique is effective in the mapping of buried structures, particularly in the detection of tiny features, even below the classical (Ricker and Rayleigh) estimates of the vertical resolution of the antenna in civil engineering and forensic applications.

The Natural History Museum of the University of Pisa hosts one of the most important osteological collections of cetaceans all over Europe, as well as a conspicuous paleontological collection, including several holotypes of Archaeoceti (archaic whales), Mysticeti (baleen whales), and Odontoceti (toothed whales). In order to valorize these collections, we used 3D technologies to digitize the most relevant specimens, create an online archive, and make the resulting models accessible and shareable with the broadest audience possible through social media profiles and internet browsers. Tens of specimens were surface-scanned using a structured-light scanner, and the resulting 3D models were processed for post-production through the 3D software Blender whenever necessary. All the 3D scans were then gathered in the online repository Sketchfab, which was chosen for its user-friendly interface and common usage among museum institutions. The result is a web page that hosts 35 surface scans of extant and extinct cetacean specimens. This Sketchfab account was linked to the social media (Facebook and Instagram) profiles of the MSNUP to increase the visibility of the museum and promote the dissemination of its outstanding collections of modern and fossil cetaceans. The preliminary results of such an effort are encouraging in terms of views and online interactions. Hopefully, this effort of digitization and online archiving will soon extend to other vertebrate collections.

We report on the geologically youngest occurrence of the softshell turtle family Trionychidae in Europe, from middle Calabrian (Emilian) strata cropping out at Montalto, Pisa Province (Tuscany, central Italy). This record indicates that the softshell turtles survived well past the glacial pulse at ca. 1.8 Ma. That the most recent finds of Trionychidae all over Europe come from mainland Italy further evokes the role of the Apennine peninsula as a refugium for humid-dwelling herpetofaunas through most of the Plio-Pleistocene. Reduced humidity associated with the intensified cool stages at the beginning of the Early–Middle Pleistocene Transition may have been behind the demise of the last European softshell turtles.

Nowadays, the infraorder Delphinida (oceanic dolphins and kin) represents the most diverse extant clade of Cetacea, with delphinids alone accounting for more than 40% of the total number of living cetacean species. As for other cetacean groups, the Early Miocene represents a key interval for the evolutionary history of Delphinida, as it was during this time span that the delphinidans became broadly distributed worldwide, first and foremost with the widespread genus Kentriodon and closely related forms. Here, we report on a new odontocete find from Burdigalian (20.4–19.0 Ma) deposits of the Friulian-Venetian Basin of northeastern Italy, consisting of the partial cranium of a small delphinidan with associated ear bones (right periotic, stapes, malleus and tympanic bulla). Osteoanatomical considerations and comparisons allow us to assign the studied specimen to the genus Kentriodon. This is the first confirmed record of Kentriodon from Europe as well as from the whole proto-Mediterranean region. Stratigraphic and phylogenetic considerations suggest that our new specimen may represent the geologically oldest member of Kentriodon. The evolutionary success of Kentriodon may correlate with the emergence of narrow-band high-frequency echolocation as a possible strategy to escape acoustic detection by large marine predators such as the squalodontids. In addition, the relatively high encephalization quotient of Kentriodon spp. may have provided these early dolphins with some kind of competitive advantage over the coeval non-delphinidan odontocetes.

Abstract A new ichnotaxonomic name, Transexcrementum cuniculus, is applied to tubular (cylindrical) tunnelings in coprolites. The type series of T. cuniculus consists of burrowed vertebrate (probably crocodilian) coprolites that originate from the Miocene Chesapeake Group of Maryland, U.S.A. These complex trace fossils exhibit the following combination of characters: burrows not lined nor backfilled; opening and transverse sections sub-circular; diameter supra-millimetric, up to ca. 20 mm, rather constant throughout; inner termination(s) rounded/conical; tunnel morphology straight or gently curved, sometimes branching; internal sculpturing sometimes present in form of short and irregularly oriented scratches and gouges. Clusters of the same kinds of gouges may also mark the outer surface of the coprolite. The tunneling tracemaker likely engaged in coprophagy; however, it is unclear what kind of organisms could have produced these burrows. Judging from the overall rarity of Transexcrementum cuniculus occurrences in the fossil record, the tracemaker responsible for the burrows might also have been rare, or fed on faeces only occasionally. ZooBank LSID: urn:lsid:zoobank.org:pub:DE755233-4F0E-48D1-A667-7AAD96B56B60

The Cenozoic succession of the East Pisco Basin preserves the sedimentary record of several episodes of deformation of the forearc crust along the Peruvian margin. The 1:50,000 scale geological map presented here encompasses an area of about 1,000 km2 lying astride the Ica River, and contributes to our understanding of the timing and mode of basin filling and deformation. Our novel two-fold megasequence framework provides a sound basis for establishing a first-order tectono-stratigraphic setting of the mid-Eocene–upper Miocene succession exposed in the study area. We interpret that the mid-Eocene to lower Oligocene succession studied in this work (megasequence P) was deposited in a single forearc basin, which was dissected into the present-day West and East Pisco basins by a fault-bounded basement high during the late Oligocene, and subsequently overlain by the Miocene fill of the East Pisco basin (megasequence N).

The lower Messinian Calcare di Rosignano Formation (Tuscany, Italy, 43° N) preserves one of the youngest and northernmost examples of coral reefs in the Mediterranean. The outcropping succession of the Acquabona quarry consists of four main facies, namely, in ascending stratigraphic order: (1) coral boundstone, (2) coralline algal rudstone, (3) serpulid floatstone to packstone, and (4) peloidal packstone to grainstone. The succession displays a trend toward increasingly more shallow conditions and progressively more restricted water circulation. The coral reef displays a limited coral biodiversity and a remarkable abundance of heterotrophs, similar to modern coral reefs developed at the edges of the ecological niche of symbiont-bearing colonial corals. The widespread presence of coral colonies pervasively encrusted by coralline algae and benthic foraminifera suggests that short-term environmental perturbations caused temporary shutdowns of the coral-dominated carbonate factory. Moving upwards, there are fewer corals and more highly adaptable carbonate producers like coralline algae and serpulids. This suggests that the decline of corals had been caused by the conditions in the basin becoming more stressful, up to the collapse of the coral community. The overall succession indicates that coral-dominated ecosystems located at the edges of the coral zone are very sensitive; they can be affected even by minor perturbations and easily collapse if negative conditions persist. [ABSTRACT FROM AUTHOR]

We present a technique for the detection of vertebrate skeletons buried at shallow depths through the use of a ground-penetrating radar (GPR). The technique is based on the acquisition of high-resolution data by medium-to-high frequency GPR antennas and the analysis of the radar profiles by a new forward modelling method that is applied on a set of representative traces. This approach allows us to obtain synthetic traces that can be used to build detailed reflectivity diagrams that plot spikes with a distinct amplitude and polarity for each reflector in the ground. The method was tested in a controlled experiment performed at the top of Cerro Los Quesos, one of the most fossiliferous localities in the Ica Desert of Peru. We acquired GPR data at the location of a partially buried fossil skeleton of a large whale and analyzed the reflections associated with the bones using the new technique, determining the possible signature of vertebrae, ribs, the cranium (including the rostrum), and mandibles. Our results show that the technique is effective in the mapping of buried structures, particularly in the detection of tiny features, even below the classical (Ricker and Rayleigh) estimates of the vertical resolution of the antenna in civil engineering and forensic applications. [ABSTRACT FROM AUTHOR]

The Messinian salinity crisis and its precursor events have been the greatest environmental perturbation of the Mediterranean Sea to date, offering an opportunity to study the response of marine ecosystems to extreme hydrological change and a large-scale biological invasion. The restriction of the marine connection between the Mediterranean and the Atlantic Ocean resulted in stratification of the water column and high-amplitude variations in seawater temperature and salinity already from the early Messinian. Here, we present a unified and revised marine fossil record of the Mediterranean (10.5281/zenodo.13358435, Agiadi et al., 2024) that covers the Tortonian stage, the pre-evaporitic Messinian stage, and the Zanclean stage and encompasses 23 032 occurrences of calcareous nannoplankton, dinoflagellates, foraminifera, corals, ostracods, bryozoans, echinoids, mollusks, fishes, and marine mammals. This record adheres to the FAIR principles, is updated in terms of taxonomy, and follows the currently accepted stratigraphic framework. Based on this record, knowledge gaps are identified, which are due to spatiotemporal inconsistencies in sampling effort and the distribution of sedimentary facies, as well as the inherent differences in the preservation potential between the groups. Additionally, sampling bias in old records may have distorted the record in favor of larger, more impressive taxa within groups. This record is now ready to be used to answer both geological and biological questions about the Mediterranean Sea and beyond and is amendable when new fossil data are brought to light. [ABSTRACT FROM AUTHOR]

A fossil mysticete was discovered along the southwestern coast of Spain, occurring in a block detached from the Neogene deposits exposed along a coastal cliff at the locality of Conil de la Frontera (Cádiz, Spain). These deposits range from Pliocene to Pleistocene in age and include shallow-marine, mixed carbonate–siliciclastic sediments, with the whale being found in occurrence of a stratigraphic unconformity marked by Thalassinoides burrows. 87Sr/86Sr analyses on oyster shells associated with the skeleton suggest an Early Pliocene age, in agreement with the age of the lowermost unit cropping out at the study site. The studied cetacean specimen consists of an articulated, almost complete balaenopteroid skeleton exposed in the field dorsal side up; being contained in an upside-down block, however, it is preserved in ventral disposition. Bones exhibit a low degree of preservation of the cortical bone tissue, which locally features shark bite marks and Osedax traces as well as abundant encrustations of barnacles and ostreids. Two shark teeth were also found near the skeleton. Bones have preserved their main histological features, even though they locally exhibit microcracks, dissolution, substitution by Fe oxides, and microborings. Sediment particles and late diagenetic cements fill the medullary cavities. We propose that the whale carcass experienced refloating before sinking to the seafloor and that the skeleton was probably exposed on the seafloor for some time before being eventually buried.

The Monti Pisani massif (Tuscany, central Italy) is an isolated mountain relief known for its rich geodiversity, including a remarkable palaeontological heritage from the Palaeozoic, Mesozoic and Cenozoic eras. The Palaeozoic record consists of exquisitely preserved plant remains and rarer invertebrates of Permo-Carboniferous age, which testify to extensive rainforests and large swamps that thrived in an alluvial system under a humid, (sub)tropical climate. In addition to invertebrate shells, invertebrate trace fossils and microbial structures, the Mesozoic record features a diverse Middle Triassic tetrapod ichnoassemblage consisting of tracks of lepidosauromorphs, archosaurs (among which are the earliest dinosauromorph fossils of Italy) and nonmammalian therapsids. These vertebrates lived in a subsiding costal setting that stretched across an expanding rift valley under a subarid climate. The Cenozoic record features abundant fossils of terrestrial vertebrates (including spectacular members of the mammalian megafauna) from karst deposits, testifying to the manifold inhabitants of the massif during the glacial and interglacial phases of the Late Pleistocene. Overall, this long-lasting fossil record remarkably demonstrates how much the Earth’s environments have been changing through the Phanerozoic. The outstanding palaeontological heritage of the Monti Pisani area is in need of specific efforts of conservation and valorisation, especially with respect to the many palaeontological sites that punctuate the massif.

The extinct “false mako” shark, Parotodus benedenii (Lamniformes: Otodontidae), is essentially known from large, robust teeth that are widespread but overall rare in Oligocene to Pliocene deposits worldwide. More than 150 years after its description, this species still represents a palaeontological conundrum, as very little is known about its body aspect and palaeoecology. Here, we describe new specimens of P. benedenii from the Pliocene of Tuscany, central Italy. These new finds comprise some of the geologically youngest finds of P. benedenii worldwide, witnessing to the survival of false makos until the Late Pliocene at least, which in turn suggests that P. benedenii may have been the latest surviving member of the family Otodontidae. Building upon a thorough literature review, we provide an updated synthesis of the palaeobiology of P. benedenii. In light of the morphological evidence, and considering previously published suggestions, P. benedenii may be reconstructed as a large-sized, carnivorous shark that dwelt in pelagic settings and fed primarily on large, soft prey and scavenging items. Thus, some ecological partitioning did likely exist between P. benedenii and other elasmobranch apex predators of the Neogene mid-latitude seas (including, in Pliocene times, the extant species Carcharodon carcharias, Carcharhinus leucas and Galeocerdo cuvier).

The white shark, Carcharodon carcharias, is the main top predator of the present-day Mediterranean Sea. The deep past of C. carcharias in the Mediterranean is witnessed by a rather conspicuous, mostly Pliocene fossil record. Here, we provide a synthesis of the palaeobiology and palaeoecology of the Mediterranean white sharks. Phenetically modern white shark teeth first appeared around the Miocene–Pliocene transition in the Pacific, and soon after in the Mediterranean. Molecular phylogenetic analyses support an origin of the Mediterranean white shark population from the dispersal of Australian/Pacific palaeopopulations, which may have occurred through the Central American Seaway. Tooth dimensions suggest that the Mediterranean white sharks could have grown up to about 7 m total length during the Pliocene. A richer-than-today marine mammal fauna was likely pivotal in supporting the Mediterranean white sharks through the Pliocene and most of the Quaternary. White sharks have seemingly become more common as other macropredators declined and disappeared, notwithstanding the concurrent demise of many potential prey items in the context of the latest Pliocene and Quaternary climatic and environmental perturbations of the Mediterranean region. The overall generalist trophic habits of C. carcharias were likely crucial for securing ecological success in the highly variable Mediterranean scenario by allowing the transition to a mostly piscivorous diet as the regional marine mammal fauna shrank.

We report 130 vertebrate fossils preserved as bony elements and the co-occurring assemblage of fish teeth and spines from the lower strata of the Pisco Formation exposed along the western side of the lower Ica Valley (East Pisco Basin, Peru). Geological mapping at 1:10,000 scale reveals that all these fossils originate from the Langhian–Serravallian P0 allomember. In the study area, P0 is up to ∼40 m thick and features a sandy lower portion, reflecting shoreface deposition, that fines upwards into a package of offshore silts. Marine vertebrates only occur in the lower sandy layers and include whales, dolphins, reptiles, birds, and bony and cartilaginous fishes. The reconstructed paleoenvironment is consistent with a warm-water, marginal-marine setting with a strong connection to the open ocean. This work helps to elucidate the rich yet still poorly understood middle Miocene portions of the Pisco Formation, and highlights the need to conserve this outstanding Fossil-Lagerstätte.

We report on the geologically youngest occurrence of the softshell turtle family Trionychidae in Europe, from middle Calabrian (Emilian) strata cropping out at Montalto, Pisa Province (Tuscany, central Italy). This record indicates that the softshell turtles survived well past the glacial pulse at ca. 1.8 Ma. That the most recent finds of Trionychidae all over Europe come from mainland Italy further evokes the role of the Apennine peninsula as a refugium for humid-dwelling herpetofaunas through most of the Plio-Pleistocene. Reduced humidity associated with the intensified cool stages at the beginning of the Early–Middle Pleistocene Transition may have been behind the demise of the last European softshell turtles. [ABSTRACT FROM AUTHOR]

Body size is of fundamental importance to our understanding of extinct organisms. Physiology, ecology and life history are all strongly influenced by body size and shape, which ultimately determine how a species interacts with its environment. Reconstruction of body size and form in extinct animals provides insight into the dynamics underlying community composition and faunal turnover in past ecosystems and broad macroevolutionary trends. Many extinct animals are known only from incomplete remains, necessitating the use of anatomical proxies to reconstruct body size and form. Numerous limitations affecting the appropriateness of these proxies are often overlooked, leading to controversy and downstream inaccuracies in studies for which reconstructions represent key input data. In this perspective, we discuss four prominent case studies (Dunkleosteus, Helicoprion, Megalodon and Perucetus) in which proxy taxa have been used to estimate body size and shape from fragmentary remains. We synthesise the results of these and other studies to discuss nuances affecting the validity of taxon selection when reconstructing extinct organisms, as well as mitigation measures that can ensure the selection of the most appropriate proxy. We argue that these precautionary measures are necessary to maximise the robustness of reconstructions in extinct taxa for better evolutionary and ecological inferences. [ABSTRACT FROM AUTHOR]

The Natural History Museum of the University of Pisa hosts the most important osteological collection of extant cetaceans in Italy as well as one of the most relevant all over Europe. Furthermore, it also preserves a significant palaeontological collection that includes several holotypes and otherwise unique specimens of Archaeoceti (archaic cetaceans), Mysticeti (baleen whales) and Odontoceti (toothed whales). Here, we provide a historical overview of these collections and the corresponding displays, with special attention paid to the origin, development and design of the 'Archaeocete Hall' and 'Cetacean Gallery'. These comprise what may be the largest exhibition worldwide among those dedicated exclusively to cetaceans—one that includes 28 complete skeletons and one skull belonging to 27 extant species as well as fossils of nine extinct species. Our review also reveals that the museum exhibitions feature the oldest known specimen of Mesoplodon bowdoini and the type specimen of Ziphius savi, the latter being a validly described species that is currently regarded as a junior synonym of Ziphius cavirostris. Also significant is the display of several holotype specimens of fossil species such as the protocetid archaeocete Aegyptocetus tarfa, the balaenid baleen whales Balaena montalionis and Balaenula astensis, and the monodontid Casatia thermophila. The Archaeocete Hall and Cetacean Gallery are highly appreciated by visitors as well as perused by the museum's educational team. The online archiving of 3D models of many of the MSNUP specimens on the open-access digital repository Sketchfab and their subsequent dissemination through the Wikimedia platforms has led to the creation of a major osteological resource—one that is broadly accessible to internet users worldwide. [ABSTRACT FROM AUTHOR]