Bollettino SPI Vol. 61 (2) - OPEN ACCESS!

Cenozoic vertebrates from the central Mediterranean basin: A tribute to the palaeontological legacy of Giovanni Capellini (1833-1922)

Published in September 2022

Guest Editors:
Giorgio Carnevale & Alberto Collareta


  • Carnevale G. & Collareta A. (2022)

Cenozoic vertebrates from the central Mediterranean Basin: A tribute to the palaeontological legacy of Giovanni Capellini (1833-1922)
pp. 87-90

  • Carnevale G., Pellegrino L., Natalicchio M. & Dela Pierre F. (2022)

The Messinian fishes of Capo di Fiume (Palena, Abruzzo): Stratigraphy, taphonomy and paleoecology
pp. 91-118


The Messinian laminated diatomites exposed in the Capo di Fiume stratigraphic section, near the town of Palena, along the slopes of Mt. Porrara in the Abruzzo Apennines, contain well-preserved articulated fish skeletal remains, often associated with plants. A large collection of fossils from this section was accumulated and donated by Erminio Di Carlo to the Museo Geopaleontologico dell’Alto Aventino, Palena, Italy. Herein, we present the stratigraphy of the Capo di Fiume section and a preliminary biosedimentological study of the fossiliferous laminated diatomites. A taphonomic and paleoecological analysis of the fish remains deriving from the oldest diatomite interval of the succession is also included. The sedimentary succession documents the transition from continental to paralic to coastal to open marine conditions, suggesting the existence of a rather narrow shelf connecting coastal and basinal areas. The deposition of the diatomite laminae took place through different biological mechanisms, most notably the so-called “fall dump”, which involved the formation of flocs (mucilaginous aggregates) that included diatom assemblages dominated by Coscinodiscus spp. and Thalassionema nitzschioides. The macrofossil assemblage of the diatomites and the biosedimentological study of the laminated fossiliferous diatomites have provided new data regarding the paleoenvironmental and paleophysiographic setting and the ecological relationships. The diatom content of the fossiliferous (biogenic) laminae as well as the structure and composition of the fish assemblage clearly indicate a depositional marine environment with depths of up to several tens of meters. The fish assemblage is largely dominated by the round herring Spratelloides lemoinei, which represented the trophic nucleus of the original fish community. This and other clupeid species and certain “adventitious visitors” occupied the upper portion of the water column. The “adventitious visitors” are represented by mesopelagic diel vertical migrants (Diaphus edwardsi, Lestidiops sphekodes, Maurolicus cf. muelleri, Myctophum columnae, Paralepis albyi), which were probably attracted in the Capo di Fiume paleobiotope by abundant planktonic organisms as well as by shoals of Spratelloides lemoinei. The lower portion of the water column and the seafloor were well aerated and occupied by a diverse community of demersal fish taxa, especially sparids. The rapid deposition of diatom mats, abundantly represented in the fossiliferous intervals, may have resulted in the swift entombment of fish carcasses, whose fossilization was promoted even under relatively well-oxygenated bottom conditions.

  • Seghetti S.M., Georgalis G.L., Tschopp E. & Delfino M. (2022)

A historical overview of the reptile fauna from the Eocene Bolca Fossil-Lagerstätte (Italy)
pp. 119-143


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 Bolca herpetofauna in recent years, and in many cases knowledge of the anatomy, taxonomy, and proper nomenclature of the Bolca reptiles still remains uncertain. Herein, we reassess the history of the discoveries, the earlier taxonomy and revisions of the crocodilians, turtles, and snakes from Bolca. To date, a total of 13 crocodilian specimens have been described in the literature or are housed in museum collections and remain unpublished. Two of the crocodilian specimens formerly cited and/or described are currently lost. All the fossil crocodilian specimens had originally been referred to two species, Crocodilus vicetinus Lioy, 1865, and Crocodilus bolcensis Sacco, 1895. In this study these identifications are considered invalid, and some specimens are referred to the genera Asiatosuchus, Boverisuchus, Diplocynodon and Hassiacosuchus while assignment at species level is still debatable. The turtles are represented by multiple specimens, which had been referred in the past to several different taxa of pleurodires and trionychids. However, only two species of turtles from Monte Bolca are currently accepted as being valid, both with species epithets dedicated to the renowned Italian palaeontologist Giovanni Capellini (1833-1922): the pleurodire Neochelys capellinii (de Zigno, 1890), which is the type species of its genus, and the trionychid “Trionyxcapellinii Negri, 1892. Both the crocodilians and the turtles had been collected at the Purga di Bolca locality. Only three snake specimens have been described from the Bolca area, representing also the oldest Cenozoic snakes from Italy: Coluber ombonii de Zigno, 1889 from Purga, and the archaeophiine Archaeophis proavus Massalongo, 1859 and anomalophiid Anomalophis bolcensis (Massalongo, 1859) from the Pesciara locality. The affinities of C. ombonii are still not clear, whereas A. proavus and A. bolcensis are considered valid and represent the type species of their genera but also are among the very few representatives of Archaeophiinae and Anomalophiidae respectively. The fossil reptiles from Bolca are housed in public collections in Italy (Turin, Verona, Padua, Rome, Pavia), the United Kingdom (London), USA (Pittsburgh, Cambridge), Germany (Darmstadt, Berlin) and Austria (Vienna).

Supplementary Online Material
  • Pavia M., Cavagna S., Pellegrino I., Pellegrino L. & Carnevale G. (2022)

The oldest fossil record of Buteo (Aves, Accipitridae) from the Late Miocene of Italy and its evolutionary implications
pp. 145-158


An isolated large tusk, belonging to the historical finds of the Collezione di Geologia “Museo Giovanni Capellini” (Bologna, Italy) and originally identified as belonging to a hippopotamus, is here described and reassigned to the genus Metaxytherium (Dugongidae, Sirenia, Mammalia). According to the museum label, this specimen originates from the now exhausted lignite deposits of Montebamboli (Tuscany, central Italy); the latter are late Tortonian to early Messinian in age and were deposited in a lacustrine environment. The Montebamboli tusk displays strong similarities with an elderly Metaxytherium subapenninum specimen from the Pliocene deposits of Bra (Piedmont, northern Italy) as well as with an isolated Metaxytherium tusk, now apparently lost, from Miocene deposits of Son Morelló (Mallorca, Spain). The Late Miocene occurrence of a large-tusked Metaxytherium in the Mediterranean Basin calls into question the anagenetic trend previously proposed for the Euro-North African species of Metaxytherium, thus also stimulating further research on the intra- and interspecific tusk size variability within this lineage. Furthermore, this specimen represents the first record of a marine species from the lignites of Montebamboli, indicating the proximity of marine settings.

  • Pandolfi L., Collareta A., Bianucci G., Contessi M., Rook L. & Sorbi S. (2022)

A large tusk of Metaxytherium (Dugongidae, Sirenia, Mammalia) from the Late Miocene of Montebamboli (southern Tuscany, Italy): evolutionary and paleoecological implications
pp. 159-166


In the Perda S’altari area, north of Fluminimaggiore, black shales of the Silurian Genna Muxerru Formation and limestones of the Devonian Mason Porcus Formation are exposed. The area is strongly tectonized, and poorly preserved fossils occur only in a few outcrops. Graptolites of Telychian and Rhuddanian age have been collected from the shales, whereas conodonts indicate the Lochkovian Icr. postwoschmidti and Ad. transitans zones. The problematic Eurytholia bohemica species is here documented from Devonian rocks for the first time outside the Czech Republic. A gradual transition between the Genna Muxerru Fm. and the Mason Porcus Fm. suggests an update to the lithostratigraphical scheme of the Silurian and Lower Devonian of SW Sardinia.

  • Citron C., Geisler J.H., Collareta A. & Bianucci G. (2022)

Systematics, phylogeny and feeding behavior of the oldest killer whale: a reappraisal of Orcinus citoniensis (Capellini, 1883) from the Pliocene of Tuscany (Italy)
pp. 167-186


Reaching body lengths of 9 m, killer whales (Orcinus orca) are the top mammalian predators of the present day oceans. These distinctive, cosmopolitan dolphins feature an extremely broad and diverse trophic spectrum, ranging from mollusks like octopuses and squids to other toothed whales and even baleen whales. Due to the lack of fossils that can be confidently assigned to Orcinus or close relatives thereof, the evolutionary origin of extant killer whales has rarely been addressed. Here, we provide an updated and thorough reappraisal of the systematics, phylogeny and feeding behavior of Orcinus citoniensis from the Pliocene of Tuscany (Italy), the only fossil species of killer whale currently known on the basis of diagnostic material. Our systematic and phylogenetic reassessment confirms that O. citoniensis is a bona fide species of Orcinus that mostly differs from the extant O. orca by virtue of a smaller body size (ca. 3.5 m). Besides Orcinus spp., the subfamily Orcininae is here recognized as a relatively early branching clade of delphinids that also includes the Italian Pliocene species “Tursiopsosennae, Hemisyntrachelus pisanus and Hemisyntrachelus cortesii. Our morphofunctional analysis supports a mainly piscivorous diet for O. citoniensis. In particular, the degree of apical tooth wear observed on the holotype is consistent with that shown by the extant generalist type of Atlantic killer whales while clearly differing from members of the cetacean-eating specialized types. The prominence of fish in the diet of O. citoniensis is further supported by the fine and shallow microwear features on the dentine exposed at the apical portion of the tooth crown. The emergence of Orcinus as one of the highest trophic level predators of the global oceans, especially at mid and high latitudes, may have involved some process of exaptation, well into the Pleistocene, when large eurytrophic sharks that used to be common and widespread in Pliocene times either became extinct or underwent a severe reduction of their biogeographic ranges.

  • Peri E., Collareta A., Aringhieri G., Caramella D., Foresi L.M. & Bianucci G. (2022)

A new physeteroid cetacean from the Lower Miocene of southern Italy: CT imaging, retrodeformation, systematics and palaeobiology of a sperm whale from the Pietra leccese
pp. 187-206


Herein we describe a new finding of a medium-sized sperm whale from the Burdigalian (Lower Miocene) of the Pietra leccese formation (southern Italy) on the basis of a partly prepared specimen that includes a partial cranium, seven detached teeth, the fragmentary right mandible and two partial vertebral bodies. Because of the overall compression of the specimen, we carried out a retro deformation of a 3D model of the cranium obtained via CT-scanning. The combined analysis of the original specimen and the retrodeformed model has allowed us to recognise that the studied specimen constitutes a new physeteroid taxon: Angelocetus cursiensis n. gen. n. sp., a longirostrine sperm whale characterised by a sideward projected supracranial basin, as evidenced by the overall displacement of its posteriormost margin. Based on a phylogenetic analysis, A. cursiensis n. gen. n. sp. is determined to be a crown physeteroid that does not belong to either the Physeteridae
or the Kogiidae. The wide temporal fossa, elongated rostrum and slender teeth, as well as the skull dimensions (estimated bizygomatic width c. 550 mm) suggest a diet based on medium to large-sized bony fish that were likely captured by a raptorial pierce feeding strategy (as for most of the coeval Burdigalian physeteroids). Despite a seemingly low ecomorphological disparity, the high degree of taxonomic diversity of the Burdigalian physeteroids suggests that this time span represents a crucial phase for the evolutionary history of sperm whales.

Supplementary Online Material
  • Bartolini-Lucenti S., Cirilli O., Pandolfi L. & Rook L. (2022)

The Montopoli site, a reference Local Fauna in the Pliocene and Pleistocene European Large Mammals Biochronology, first discovered by Giovanni Capellini (1833-1922)
pp. 207-222


The Italian paleontologist Giovanni Capellini (1833-1922), internationally renowned for his studies on cetaceans, was the first to discover one of the most important sites for European land mammals biochronology of the Pliocene and Pleistocene, Montopoli (Pisa province; Tuscany). Excavated during the second half of the nineteenth century by the Swiss paleontologist Charles Immanuel Forsyth Major (1843-1923), the fauna from Montopoli differs greatly from the older sites of Italy and Europe. The presence of taxa adapted to more open country environments, typical of arid and progressively cooler habits, contrasts with the faunas of the Early Villafranchian (generally characterized by wooded, tropical/subtropical taxa). This difference led several scholars to regard the fauna of Montopoli as the base of the Middle s. The present study aims to investigate the relationships, in terms of similarity/differences, between the faunal association of Montopoli as compared to other Pliocene and Pleistocene ones from the Old World. Toward achieving this objective we used generic-based resemblance indices and permutative clustering methods attested in literature for their discrimination power. The results of our analysis strengthen the interpretation of the pivotal importance of the faunal association of Montopoli, not only at a regional level but also within a pan-Eurasian Pliocene and Pleistocene framework.

Supplementary Online Material
  • Azzarà B., Breda M., Cirilli O., Madurell-Malapeira J., Ruzza F., Sorbelli L., Tancredi D. & Cherin M. (2022)

Vigna Nuova: the first Middle Villafranchian mammal assemblage from the Valdichiana Basin, Perugia (Italy)
pp. 223-247


The region of Umbria (Central Italy) represents one of the most interesting areas for the study of Mediterranean Plio-Pleistocene mammal faunas due to the occurrence of numerous paleontological deposits. Most of the fossils have been discovered within the Tiberino Basin, which extends across the centre of the territory for ca. 1800 km2, and the small Tavernelle-Pietrafitta Basin, south of Lake Trasimeno. Herein, we provide an additional report on an assemblage within continental deposits from the locality of Vigna Nuova (Piegaro, Perugia) in the Valdichiana Basin, which are observed to crop out in only a few areas of western Umbria. Taphonomic and geological evidence, coupled with rare excavation notes, have allowed the sample to be divided into two assemblages. The younger was collected from the conglomerates (here called “upper layer”) which are still visible on the outcrop and is composed only of remains of cf. Leptobos. The data obtained from this fragmentary sample together with some sedimentological-stratigraphic considerations suggest that it can be correlated with other well-known Valdichiana assemblages (e.g., Selvella, Farneta), dated at ca. 1.5 Ma. The older sub-sample comes from a putative paleosol (here called “lower layer”) no longer visible in the field. It includes cf. Megantereon, Canis sp., Mustelidae indet., Proboscidea indet., Equus cf. senezensis, cf. Leptobos, Croizetoceros ramosus, Pseudodama sp., Cervidae indet. (large size), and Sus cf. strozzii. This assemblage can be referred to the Coste San Giacomo Faunal Unit (late Middle Villafranchian; ca. 2.2-2.1 Ma). It represents the first wellpreserved sample of this age in Umbria and one of the few recorded in Italy, opening new interesting research perspectives on the distribution of mammals within the Peninsula and also on the stratigraphy and evolution of the Valdichiana Basin.

Aggiungeva che per scavare le poche ossa che mi avrebbe subito inviate aveva dovuto faticare enormemente e lottare per due giorni contro ogni sorta di difficoltà. Ricevuto il graditissimo invio mi affrettai a consolidare e restaurare con ogni cura quelle primizie ed oggi sono lieto di poter annunziare, in così solenne adunanza, la interessante scoperta…” (“He added that in order to excavate the few bones that he would immediately send me he had had to work enormously and struggle for two days against all sorts of difficulties. Having received the very welcome shipment, I hastened to consolidate and restore those fruits of the soil with every care and today I am pleased to be able to announce, in such a solemn meeting, the interesting discovery…”).

Giovanni Capellini, 1890

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