First Attempt to Infer Sound Hearing and Its Paleoenvironmental Implications in the Extinct Insular Canid Cynotherium sardous Studiati, 1857 (Sardinia, Italy).

This is the first study on the bony labyrinth of Cynotherium sardous, an intriguing extinct canid that inhabited Sardinia in the late Middle and Late Pleistocene. The morphological features of the cochlea indicate that C. sardous had a lower number of cochlear turns (2.25) than all extant canids. This feature, as well as the reduced length of the spiral canal, the cochlear curvature rate, and the narrow basal membrane, indicates that C. sardous had poor hearing abilities limited to high-frequency sounds with a low limit of 250 Hz and poor echolocalization skills. From the data available, it is not possible to infer whether C. sardous was unable to echolocalize its prey and relied on other senses (e.g., smell and sight) to locate them or whether the acoustic range of C. sardous was specialized for identifying the sounds produced by its most common prey to transmit signals for predator warnings or group communication. All things considered, the results obtained confirm the utility of cochlea morphological studies in reconstructing the hearing abilities of this species and in providing some suggestions about its ethology, but they fall short of providing any new sound evidence regarding the ecological role of C. sardous in the Late Pleistocene Sardinian ecosystem.

The intriguing giant deer from the Bate cave (Crete): could paleohistological evidence question its taxonomy and nomenclature?

The research describes for the first time a possible case of pituitary gigantism in fossil mammals, precisely in deer. The pathology was detected in 2 long bones (tibia and metatarsus) belonging to an individual of an unusual large size found at the Bate cave (Rethymnon, Northern Crete). It formed the basis of Candiacervus major, the largest among the endemic deer species recorded in the Pleistocene-Early Holocene of Crete. Radiological and histomorphological examinations highlighted a reduction in cortical bone thickness and the presence of wide lacunae inside of the bone tissue. The pathological conditions suggest a pituitary gigantism diagnosis also supported by some morphological evidence, such as the extremely elongated distal part of the metatarsal diaphysis, the proportionally small proximal epiphysis, and some bone gracility. The diagnosis of a case of pituitary gigantism as presumed responsible for the extraordinary elongation of the tibia and the metatarsal bone is intriguing as they are, respectively, the paratype and the holotype of the C. major. The species represents a case of a deviation from the "island rule" in Pleistocene large mammals. The new evidence recommends a taxonomic and nomenclatural revision of this species. The main outcomes of this research are as follows: (i) a case of pituitary gigantism is described for the first time in an extinct mammal; (ii) it is underlined that paleohistology may provide interesting clues for disentangling taxonomic and nomenclatural issues; (iii) one of the very few cases of gigantism in insular mammals is being questioned.

Palaeohistology reveals a slow pace of life for the dwarfed Sicilian elephant.

The 1-m-tall dwarf elephant Palaeoloxodon falconeri from the Pleistocene of Sicily (Italy) is an extreme example of insular dwarfism and epitomizes the Island Rule. Based on scaling of life-history (LH) traits with body mass, P. falconeri is widely considered to be 'r-selected' by truncation of the growth period, associated with an early onset of reproduction and an abbreviated lifespan. These conjectures are, however, at odds with predictions from LH models for adaptive shifts in body size on islands. To settle the LH strategy of P. falconeri, we used bone, molar, and tusk histology to infer growth rates, age at first reproduction, and longevity. Our results from all approaches are congruent and provide evidence that the insular dwarf elephant grew at very slow rates over an extended period; attained maturity at the age of 15 years; and had a minimum lifespan of 68 years. This surpasses not only the values predicted from body mass but even those of both its giant sister taxon (P. antiquus) and its large mainland cousin (L. africana). The suite of LH traits of P. falconeri is consistent with the LH data hitherto inferred for other dwarfed insular mammals. P. falconeri, thus, not only epitomizes the Island Rule but it can also be viewed as a paradigm of evolutionary change towards a slow LH that accompanies the process of dwarfing in insular mammals.

A first comparison of bone histomorphometry in extant domestic horses (Equus caballus) and a Pleistocene Indian wild horse (Equus namadicus).

The microstructural features of the tissue of long bones subjected to different biomechanical stresses could be a helpful tool for a better understanding of locomotor behavior in extant and extinct mammals, including equids. However, few researches have attempted to describe the bone tissue of extinct horses. In our study, we analyze and compare the histomorphometric features of the bone tissue in extant modern horses, Equus caballus, and Equus namadicus, a Pleistocene Indian extinct wild horse. The number, position, and size of the osteons and Haversian canals of the bone tissue, classifiable as dense Haversian tissue, were considered for the comparison. The results obtained highlight some differences between the analyzed species, E. caballus having fewer and bigger osteons than E. namadicus. The microstructural differences may depend on the different lifestyles and environmental conditions characterizing the two species. The results obtained suggest that comparing the biomechanical properties of extinct and modern horse species may provide indirect information on their paleoenvironment.

Insular mammalian fauna dynamics and paleogeography: A lesson from the Western Mediterranean islands.

Since the time of Darwin (1859) and Wallace (1869), islands have been regarded by scientists as a prime target for scrutinizing the forces that may influence evolution and diversification and important elements in biogeographic studies. This research aims to scrutinize whether and to what extent the composition and structure of past mammal insular faunas and their changes through time may provide sound clues for inferring the paleogeographical evolution of a region. As a case study, I critically analyzed the dynamics shown by the Plio-Pleistocene mammalian fauna of 3 Western Mediterranean insular districts, the Balearic Islands, Sardinia and Sicily, each characterized by its own peculiar paleobiogeographical evolutionary history. The revision of faunas and the critical analysis of the dispersal ability of the ancestors of island settlers have allowed hypothesizing on the time and mode of island colonization. The results obtained confirm that the early isolation of the Balearic Islands from the mainland led to the establishment of an endemic fauna since the pre-Messinian Miocene (?Astaracian European Land Mammal Age, MN7/8), and that Sardinia has definitely been isolated since the Pliocene, although dispersal events led to some faunal turnovers during the Pleistocene. In addition, the results suggest for Sicily a complex, still imperfectly disentangled history of alternate phases of complete separation and sporadic, more or less difficult connections with southern Italy.

Palaeoloxodon and human interaction: depositional setting, chronology and archaeology at the Middle Pleistocene Ficoncella site (Tarquinia, Italy).

The Ficoncella site in northern Latium (Italy) represents a unique opportunity to investigate the modalities of a short occupation in an alluvial setting during the Lower Palaeolithic. The small excavation area yielded a lithic assemblage, a carcass of Palaeoloxodon antiquus, and some other faunal remains. The main objectives of the study are to better characterize the depositional context where the Palaeoloxodon and the lithic assemblage occur, and to evaluate with greater precision the occupation dynamics. A 25 m-long well was drilled just above the top of the terrace of the Ficoncella site and faunal and lithic remains were analyzed with current and innovative techniques. The archaeological site contains floodplain deposits as it is located next to a small incised valley that feeds into a larger valley of the Mignone River. A tephra layer capping the site is 40Ar/39Ar dated to 441± 8 ka. Collectively, the geochronologic, tephrochronologic and geologic data, suggest the site was occupied during MIS 13. The new results should prompt further research at Ficoncella in order to improve our understanding of the dynamics of human settlement in Europe during the Early to Middle Pleistocene.

Lights and shadows in the evolutionary patterns of insular bovids.

Endemic bovids are intriguing elements of insular faunas. The living species include the Japanese serow (Capricornis crispus) and the Formosan serow (C. swinhoei), the tamaraw from Mindoro, Philippines, (Bubalus mindorensis) and the anoas (B. depressicornis and B. quarlesi), 2 species of dwarf buffalos endemic to Sulawesi, Indonesia. Fossil endemic bovids are only recorded in some Asian, North American and Western Mediterranean islands. Here we present a comprehensive overview of the changes in body size and evolutionary patterns exhibited by both extant and extinct insular bovids. Our appraisal indicates that each insular representative of Bovidae shows its own peculiar evolutionary model, albeit some parallel trends exist (e.g. reduction in body size, allometric changes in limb bones, alteration of the life history traits). Some changes in morphology (e.g. the simplification of horn cores, the increase in hypsodonty, the acquisition of a 'low-gear' locomotion), for instance, appear as common, albeit not general, patterns triggered by a combination of selective forces. Body size patterns support the 'generality of the island rule' and suggest that biotic interaction had/have a major role in influencing body size evolution in these species, although in different ways on different islands. All things considered, available evidence suggest that a major role in the evolution of insular bovids is played by the structure of the insular community, the nature of available niches and by the dynamics of ecological interactions.

Do methods for predicting paleohabitats apply for mountain and insular fossil bovids?

A variety of methods have been developed to use bovid postcranial elements in the functional morphology approach to paleohabitat prediction. This study represents a first attempt at testing morphometric methods based on astragalus and phalanges on extant mountain-dwelling bovids and insular fossil bovids from Sardinia assigned to the so-called 'Nesogoral group', already regarded as close to the Caprini tribe. We intended to answer the questions whether a classic four-habitats model could be successfully applied to extant mountain-dwelling bovids, and whether results obtained could support the hypothesis of a radiative evolution for the Sardinian bovids. Results obtained, on the one hand, highlighted some inadequacies of the method if not applied to African bovids; on the other hand, they stressed the difficulties of discriminating habitat preferences of Sardinian taxa only based on biometry of astragalus and phalanges. Nonetheless, statistical habitat predictions suggest the contemporaneous presence in Sardinia of bovids having about the same size, but inhabiting different environments, giving support to the hypothesis that Sardinian representatives of the genus Nesogoral originated from a still unknown ancestor by an adaptive radiation evolutionary process.