Massimo Delfino

• PhD
• University of Turin

Transmarine distribution and gigantism in the Late Cretaceous North American crocodyliform Deinosuchus has been difficult to reconcile with consistently inferred phylogenetic relationships to alligatorids, an otherwise freshwater and smaller-bodied group. We present an expanded phylogeny with increased spatiotemporally coherence that reinterprets s...

The collection of fossils supports a thriving market that is largely unknown to the academic community. The fossil market is characterised by fairs, online shops, physical stores and auction sales that attract hundreds of thousands of people each year. The sale of fossils to private individuals is often perceived as a threat by professional palaeon...

We describe a new species of Apus (Aves, Apodidae) from the Early Pliocene of Langebaanweg (Cape Province, South Africa). A single swift humerus was reported from Langebaanweg’s extraordinary rich fossil bird association, but it was never analyzed in detail. More recently, we located a second bone, a carpometacarpus, and completed the comparison of...

The Western whip snake (Hierophis viridiflavus) is a colubroid snake typically considered aglyphous and non-venomous, although this is currently a subject of debate. Indeed, the maxillary dentition of this species has recently been described as opisthoglyphous due to the presence of enlarged rear maxillary teeth, reportedly characterised by promine...

The Italian record of Metriorhynchoidea, a Mesozoic clade of marine crocodylomorphs, is represented by fossils from the Veneto, Lombardy and Sicily regions. A new specimen (FMCR FOS03817) from the Lower Cretaceous of Asiago area (Veneto region, northeastern Italy) is here described for the first time and comparatively studied with a Middle Jurassic...

Urodele osteology is characterised by simplified skulls, loss of several bones and a specific sequence of cranial and limb ossification. The relatively few studies devoted to the comparative analysis of isolated urodele bones are mostly focused on the vertebrae and occipital complexes, and to a lesser extent humeri. The tooth-bearing skeletal eleme...

South Africa is currently home to a highly diverse tortoise assemblage whose evolutionary history has been investigated mostly as a result of molecular studies. The fossil record is of little help because the remains of extant taxa are relatively recent, and only three extinct species, of uncertain relationships, have been described so far. An Earl...

We here describe abundant new fossil material of amphibians and reptiles from different late Neogene localities of northern Greece: the Early Pliocene (MN 14) of Spilia 0, Spilia 1, and Spilia 2; the Early Pliocene (MN 15) of Spilia 3, Spilia 4, Spilia 5, and Vevi; and the Late Miocene or Pliocene of Chalicorrema and Rema Marmara. These new late Ne...

A new parrot species of the genus Agapornis (Aves, Psittaculidae), namely Agapornis longipes nov. sp., is described from the Plio-Pleistocene of Kromdraai, Cooper’s Cave, and Swartkrans in the Cradle of Humankind, South Africa. The new species is represented by all major wing bones (humerus, ulna, and carpometacarpus) and by the tarsometatarsus, to...

This chapter aims to summarize the most relevant peculiarities of the reptile skeleton (organized per major groups) and provide a comprehensive list of reference literature for further deepening autonomously the topic.

Very few remains of elapid snakes are known from the Iberian Peninsula, but these include a probable endemic extinct species of cobra, Naja iberica from the Late Miocene. We here describe isolated cobra vertebrae from several Middle-Late Miocene localities in the Vall es-Pened es Basin (Catalonia, Spain). All of these fossils are herein referred to...

The Late Miocene (Messinian) Gargano “Terre Rosse” in southern Italy has yielded a diverse and peculiar insular vertebrate fauna, comprising an array of endemic taxa, including forms characterized by extreme sizes and unique morphologies. We here present a new testudinid from this insular assemblage. The material comprises few shell remains (both c...

The holotype and only specimen referred to the Early Miocene shark Acanthias bicarinatus Sismonda, 1849 is housed in the collections of the Museo di Geologia e Paleontologia dell’Università degli Studi di Torino and was collected from the serpentinite sandstone of the middle-late Burdigalian Termofourà Formation of the Torino Hill. The specimen, fo...

I geositi sono luoghi di importanza geo-paleontologica e la raccolta di dati inerenti ad essi può dare impulso a future indagini per la loro gestione e conservazione. Qui viene applicata a due siti piemontesi, Cava Ex Fornace RDB e Valle Botto (Pavia et al. 2004), un’innovativa procedura per la valutazione analitica della rilevanza di siti fossilif...

The Early Pleistocene site of Dmanisi (Georgia, Lesser Caucasus), dated to 1.8 Ma, is a relevant locality not only for documenting one of the earliest human evidences in Western Europe, but also for its geographical placement, at the end of a natural corridor that allowed fauna communication between Africa and Eurasia. The amphibian and reptile rem...

This study describes the first fossils of Testudo hermanni hermanni from the island of Sicily (Late Pleistocene, Contrada Pianetti, Ragusa). They are among the most complete Pleistocene tortoise remains of Italy and by far the most informative ones from Sicily. The presence of two complete plastra permits the detection of a character typical of the...

These are the "proceedings" of the SHI scientific congress held in Turin in September 2022. It is a compilation of 21 papers dealing with several aspects of amphibian and reptile biology. Further and separate articles will be shared on Research Gate as well. Those interested in receiving more information could contact the editors.

A chiusura del volume degli atti del XIV Congresso Nazionale della Societas Herpetologica Italica, tenutosi a Torino dal 13 al 17 settembre 2022, vengono celebrati due erpetologi piemontesi di spicco in occasione del bicentenario della nascita di Michele Lessona (1823-1894) e del centenario della scomparsa di Mario Giacinto Peracca (1861-1923).

The Buia study area is in the Dandiero basin (the northernmost part of the Eritrean Danakil depression) and is named after the Buia village situated 100 km SSE of Massawa. In this area a paleoanthropological landmark discovery was made in 1995 when a cranium of an adult individual Homo erectus/ergaster was uncovered in the alternating fluvial, delt...

The hylaeochampsid crocodylomorph Acynodon adriaticus, from the uppermost Cretaceous ‘Villaggio del Pescatore’ site, belongs to an early diverging lineage in Eusuchia. Here an additional specimen, MCSNT 57031, is osteologically and osteohistologically described in detail. After integrating this morphological information together with the recent chr...

The northern Mediterranean reptiles and amphibians show contrasting biogeographic histories during the Plio- Pleistocene. The influence of European climate changes on the evolution of the biogeographic ranges of taxa with a rich fossil record is determined herein combining different proxies to obtain well-supported palaeoclimatic scenarios. Ecolo...

Northern Armenia and southern Georgia, divided in the Haghtanak-Bagratashen area by the Debed River, witnessed considerable volcanic activity between ~2.1 and 1.6 Ma, towards the end of which the earliest evidence of Homo outside Africa is found at Dmanisi, Georgia (~1.77 ma). The rich assemblages of lithic, faunal, and human fossil materials found...

The osteology of European urodeles was studied extensively in the past, but comparative analyses of isolated bones are rare, despite being the most useful tool for the identification of fossil remains. The present work is focused on the most robust skeletal elements (therefore, common in the fossil record: otic–occipitum complexes and vertebrae) an...

This chapter introduces the beginning of Pleistocene vertebrate records, starting from an overview of the Plio-Pleistocene transition. Understanding the Early Pleistocene vertebrate record is impossible without the knowledge of the events characterizing the setting of Early Pleistocene land vertebrate communities. A number of taxa are selected to d...

The disjunct geographical range of many lineages of caudates points to a complex evolutionary and biogeographic history that cannot be disentangled by only considering the present-day distribution of salamander biodiversity. Here, we provide a critical reappraisal of the published fossil record of caudates from the Palearctic and quantitatively eva...

The Eocene fossil reptiles from the Bolca Fossil-Lagerstätte (Verona, Italy) have been known in the literature since at least the 1850’ and were the subject of many studies during the second half of the XIX century and the first decades of the XX century. However, with the exception of a few papers, only rare works have been published on the Monte...

The abstract book of the XIV herpetological congress of the Societas Herpetologica Italica (SHI), with plenary lectures, ordinary talks and poster presentations.

We review in detail the published fossil record of turtles from Sardinia and, in addition, we document previously undescribed specimens for the first time. Among these undescribed specimens, is the oldest occurrence of Testudo hermanni on the island, from the Early Pleistocene of Monte Tuttavista. The turtle fossil record in Sardinia goes back to t...

Archaeological investigations carried out in the cave Zubbio di Cozzo San Pietro, Bagheria, Sicily, revealed the presence of a few skeletal elements of a large-sized tortoise in a funerary area dating to the Copper/Bronze Age. The tortoise has been AMS-dated revealing an age of 12.5 ± 0.5 kyr BP and therefore it pre-dates the funerary activities. T...

Nexus File

Els Casots is one of the richest fossil vertebrate sites of the Vallès-Penedès Basin (Catalonia, Spain). It was discovered in 1989 and excavated briefly during the 1990s, resulting in the recovery of thousands of remains and the erection of several new mammal species. Excavations were resumed in 2018 and continue to date. Here we provide updated re...

In the ‘50s of the XX century, a German expedition lead by Richard Dehm collected a large amount of fossil remains from northern Pakistan. Among these was an isolated trunk vertebra of a lizard, which is here referred to Varanus sp. The collecting site of this specimen is not precisely known, but it most likely comes from middle to early late Mioce...

The fossil record of squamates (i.e. lizards and snakes) from Greece is reviewed herein. Fossil remains of lizards and snakes in Greece date as back as the early Miocene. Their fossil record spans throughout most of the Neogene and the Quaternary and extends in several localities across both continental and insular parts of the country. All major s...

The fossil record of crocodylians from Greece is reviewed herein. Currently absent from the extant herpetofauna of the country (and the whole Europe), however, fossil record attests that crocodylians were present, even if rare and poorly represented (only isolated teeth and one osteoderm have been found so far) during the Miocene. The oldest crocod...

The oldest fossil amphibians from Greece are early Miocene in age, and therefore, only the last few millions of years of amphibian evolution are recorded in this country, and all but one (Mioproteus) of the identified taxa are referable to extant genera. No extinct species of amphibians have been described so far on the basis of Greek fossils. Howe...

Dmanisi (Georgia) is one of the oldest Early Paleolithic sites discovered out of Africa. In addition, it is the best site to understand the first Homo deme out of Africa and the first hominin occupation of Central to Western Eurasia. It has produced more than 40 hominin remains, including several very informative skulls, found in direct association...

Salamandrina is the only extant representative of the Salamandrininae, characterised by a unique suite of morphological, ecological, and ethological features. The Salamandrina species are endemic of Italy, although their past distribution included different areas of Europe. Fossil remains belonging to this genus were previously described from the M...

The sedimentary basins of Greece contain an important record of fossil vertebrates that has been known and studied for nearly two centuries. Here, we present our collective effort to review and summarize this fossil record. A combination of our original research and previously published records permits the complete reassessment of the identified ve...

The two extant Salamandrina species represent a unique case of morphology, ecology, and ethology among urodeles. The range of this genus is currently limited to Italy, where it represents the only endemic vertebrate genus, but its past range extended over a much broader area of Europe, including the Iberian and Balkan peninsulas. ENM analyses using...

Generally, the species is considered to be the only naturally occurring taxon. However, species recognised and defined using different species delimitation criteria cannot readily be compared, impacting studies of biodiversity through Deep Time. This comparability issue is particularly marked when comparing extant with extinct species, because the...

Generally, the species is considered to be the only naturally occurring taxon. However, species recognised and defined using different species delimitation criteria cannot readily be compared, impacting studies of biodiversity through Deep Time. This comparability issue is particularly marked when comparing extant with extinct species, because the...

The title of this book, written in Italian, may be translated as: “Turtles in people’s Culture – Myths, Legends and Folklore”. The author, Mariella Turrini, is an Italian literature teacher loving animals and travels and this book is clearly the outcome of the cross-over of these two beloved passions.

In this study, we describe a new crocodile skull from the world-renowned palaeontological and archaeological site of Olduvai Gorge (Tanzania). The fossil, discovered and collected in 2016 during an annular solar eclipse and therefore nicknamed “Black Sun,” was found in a new trench excavated by the THOR (Tanzania Human Origins Research) team at sit...

The latest Messinian Monticino Quarry fissure fillings, near Brisighella in Italy, are well known for their diverse mammal fauna. Conversely, little is known about other vertebrates from this rich site. beside presenting an overview of the mammals, here we describe fish, amphibian, reptile, and bird remains, identifying three, eight, at least 18, a...

Albanerpetontids are an extinct clade of superficially salamander-like lissamphibians that range from the Middle Jurassic (Bathonian)-Early Pleistocene and have a primarily Laurasian distribution. The best Cenozoic record for the clade is in Europe, where two species in the type genus Albanerpeton Estes & Hoffstetter, 1976 occur in over 40 localiti...

The sedimentary basins of Greece contain an important record of fossil vertebrates that has been known and studied for nearly two centuries. Here, we present our collective effort to review and summarize this fossil record. A combination of our original research and previously published records permits the complete reassessment of the identified ve...

A small sample of fossil vertebrae of amphibians and reptiles found in the historical collection of the Museum of Geology and Palaeontology of the University of Torino is here described. The fossils most likely originate from the same deposits as the old collections from Weisenau, in the Mainz Basin. In spite of the small number of remains, at leas...

Warm-adapted species survived the cold stages of the past glacial-interglacial cycles in southern European peninsulas and recolonized major parts of Central and Northern Europe in the wake of postglacial warming. However, many of the genetic lineages which differentiated within these refugia predate even the Pleistocene. One of these refugia is the...

The Early Pleistocene site of Pietrafitta (central Italy) produced a rich vertebrate assemblage from the Late Villafranchian Land Mammal Age (late MN18). Geological and paleobotanical data from Pietrafitta indicate a lacustrine environment, surrounded by a humid deciduous broadleaved forest with a temperate climate. The vertebrate assemblage consis...

Olduvai Gorge (northern Tanzania) is one of the best known and most iconic palaeontological and archaeological sites in the world. in more than a century of research it has yielded an impressive record of fossils and stone tools which stands as a compendium of human evolution in the context of environmental changes of east Africa in the last 2 Ma....

We here describe a small turtle assemblage originating from the early Tortonian (late Miocene) palaeoisland of Scontrone, central Italy, a locality previously known mostly for its endemic mammals and giant birds, which were otherwise shared only with the Gargano localities, another fossiferous area belonging to the same palaeobioprovince. The fossi...

Several extinct species are known from the family Lacertidae, but due to poor preservation, many of them are based on single bones. Here, we compare phylogenetic signals of disarticulated premaxillae, maxillae and dentaries of lacertids from four French Oligocene localities (Coderet, La Colombière, Roqueprune 2, Mas de Got B). We identified five mo...

The Pliocene–Pleistocene transition in the Neotropics is poorly understood despite the major climatic changes that occurred at the onset of the Quaternary. The San Gregorio Formation, the younger unit of the Urumaco Sequence, preserves a fauna that documents this critical transition. We report stingrays, freshwater bony fishes, amphibians, crocodil...

Turtles of the clade Testudinoidea have a rather scarce fossil record in Afro-Arabia, ranging from the late Eocene up to the Quaternary. The vast majority of testudinoid fossils from Afro-Arabia are ascribed to Testudinidae, which has had a continuous presence in the area since the late Eocene. Geoemydidae is poorly documented by fragments found th...

Once a much more globally widespread crocodylian clade, Tomistominae is today represented by just one species, Tomistoma schlegelii (the false gharial), restricted to south-east Asia. Although tomistomine fossil occurrences are recognized from the early Eocene (∼55 Ma) onwards, their remains are often incomplete, making appropriate taxonomic classi...

The most accepted killing model for the Permian-Triassic mass extinction (PTME) postulates that massive volcanic eruption (i.e., the Siberian Traps Large Igneous Province) led to geologically rapid global warming, acid rain and ocean anoxia. On land, habitable zones were drastically reduced, due to the combined effects of heating, drought and acid...

The site of Coste San Giacomo, in the Anagni Basin, is renowned for its mammal fauna of Early Pleistocene (Gelasian) age, including 32 taxa of small and large mammals. Adding to these, a small amount of remains of amphibians and reptiles was also recovered during recent excavations focused on collecting small vertebrates as well. The analysis of th...

We here describe turtle remains from the late Miocene (Tortonian) of Cessaniti (Calabria, southern Italy), an area that recently has been palaeogeographically reconstructed as being, at that time of the Neogene, directly connected (or at least rather proximate) to northern Africa, instead of Europe. The material pertains to three different turtle c...

In this paper, we describe pleurodire turtle material from the island of Sardinia, Italy, originating from two Eocene localities of the Cixerri Formation. The more complete among the two specimens bears strong resemblance with the continental Western European Eocenochelus eremberti and is tentatively referred to the same species, while the second,...

Vipera walser is the most recently recognized European viper. This rare species is endemic to a small area in the Piedmont Alps of Italy, but its closest relatives are found among the Caucasian viper species. In order to provide a starting point for a phylogenetic and biogeographic investigation based on osteology, and including fossils remains, we...

Recently described species suffer lack of information that hampers setting up appropriate conservation strategies. The situation is particularly complex with microendemic snakes, for which detection and monitoring are particularly challenging. The Walser viper Vipera walser is a recently described snake inhabiting a small area of the SW Italian alp...

Salamandrina perspicillata, from Northern and Central Italy, and Salamandrina ter-digitata, from Southern Italy, represent a unique case of endemism among the amphibians of Europe. Some efforts were made to study the biology and ecology of these species, but only few studies focused on their comparative osteology. In particular , detailed descripti...

This paper presents the recent excavations at the previously unexplored site of RJ-3, in the bay of Ras al-Jinz, one of the key archaeological areas in the Omani Sharquiyyah. During the Umm an-Nar period, RJ-3 most likely formed a single, large settlement with the nearby – and well known – site of RJ-2, located on the other side of the bay. Results...

The dispersal of Crocodylus from Africa to Europe during the Miocene is not well understood. A small collection of cranial fragments and postcranial elements from the latest Miocene (6.2 Ma) site of Venta del Moro (Valencia, Spain) have previously been referred to Crocodylus cf. C. checchiai Maccagno, 1947 without accompanying descriptions. Here we...

Together with the rest of the Balkan Peninsula, Greece acted as a refuge for thermophilic amphibians and reptiles during the Quaternary Glaciations. In this work we provide new herpetological data on the latest Early Pleistocene locality of Kaiafas, in western Peloponnese. The site yielded one indeterminate salamandrid urodelan (Salamandridae indet...

Several localities around the world expose successions of rocks that straddle the Permian–Triassic boundary documenting a common pattern of environmental change. This change testifies to a large-scale event that led to the extinction of a significant portion of biodiversity, the most severe mass extinction of all times. This event is called the End...

Molecular and morphological phylogenies concur in indicating that the African lineages formerly referred to Crocodylus niloticus are the sister taxon the four Neotropical crocodiles (Crocodylus intermedius, C. moreleti, C. acutus and C. rhombifer), implying a transoceanic dispersal from Africa to America. So far the fossil record did not contribute...

This paper presents the recent excavations at the previously unexplored site of RJ-3, in the bay of Ras al-Jinz, one of the key archaeological areas in the Omani Sharquiyyah. During the Umm an-Nar period, RJ-3 most likely formed a single, large settlement with the nearby –– and well known –– site of RJ-2, located on the other side of the bay. Resul...

A certain stability in the Quaternary herpetological assemblages of the Veneto area has been recognised in recent years, however further studies are needed in order to better comprehend their evolutionary dynamics. Amphibians and reptiles from the early Biharian site of Monte La Mesa can prove useful in this respect as the assemblage is composed of...

The skull osteology of Hierophis viridiflavus is here described and figured in detail on the basis of 18 specimens. The sample includes specimens from the ranges of both H. viridiflavus viridiflavus and H. viridiflavus carbonarius as well as specimens not identified at sub-specific level. The main characters that define H. viridiflavus in compariso...

We here redescribe Palaeophis oweni, an almost neglected species of palaeophiid snakes from the Eocene of Monte Duello, northeastern Italy. Despite having been described since the 19th century and being the only known named species of its genus from Mediterranean Europe, P. oweni has so far received only minor and sporadic attention in the literatu...

We here describe turtle remains from the late Miocene (Tortonian) of Cessaniti (Calabria, southern Italy), an area that recently has been palaeogeographically reconstructed as being, at that time of the Neogene, directly connected (or at least rather proximate) to northern Africa, instead of Europe. The material pertains to three different turtle c...

Since the description of Isisfordia duncani, a number of new extinct species and revisions of previously described species have prompted a variety of contradicting phylogenetic hypotheses on the topology of Neosuchia. As a consequence, a consensus on the rooting of Eusuchia in relation to other neosuchian clades has not been reached and the origin...

We herein describe the amphibians and reptiles from the Ptolemais fossil assemblage, originating from 12 nearby localities in northwestern Greece, spanning from the late Miocene (MN 13) to the early Pliocene (MN 15). Amphibians are known exclusively of anurans, with the genera Latonia and Rana being identified, the latter constituting the oldest so...

We herein describe Sardophis elaphoides Georgalis & Delfino n. gen. n. sp., a new snake taxon from the early Pleistocene of Monte Tuttavista VI, Sardinia, Italy. Sardophis elaphoides Georgalis & Delfino n. gen. n. sp. possesses a distinct vertebral anatomy and is diagnosed by a unique combination of features. The new Sardinian taxon is further comp...

The number of precaudal vertebrae in all extant crocodylians is remarkably conservative, with nine cervicals, 15 dorsals and two sacrals, a pattern present also in their closest extinct relatives. The consistent vertebral count indicates a tight control of axial patterning by Hox genes during development. Here we report on a deviation from this pat...

We herein describe the fossil amphibians and reptiles from the Neogene (latest Miocene or earliest Pliocene; MN 13/14) locality of Maramena, in northern Greece. The herpetofauna is shown to be extremely diverse, comprising at least 30 different taxa. Amphibians include at least six urodelan (Cryptobranchidae indet., Salamandrina sp., Lissotriton sp...

Osteological information for extant amphibians and reptiles is scattered through numerous papers and books; this hinders a precise perception of what is known and what is not. In order to aggregate in a synthetic way the available published information and to determine which taxa and topics warrant further attention, we started to compile an online...

Burrowing habits or complex environments have generally been considered as potential drivers acting on reduction and loss of the appendicular skeleton among vertebrates. Herein, we suggest that this might be the case for lissamphibians and squamates, but that fin loss in fishes is usually prevented by important structural constraints, because pecto...

Burrowing habits or complex environments have generally been considered as potential drivers acting on reduction and loss of the appendicular skeleton among vertebrates. Herein, we suggest that this might be the case for lissamphibians and squamates, but that fin loss in fishes is usually prevented by important structural constraints, because pecto...

The fossil record provides evidence of a long evolutionary history of European lizards. Since fossil lizards are regularly represented by bone remains, the knowledge of the origins of extant taxa and their distribution in time and space is hindered by the fact that their comparative osteology is not yet completely and adequately known. In spite of...

We here describe new remains of amphibians and reptiles from the early Miocene (MN 4) of two different Greek localities, Aliveri and Karydia. The newly described material consists of urodelans, alytids, indeterminate anurans, turtles, crocodylians, lacertids, indeterminate scincomorphs, anguids, colubrids, viperids, and indeterminate snakes. The pr...

Diplocynodon is a genus of basal alligatoroid comprising nine species, which spanned the late Paleocene to middle Miocene of Europe. Despite recent revisions of most Diplocynodon species, one of the earliest named and most complete, Diplocynodon hantoniensis, has not been redescribed for over 150 years. This species is known from the remains of num...

Diplocynodon is a genus of basal alligatoroid comprising nine species, which spanned the late Palaeocene to middle Miocene of Europe. Despite recent revisions of most Diplocynodon species, one of the earliest named and most complete, Diplocynodon hantoniensis, has not been re-described for over 150 years. This species is known from the remains of n...

Early Pleistocene terrestrial climate conditions in the Mediterranean region, especially between 1.3 and 1.7 Ma, are poorly understood. Here, the amphibian and reptile fossil record from 24 fissures (Cava Pirro) of the Pirro Nord karstic complex (southern Italy) is used to infer quantitative paleoclimatic and paleoenvironmental reconstructions. The...

Crocodilian remains from the Oligocene fossil locality of Monteviale, northeastern Italy, have historically been referred to different genera, but all material was recently assigned to Diplocynodon cf. D. ratelii Pomel, 1847. The purpose of the present work is to clarify the systematics of the known crocodilian remains from Monteviale. The largest...

• Amphibians form a major component of the diet of the otter Lutra lutra in several areas of its wide geographic range. Yet, amphibian remains are rarely identified to species level and therefore information on the diversity of this food resource is generally scarce. • The aims of this study were: 1) to assess the overall pattern and trends in the...