| Comparative CT images in dorsal view of the paranasal pneumatization and brain morphology of MPND1083 (A-C), Diceros bicornis (D,E), and Ceratotherium simum (F,G). Digitalized natural endocast MPND1083 (H); brain endocasts of D. bicornis (I); and C. simum (J). Sale bars 5 cm.

| Comparative CT images in dorsal view of the paranasal pneumatization and brain morphology of MPND1083 (A-C), Diceros bicornis (D,E), and Ceratotherium simum (F,G). Digitalized natural endocast MPND1083 (H); brain endocasts of D. bicornis (I); and C. simum (J). Sale bars 5 cm.

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Cranial remains of juvenile fossil rhinoceroses are rarely described in literature and very few is known about the ontogenetic development of their inner anatomy. In this study, we report the first CT based description of a juvenile braincase and its natural brain endocast of a late Middle Pleistocene Rhinocerotinae from Melpignano (Apulia, Italy)....

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... the paranasal sinuses, they are characteristic air-filled chambers developed in different cranial bones of placental mammals and are commonly divided into maxillary, frontal and sphenoidal sinuses (Falk, 2009;Boscaini et al., 2018). Processing the CT images of MPND1082, multiple chambers with a diameter just over 1 cm have been evidenced only in the frontal bones, thus corresponding to the frontal sinuses ( Figure 5C). Such a pneumatization is missing inside the occipital, the parietals and the temporals, where instead, abundant spongy bone tissue is observed (Figure 4). ...
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... a pneumatization is missing inside the occipital, the parietals and the temporals, where instead, abundant spongy bone tissue is observed (Figure 4). In dorsal view, the frontal sinuses of MPND1082 are posteriorly delimited by the coronal suture with some terminal lobes that reach the anteriormost portion of the parietals, covering a large part of the frontal lobes of the brain ( Figure 5C). Anteriorly, the air-filled chambers end 2 cm before the frontonasal suture in a completely different assessment compared to those of adult African rhinoceroses, where the frontal sinuses are part of a much extended and more complex system of cranial pneumatization (Figures 5D-G). ...
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... dorsal view, the frontal sinuses of MPND1082 are posteriorly delimited by the coronal suture with some terminal lobes that reach the anteriormost portion of the parietals, covering a large part of the frontal lobes of the brain ( Figure 5C). Anteriorly, the air-filled chambers end 2 cm before the frontonasal suture in a completely different assessment compared to those of adult African rhinoceroses, where the frontal sinuses are part of a much extended and more complex system of cranial pneumatization (Figures 5D-G). ...
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... addition to the digitalized natural endocast MPND1083, a second replica of the brain has been made using the braincase MPND1082 as a digital mold, which includes also a partial 3D model of the paranasal pneumatization (Figures 5A-C). ...
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... this second endocast the size and the anatomical details of the surface are better appreciable. The cerebrum is globular, divided in two hemispheres by a longitudinal fissure both with smooth surface and free of convolutions ( Figure 5C). Some tubular marks interpreted as blood vessels are noticeable along the parietal lobes close to the transverse fissure ( Figure 5C). ...
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... cerebrum is globular, divided in two hemispheres by a longitudinal fissure both with smooth surface and free of convolutions ( Figure 5C). Some tubular marks interpreted as blood vessels are noticeable along the parietal lobes close to the transverse fissure ( Figure 5C). As the consequence of the weak fusion of the sagittal suture, the longitudinal fissure impressed on the digital cast would seem to be projected upwards, but this effect is a graphic artifact originated during the segmentation process. ...
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... the consequence of the weak fusion of the sagittal suture, the longitudinal fissure impressed on the digital cast would seem to be projected upwards, but this effect is a graphic artifact originated during the segmentation process. Indeed, the two main protrusions of the longitudinal fissure correspond with the slits observed along the sagittal suture ( Figure 5C). ...
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... unfused suture of the braincase and the volume of the natural endocast (313 cm 3 ), without the olfactory bulbs and cerebellum, of about half of those obtained for extant adult species (Figures 5H-J) indicate the early ontogenetic status of the fossil specimen. Considering the skull features, the development of the cranial sutures is similar in extinct and extant rhinoceroses species (Hagge, 2010). ...
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... the MPND1083 represents one a few natural brain endocasts described in extinct Rhinocerotinae (Figure 6). MPND1083 consists of a globular telencephalon, free of convolutions with the frontal lobes less expanded than the parietal ones resembling in the general shape those reported on extant Ceratotherium simum and Dicerorhinus sumatrensis (Garrod, 1878;Bhagwandin et al., 2017), whereas it differs from that of Diceros bicornis ( Figure 5I). In dorsal view, the brain of the black rhino shows a very rounded shape without the narrowing at the level of the lateral sulcus, which is evident in the other taxa. ...
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... dorsal view, the brain of the black rhino shows a very rounded shape without the narrowing at the level of the lateral sulcus, which is evident in the other taxa. The different brain morphology of extant D. bicornis and C. simum (Figures 5I,J) has been studied with Magnetic Resonance Imaging by Bhagwandin et al. (2017) and related to diet. According to the authors, the shape of the brains reflects the overall architecture of the skulls, which in turn are related to the feeding habits of the two species, browsing for the black rhinoceros and grazing for the white rhinoceros. ...
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... lack of a detectable pattern of convolution observable on the Melpignano specimen as well as on both the adult African species (Figures 5C,I,J), seem not to be an agerelated trait, since the brain endocasts of juvenile fossil mammals strongly resembles that of adults, indicating that the general brain morphology is almost completely formed in newborn individuals (Falk, 2009;Petrovič et al., 2018). Among gyrencephalic mammals, large herbivores have less complex and very prominent convolutions compared to carnivorans (Roth and Dicke, 2005;Macrini et al., 2007;Hu et al., 2014), consequently, their brain endocasts have fairly smooth appearance amplified by the covering effect of the meninges. ...
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... studies on rhinoceroses are almost completely missing (Gerard et al., 2018) and the development of cranial pneumatization in fossil and extant taxa was poorly investigated. The paranasal sinuses of adult C. simum and D. bicornis form a complex asymmetrical system of breached pneumatization in almost all the cranial bones, including the parietals and the occipital (Figures 5D-G). In contrast, in the MPND1083 specimen the pneumatization concerns only the frontal bones ( Figure 5C) whereas the occipital and the parietals are filled by spongy-bone tissue without pneumatic cavities (Figure 4). ...
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... paranasal sinuses of adult C. simum and D. bicornis form a complex asymmetrical system of breached pneumatization in almost all the cranial bones, including the parietals and the occipital (Figures 5D-G). In contrast, in the MPND1083 specimen the pneumatization concerns only the frontal bones ( Figure 5C) whereas the occipital and the parietals are filled by spongy-bone tissue without pneumatic cavities (Figure 4). It is likely that the areas filled with spongy bones could be developing in paranasal sinuses. ...

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... Paranasal sinuses are air-filled chambers that form inside the cranial bones of many vertebrates and are connected to the nasal cavity (Farke 2010;Iurino et al. 2020;Boscaini et al. 2020b). They are typically found within the frontal and sphenoidal bones but, in several mammals, they extend into other cranial bones, forming in some cases large and complex pneumatic systems (Boscaini et al. 2020b). ...
... Considering this, although caution is needed given the paucity of the sample, among Suidae the shape of the brain would appear to be independent from body size and phylogeny. Indeed, paranasal sinuses and even more brain endocasts are related to several morphofunctional and eco-ethological aspects, and therefore are widely investigated for inferring these characteristics in extinct species (Sakai et al. 2011;Vinuesa et al. 2016;Iurino et al. 2020;Pérez-Ramos et al. 2020;Boscaini et al. 2020a). In a recent work (Bhagwandin et al. 2017), the differences in brain morphology of the extant rhinoceros Diceros bicornis (Linnaeus, 1758) and Ceratotherium simum (Burchell, 1817) have been related to diet. ...
... According to the authors, the shape of the brains reflects the overall architecture of the skulls, which in turn is related to the behaviour and feeding habits of the two species, browsing for the black rhinoceros, and grazing for the white rhinoceros (Bhagwandin et al. 2017). More recently, Iurino et al. (2020) using CT analyses, confirmed the morphological differences in the brain endocasts of D. bicornis and C. simum, and documented a similar arrangement of the cranial pneumatisation in both species. Similarly, S. arvernensis and S. scrofa share similar paranasal sinuses and differently shaped brains. ...
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... Besides comparing semaphoronts of different taxa, the threedimensional reconstruction of the endocranial spaces of individuals of distinct ontogenetic stages can provide key information on the growth and ontogenetic variation in a non-invasive way (Dumont et al., 2020;Hoffmann et al., 2019;Hu et al., 2020;Hublin et al., 2015;Hurlburt et al., 2013;Iurino et al., 2020;Lautenschlager & Hübner, 2013;Macrini et al., 2007;Petroviĉ et al., 2018;Picasso et al., 2010). Therefore, the study of ontogenetic growth of cranial cavities is not only important for understanding species developmental patterns, but also for identifying age-related bias when applying these methodologies to fossil taxa. ...
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... Formal descriptions and studies of selected material of the MSS fossil collection have been reported in a number of papers: equids (Mecozzi & Strani 2021); rhinocerotids (Petronio & Pandolfi 2008;Pandolfi & Petronio 2011;Iurino et al. 2020); bovids ; suids (Iannucci et al. 2020); hyaenids Vinuesa et al. 2016); canids (Iurino et al. 2013;Sardella et al. 2014;Mecozzi & Bartolini Lucenti 2018;Mecozzi et al. 2020) and mustelids Mecozzi 2021). ...
... In the last decades, several authors carried out morphological and biometric studies on the fossil remains from the "terre rosse" of MSS (Petronio & Pandolfi 2008;Pandolfi & Petronio 2011;Petrucci et al. 2012;Iurino et al. 2013Di Stefano et al. 2015;Vinuesa et al. 2016;Iannucci et al. 2020;Iurino et al. 2020;Mecozzi 2021;Mecozzi & Strani 2021). ...
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... The fossiliferous area of Melpignano was finely investigated after the first description provided by Mirigliano (1941) (de Lorentiis 1958Cardini 1962;Bologna et al. 1994). Previously referred to the early Late Pleistocene, the age of the terre rosse has been extended recently to the late Middle Pleistocene (Mecozzi et al. 2019a;Iannucci et al. 2020;Iurino et al. 2020). The badger remains were recovered from the terre rosse of karst cavities from Cava Bianco and Cava Nuzzo (Text- fig. ...
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