Search

The urban area of Rome and its surroundings (the Roman Campaign, “Campagna Romana”) yielded an exceptional amount of fossil remains of terrestrial large mammals, which are of prime relevance for biochronological correlations and for reconstructing the paleoenvironmental conditions in the Italian Peninsula of the Middle Pleistocene. Apart from the mere quantity, the scientific importance of the Roman fossil record rests on the geochronological constraints that allow to date or correlate several findings, as well as on the dense chronological cover of continental deposits, for the last ~0.8 Ma. On the other hand, the case of Rome is emblematic of the challenges posed by investigating historical fossil collections, for instance, the necessity of a thoughtful integration of historical documentation (e.g., geological maps, museum labels) and indirect geological information (e.g., borehole lithology, the correlation between obliterated fossil sites and extant outcrops), owing to the intense urbanization occurred especially since the 1800s. Fossil-rich deposits of Rome have supplied collections housed in the city’s major museums, geosites, and universities, creating a valuable and unique paleontological heritage. Synthesis and perspective on these entwined aspects are provided herein, offering a geological and historical background alongside an overview of Middle Pleistocene mammal faunas of the area of Rome, with special emphasis on recent results that offer examples of - and how to deal with - different kinds of recoveries (from sporadic finds to systematic excavations), and ongoing work on the collection of the University Museum of Earth Sciences of Sapienza University of Rome (MUST). Reviewing the history of the MUST collection underlines the profound link between the history of the research on large mammal faunas of Rome and the history of the collection itself. The management of the paleontological heritage of Rome, consisting of thousands of remains spanning from isolated teeth to complete skeletons, is a crucial task for providing new data and support for research and dissemination, both of which are carried out at MUST accompanying traditional and yet fundamental efforts, such as cataloging and restoration, with the digital enhancement of the collection., Journal of Mediterranean Earth Sciences, Vol. 15 (2023): In progress

LATE MIDDLE TO LATE PLEISTOCENE APULIAN FOSSIL EQUIDAE MELPIGNANO The karst infilling deposits of Melpignano, locally known as ���ventarole���, are located in the area of the village of Maglie. These karst deposits were firstly described by Mirigliano (1941), since then several Institutions, as the IsIPU and Italian Institute of Prehistory and Protohistory (IIPP), with the support of local Salentine Speleological groups, investigated this area (de Lorentiis 1962; Cardini 1962a). The ���ventarole��� are generally filled with reddish sediments (called ���terre rosse���) in the lower part, and brownish sediments (called ���terre brune���) in the upper, particularly rich in vertebrate fossil remains (Bologna et al. 1994) (Fig. 1). The Equus sample was recovered from the ���terre rosse��� of the ���ventarole��� of Mirigliano, Cava Nuzzo and Cava Bianco. Repository and studied material MPUN Mirigliano collection ��� 17 upper teeth, 24 lower teeth; Cava Nuzzo ��� PF 2 hemimandibles, 4 lower teeth (Fig. 3C); Cava Bianco ��� PF 17 upper teeth, 2 hemimandibles, 9 lower teeth. SAN SIDERO The ���ventarole��� of San Sidero are located along the state road (SS16) between the villages of Corigliano d���Otranto and Maglie. The first description of the deposit and its faunal assemblage was reported by Cardini (1962a). The mammal fauna from San Sidero was also studied by other authors (De Giuli 1980, 1983; Petrucci et al. 2012; Iurino et al. 2013, 2015). Similarly to the Melpignano sediments, these ���ventarole��� include ���terre rosse��� layers in the lower part and ���terre brune��� layers in the upper part. The Equidae sample was collected from the ���terre rosse��� of the ���ventarole��� called SS6 and Cava L. Repository and studied material SS6 ��� IGF one skull (Fig. 2A), 11 upper teeth, 10 lower teeth (Fig. 3A); Cava L ��� PF five upper teeth, one hemimandible, six lower teeth. GROTTA DI CAPELVENERE The site, located near the town of Santa Caterina (Lecce), occurs in a Cretaceous limestone (Calcari di Melissano Formation) at 20 m a.s.l. and about 100 m from the current seashore. The cave was discovery in 1960 and was only partially excavated in 1971, 1974 and 1975 (Borzatti von L��wenstern 1961; Giusti 1979, 1980). Outside the cave, a nearby conglomerate deposit at about 8 m a.s.l. has been referred to Tyrrhenian beach (MIS 5) (Patriarchi 1980). The stratigraphic sequence can be divided into two main complexes separated from a speleothem: in the upper part, the brownish sediment, where domestic fauna, ceramics and artefacts appeared, has been referred to Iron age. Instead, in the lower part ten levels including vertebrate fossils and artefacts have been referred to Mousterian (Borzatti von L��wenstern 1961; Giusti 1979, 1980; Patriarchi 1980). The studied sample comes from the lower part of the sedimentary succession. Repository and studied material IGF 18 upper teeth, five hemimandibles, 26 lower teeth. GROTTA DELLE TRE PORTE The fossiliferous site was discovered in 1936during a field survey of the IsIPU conducted by Gian Alberto Blanc (Blanc 1958) (Fig. 1). The coastal cave, located at Punta Ristola, includes three different saloons, but only two contained Pleistocene deposits, Antro del Bambino and Grotta Titti. The Equus material was collected from the level F of Antro del Bambino, associated to Middle Palaeolithic artefacts. Repository and studied material IsIPU eight upper teeth, one hemimandible, nine lower teeth. GROTTA MARIO BERNARDINI The cave located along the Ionian coast near the village of Santa Caterina, also known as Grotta di Santa Margherita, was discovered in 1961 (Borzatti von L��wenstern 1970, 1971) (Fig. 1). The stratigraphic succession consists of four main archeological levels: D, C, B and A, excepted for the top of the sequence, where ceramic and artefacts were referred to Iron and Bronze age (Borzatti von L��wenstern 1970, 1971; Carmigiani & Romagnoli 2017). From the complex VI-III artefacts referred to Mousterian facies were found, whereas those from the complex II were attributed to Uluzzian (Borzatti von L��wenstern 1970, 1971; Carmigiani & Romagnoli 2017). The volcanic material recovered in the top of complex IV (firstly indicated as ss) could be correlated with that found in the level G of the Grotta del Cavallo dated approximately to 109 100 �� 900 ka (Sarti et al. 2002; Spinapolice 2008; Spinapolice 2018; Douka & Spinapolice 2012; Zanchetta et al. 2018). Instead, the volcanic material from the bottom of the complex II (initially indicated as ss ) can be correlated with that found at the top of the level F of the Grotta del Cavallo dated approssimatively to 45700 �� 1 000 ka (Sarti et al. 2002; Spinapolice 2008, 2018; Douka & Spinapolice 2012; Zanchetta et al. 2018). Albeit the mammal remains have never been studied in detail, a preliminary mammal list was provided byBorzatti von L��wenstern (1970, 1971). Equus is found from different levels (VI, IV, III, II), where also lithic artefacts referred to Middle Palaeolithic from the complexes VI-III and Upper Palaeolithic from the complex II were found (Table 1; Appendix 1). Repository and studied material IGF 27 upper teeth, one hemimandible, 27 lower teeth. GROTTA ULUZZO C The cave is located in the Uluzzo Bay near the village of Nard��, opening into the Cretaceous limestone. The stratigraphic succession was described by Borzatti von L��western (1965, 1966) and Borzatti von L��western & Magaldi (1969). In particular, the green volcanic sand from the bottom of the complex II could be correlated with the tephra found at the top of the level F of Grotta del Cavallo, dated at 45700 �� 1000 ka by Zanchetta et al. (2018). Instead, the complex II-I transition, consisting of a grey volcanic sand, could be correlated with the Ignimbrite Campana (CI) identified at the bottom of the level C of Grotta del Cavallo, dated at 39 850 �� 140 ka Zanchetta et al. (2018). The presence of a reworked tephra in the top of the complex IV could represent a marker for the lower deposit. Further investigations needed to confirm the age of these volcanic levels. Moreover, Borzatti von L��western (1965, 1966) reported a preliminary list of the fossil mammals recovered from this locality. The studied sample come from the complex IV, III and II (Table 1; Appendix 2). Whereas, artefacts from the complex IV and III were attributed to Middle Palaeolithic, instead those from the complex II to early Upper Palaeolithic. Repository and studied material IGF 21 upper teeth, two hemimandibles, 33 lower teeth. CASTELLANETA The Castellaneta deposit is a karst infilling into the Pleistocene Calcarenite (Monte Castiglione Formation) (De Giorgi 1877). The fossil material was recovered during the geological survey in the area conducted by Cosimo De Giorgi in the 1870s. Albeit the author delegated the study of the collected sample to Ulderigo Botti, the remains from the karst fissure were never described. Repository and studied material ITCGC 28 upper teeth, 1 hemimandible, 28 lower teeth. GROTTA DEI GIGANTI The coastal cave, located between Punta Ristola and Punta Marchiello, was discovered by the Italian Institute of Human Paleontology during a field survey conducted by Gian Alberto Blanc in the 1930s (Blanc 1958). The authors also reported a mammal faunal list.In addition, Alessio et al. (1978) attempted to carry out radiocarbon dating without success.However, the fossil material was found associated to Middle Palaeolithic artefacts. Repository and studied material IsIPU 12 upper teeth, 20 lower teeth; ITCGC four upper teeth, three lower teeth. GROTTA SANTA CROCE The cave, located near the small village of Bisceglie, was discovered in the 1937 (Segre & Cassoli 1987). The first systematic excavations directed by Luigi Cardini were carried out by Italian Institute of Human Palaeontology (IsIPU) dur - ing the 1950s (Mallegni et al. 1987; Segre & Cassoli 1987; Boscato et al. 2006). The sedimentary succession includes several levels attributed to Middle and Upper Paleolithic and Neolithic (Segre & Cassoli 1987; Arrighi et al. 2009; Ranaldo et al. 2017). Radiocarbon dated indicated an age of 24 900 �� 150 BP for the level B and 31 500 �� 400 BP for the level C (Arrighi et al. 2009; Ranaldo et al. 2017). There are no radiocarbon dating results for levels H-D that are referred to MIS 5 and MIS 4, respectively (Arrighi et al. 2009; Ranaldo et al. 2017). The fossil remains of mammals from the level D were studied by Segre & Cassoli (1987). Recently, new material also was discovered from this level and Equus ferus and Bos primigenius Bojanus, 1827 were identified (Boscato et al. 2006). The studied material was recovered from the level D, associated with lithic artefacts attributed to Middle Palaeolithic. Repository and studied material IsIPU seven upper teeth, seven lower teeth. GROTTA LACEDUZZA The cave deposit, located near the village of San Michele Salentino, was discovered by the ���Gruppo Speleologico Salentino Pasquale de Lorentiis��� in 1970 (Coppola 2005, 2012). The lower part of the sedimentary succession includes fossil remains of mammals and artefacts referred to Middle Palaeolithic. A preliminary mammal list was recently reported, including a rich sample of Meles meles (Linnaeus, 1758) (Mecozzi et al. 2019). The Equus material was recovered from the lower part of the deposit, associated to Middle Palaeolithic artefacts. Repository and studied material MPCCSM nine upper teeth, 12 lower teeth. GROTTA ZINZULUSA The cave is located along the Adriatic coast near the town of Castro. In 1793, Francesco Antonio Del Duca, bishop of the diocese of Castro described the cave in a letter to Ferdinando IV, king of the Kingdom of the Two Sicilies. Nevertheless, the Pleistocene infilling deposit was presented only after a century from the discovery by Botti (1874). The cave consists of several saloons, where Pleistocene deposits were found (Blanc 1962). The sedimentary succession can be divided in two complexes, the lower part, the level B6, where fossil remains of mammals and artefacts referred to Middle Palaeolithic were found, and the upper part, the levels B5-3 including fossil remains of mammals and artefacts referred to Upper Palaeolithic. The Equus material comes from the levels B5-3, where also artefacts referred to Middle Palaeolithic were found. Repository and studied material IsIPU B5-6 upper teeth, five lower teeth. CARDAMONE The karst infilling deposit was discovered by Cosimo De Giorgi in 1872 (Botti 1890). The site is located in a region where several quarries are opened for the extraction of a Plio-Pleistocene calcarenite, and, unfortunately, the deposit was destroyed. The mammal assemblage from Cardamone, initially described by Botti (1890), was recently revised by Rustioni et al. (2003). Based on the presence of the wholly rhino (Coelodonta antiquitatis (Blumenbach, 1799)) and the wholly mammoth (Mammuthus primigenius (Blumenbach, 1799)), the association was referred to ��� Mammuthus-Coelodonta Faunal Complex��� and chronologically attributed to climax of the Last Glacial Maximum (22-18 kyr). Repository and studied material IGF 16 upper teeth, 33 lower teeth; ITCGC two crania (Fig. 2B, C), 13 maxillaries, 83 upper teeth, four hemimandibles, 112 lower teeth (Fig. 3B). FONDO FOCONE The site, discovered during a survey conducted by Decio de Lorentiis in the early 1960s, is located near the village of Ugento. The first excavation campaign was carried out by Luigi Cardini (Cardini 1962b). The fieldwork, direct by Eugenia Segre Naldini, continued during the 1970s, who opened a trench, 3 �� 3 m, called ���Trincea B��� (Cancellieri 2017). Fossil remains of mammals and the artefacts referred to Upper Palaeolithic (early Epigravettian) were found (Cardini 1962b; Cancellieri 2017). Repository and studied material IsIPU one maxillary, 37 upper tooth, one hemimandible, 27 lower teeth. SANTA MARIA D��� AGNANO ��� ESTERNO (SMA-ESTERNO) The Grotta di Santa Maria di Agnano, located near the village of Ostuni, on the north-western margin of Risieddi promontory, was discovered during the 1960s by the Associazione Studi e Ricerche (Studies and Research Association) (Coppola 2012). The excavations in the cave deposit started in 1991 and continued still today, whereas from 2007 the area outside the cave, known as SMA-esterno, was also investigated (Coppola 1992, 2012; Vacca et al. 1992; Vacca & Coppola 1993; Baills 2015; Coppola et al. 2017; Chakroun et al. 2018). The fossil remains come from level 8, dated 25221-24549 cal BP and 26338-25779 cal BP, and levels 6A-4C dated 18013- 17587 cal BP and 16745-16401 cal BP with radiocarbon method (Renault-Miskovsky et al. 2011; Baills 2015). Repository and studied material MPCCSM level eight ��� 10 maxillary, 59 upper tooth, 35 hemimandibles, 67 lower teeth (Fig. 3 D-E). RESULTS Equus samples from the late Middle Pleistocene to Early Holocene of AP show a large variation in dental dimensions and proportions. In particular, the length of P2 and M3 are highly variable both within and between samples (Appendices 3; 4; 7). On the contrary, the length of P 3-4 from San Sidero is closer (p- value> 0.05) to those from Cava Spagnulo, Grotta di Capelvenere, Grotta Mario Bernardini ��� II and Grotta Uluzzo C (Appendix 7). The length of M1-2 from San Sidero is similar (p- value> 0.05) to those from Castellaneta, Fondo Catt��e, Grotta di Capelvenere, Grotta Mario Bernardini ��� III, Grotta Santa Croce, Grotta Zinzulusa, Melpignano and Tana delle Iene (Appendix 7). As in the case of the upper teeth, even the length of P 2 and M 3 show a huge variability (Appendices 5; 6). The P 3-4 from San Sidero is closer (p-value> 0.05) to those from Grotta di Capelvenere, Grotta dei Giganti, Grotta Mario Bernardini ��� IV, Grotta delle Tre Porte, Grotta Zinzulusa and Tana delle Iene (Appendices 5; 6; 8). Whilst, the M1-2 from San Sidero is closer (p-value> 0.05) to Grotta Santa Croce, Grotta delle Tre Porte, Grotta Zinzulusa, Grotta Uluzzo C ��� II, Melpignano and Tana delle Iene (Appendices 5; 6; 8). Considering the MIS chronology, the length of P2 and M3 of the sample from MIS 9-8 is larger only of samples from MIS 3 (P2, p-value 0.05), P3-4 (p-value> 0.05) and M3 (p-value> 0.05) (Fig. 5, Table 3). The protocone index of the Grotta di Capelvenere, Melpignano and San Sidero shows a trend closer to that reported for E. mosbachensis, with M3 value higher than M1-2 one and P3-4 value higher or sub-equal than M1-2 one (Fig. 6). On the contrary, the pattern of the other samples is closer to that reported for E. ferus, with M1-2 value higher than both P3-4 and M3. Finally, the sample from Tana delle Iene and Grotta delle Mura possesses a well different values compared to the others, resembling those reported for Equus hydruntinus. In fact, in E. hydruntinus the values for P3-4 and M1-2 are significantly lower than those of both E. mosbachensis and E. ferus, whereas they are similar in P 2 and M 3 values. Whereas, the postflexid index has been investigated, but no trend can be detected through the time and/or differences among the considered taxa (Fig. 7). Finally, in the standard bivariate plot of muzzle proportions (Fig. 8), two groups can be recognized. The first includes the specimen from the fossiliferous sites referred to glacial stages (Appendix 9), which display a large muzzle in relation to their length. An exception is the cranium of Equus ferus antunesi Cardoso and Eisenmann, 1989 from Fontainhas (Portugal), where the muzzle is longest. The specimen from Cardamone falls in the variability of the glacial horses, and is similar to that from Cuane de l���Arago (Fig. 8). A second group is composed by crania from deposits referred to interglacial stages, where the muzzle is narrow compared to its total length. The cranium from San Sidero falls in this variability. DISCUSSION The taxonomy of Middle to Late Pleistocene European Equus remains controversial (Forst��n 1991; van Asperen 2012). The large variability of the morphological features and biometric traits of caballoid horses has been the subject of controversy amongst many authors. No consensus exists on how to define this variability, as it is either treated as being intra-specific or inter-specific events (Azzaroli 1983; Forst��n 1988; Cramer 2002; van Asperen 2012). This unresolved taxonomic issue has led a proliferation of taxa, identified as either species or subspecies (E. mosbachensis, E. steinheimensis Von Reichenau, 1915, E. achenheimensis Nobis, 1971, E. taubachensis Freudenberg, 1911) (see van Asperen 2012 for discussion). In the Italian fossil record, the specific attribution of the fossil samples from Middle Pleistocene sites reflects this uncertainty (Equus altidens von Reichenau, 1915, Equus aff. sussebornensis, Equus caballus ssp., E. cf. E. mosbachensis Equus caballus cf. mosbachensis and Equus sp.) (Berzi 1972; Caloi & Palombo 1987; Strani et al. 2018, 2019). Preliminary studies on Equus samples from several AP sites led to the identification of E. ferus from the deposits of Cardamone (Rustioni 1998), Cava Spagnulo (Mecozzi et al. 2018), Grotta del Cavallo (Sarti et al. 1998, 2002), Grotta Paglicci (Boscato 1994), Melpignano (Rustioni 1998) and Tana delle Iene (Conti et al. 2010), whereas the previous taxonomic attribution of the material from San Sidero has been questioned. In the revision of the Middle to Late Pleistocene Equus material from Europe, Eisenmann (1991b) attributed the material from San Sidero to the large-sized Equus chosaricus, whereas other authors (De Giuli 1983; Rustioni 1998) classified it as Equus ferus. Unlike, the results of the statistical analysis and the comparison of the protocone index carried out in this work allow to refer the material from San Sidero to E. mosbachensis. In this scenario, the analysis of a relatively large sample of Equus fossils from late Middle to Late Pleistocene localities of AP allows us to reassess the taxonomy and the evolutionary trend of local horse spec, Published as part of Mecozzi, Beniamino & Strani, Flavia, 2022, Equids from the late Middle Pleistocene to Early Holocene of the Apulia Peninsula (southern Italy): reassessment of their taxonomy and biochronology, pp. 17-45 in Geodiversitas 44 (2) on pages 21-32, DOI: 10.5252/geodiversitas2022v44a2, http://zenodo.org/record/5864283, {"references":["MIRIGLIANO G. 1941. - Avanzi di vertebrati quaternari di Melpignano (Lecce). Atti della Regia Academia di Scienze fisiche e matematiche Napoli 2 (4): 2 - 48.","DE LORENTIIS D. 1962. - Nuovi giacimenti nelle fessure della pietra leccese: le ventole ossifere di S. Isidoro e Mepignano a Maglie (Lecce). Quaternaria 5: 297 - 298.","CARDINI L. 1962 a. - Prime determinazioni delle ventole ossifere di S. Isidoro e Melpignano, Maglie. Quaternaria 5: 315 - 318.","BOLOGNA P., DI STEFANO G., MANZI G., PETRONIO C., SARDELLA R. & SQUAZZINI E. 1994. - Late Pleistocene mammals from the Melpignano (LE) \" Ventarole \": preliminary analysis and correlations correlations. Bolletino delaa Societa Paleontologica Italiana It. 33 (2): 265 - 274.","BODDAERT P. 1785. - Elenchus Animalium. Sistens quadrupedia huc usque nota, eorumque varietates. CR Hake. Rotterodami [Rotterdam] 1: 1 - 173. https: // www. biodiversitylibrary. org / page / 28229983","DE GIULI C. 1980. - La fauna di Maglie (Lecce). I Vertebrati fossili italiani, catalogo della Mostra, Verona, 241 p.","DE GIULI C. 1983. - Le faune pleistoceniche del Salento 1. La fauna di San Sidero 3. I Quaderni - Edizioni scientifiche del Museo Comunale di Paleontologia di Maglie 1: 45 - 79.","PETRUCCI M., ROMITI S. & SARDELLA R. 2012. - The Middle- Late Pleistocene Cuon Hodgson, 1838 (Carnivora, Canidae) from Italy. Bollettino della Societa Paleontologica Italiana 51 (2): 137 - 148. https: // doi. org / 10.4435 / BSPI. 2012.15","IURINO D. A., FICO R., PETRUCCI M. & SARDELLA R. 2013. - A pathological Late Pleistocene canid from San Sidero (Italy): implications for social and feeding behaviour. Naturwissenschaften 100 (3): 235 - 243.","IURINO D. A., FICO R. & SARDELLA R. 2015. - A pathological Late Pleistocene badger from San Sidero (Apulia, Southern Italy): Implications for developmental pathology and feeding behaviour. Quaternary International 366: 96 - 101. https: // doi. org / 10.1016 % 2 Fj. quaint. 2014.12.030","BORZATTI VON LOWENSTERN E. 1961. - Un saggio di scavo nella Grotta di Capelvenere, a Santa Caterina di Nardo. Rivista di Scienze Preistoriche 16: 207 - 216.","GIUSTI M. 1979. - La Grotta di Capelvenere a S. Caterina-Nardo (Campagna di scavo 1974). Studi Ecologia Quaternario 1: 19 - 40.","GIUSTI M. 1980. - La Grotta di Capelvenere a S. Caterina-Nardo (Campagna di scavo 1975). Studi Ecologia Quaternario 2: 77 - 85.","PATRIARCHI G. 1980. - Studio sedimentologico e geochimico del paleosuolo dela Grotta di Capelvenere (Nardo-Lecce). S tudi Ecologia Quaternario 2: 87 - 96.","BLANC A. C. 1958. - Giacimenti musteriani con fauna ad elefante, rinoceronti e leone sulla scogliera del Capo di Leuca. La Zagaglia 1: 5 - 14.","BORZATTI VON LOWENSTERN E. 1970. - Prima campagna di scavi nella grotta Mario Bernardini (Nardo Lecce). Rivista di scienze preistoriche 25: 89 - 125.","BORZATTI VON LOWENSTERN E. 1971. - Seconda campagna di scavi nella grotta \" Mario Bernardini \" (Nardo-Lecce). Rivista di scienze preistoriche 26: 31 - 62.","CARMIGIANI L. & ROMAGNOLI F. 2017. - Revisione dell'industria litica musteriana di Grotta Mario Bernardini (Nardo, Lecce). Atti della XLVII Riunione Scientifica Preistoria e Protostoria della Puglia: 1 - 6.","SARTI L., BOSCATO P., MARTINI F. & SPAGNOLETTI A. P. 2002. - Il Musteriano di Grotta del Cavallo-strati H e I: studio preliminare. Rivista di Scienze Preistoriche 52 (21): 45 - 109.","SPINAPOLICE E. E. 2008. - Technologie lithique et circulation des matieres premieres au Paleolithique moyen dans le Salento (Pouilles, Italie meridionale): perspectives comportementales. Doctoral dissertation, Sapienza Universita di Roma and Universite de Bordeaux 1, Rome / Bordeaux, 480 p.","SPINAPOLICE E. E. 2018. - Neanderthal mobility pattern and technological organization in the Salento (Apulia, Italy), in BORGIA V. & CRISTIANI E. (eds), Palaeolithic Italy Advanced Studies on Early Human Adaptations in the Apennine Peninsula. Sidestone Press Acadmics, Leiden, Netherlands: 95 - 124.","DOUKA K. & SPINAPOLICE E. E. 2012. - Neanderthal Shell Tool Production: Evidence from Middle Palaeolithic Italy and Greece. Journal of World Prehistory. 25: 45 - 79. https: // doi. org / 10.1007 / s 10963 - 012 - 9056 - z","ZANCHETTA G., GIACCIO B., BINI M. & SARTI L. 2018. - Tephrostratigraphy of Grotta del Cavallo, Southern Italy: Insights on the chronology of Middle to Upper Palaeolithic transition in the Mediterranean. Quaternary Science Reviews 182: 65 - 77. https: // doi. org / 10.1016 / j. quascirev. 2017.12.014","DE GIORGI C. 1877. - Da Bari al Mar Ionio; appunti geologici per C. De Giorgi. Bolletiono del Reale Comitato Geololigico Italia 7: 239 - 247.","ALESSIO M., ALLEGRI L., BELLA F., IMPROTA S., BELLUOMINI G., CALDERONI G., CORTESI C., MANFRA L. & TURI B. 1978. - University of Rome carbon- 14 dates XVI. Radiocarbon 20: 79 - 104. https: // doi. org / 10.1017 / S 0033822200003921","SEGRE A. G. & CASSOLI P. F. 1987. - Giacimento preistorico del Pleistocene medio e superiore della grotto di S. Croce, Bisceglie (Bari). Atti dell'Istituto Italiano di Preistoria e Protostoria 25: 111 - 118.","MALLEGNI F., PIPERNO M. & SEGRE A. 1987. - Human remains of Homo sapiens neanderthalensis from the Pleistocene deposit of Santa Croce Cave, Bisceglie (Apulia), Italy. American Journal of Physical Anthropology 72: 421 - 429. https: // doi. org / 10.1002 / ajpa. 1330720402","BOSCATO P., CREZZINI J. & PELLEGRINI A. 2006. - Le parti mancanti: faune del Paleolitico Medio nel deposito esterno della Grotta di Santa Croce, Bisceglie (BA), in Tecchiati u. & Sala B. (eds), Studi di Archeozoologia in onore di Alfredo Riedel. Ripartizione Beni Culturali, Ufficio Beni Archeologici, Bolzano: 39 - 50.","ARRIGHI S., FREGUGLIA M., RANALDO F. & RONCHITELLI A. 2009. - Production and use in the lithic industry of the Mousterian in Santa Croce (Bisceglie, Italy). Human evolution 24 (2): 91 - 106.","RANALDO F., ARRIGHI S., FREGUGLIA M., BOSCATO P. & RONCHI- TELLI A. 2017. - La variabilita tecnica del Musteriano della Grotta di Santa Croce (Bisceglie - BT). Studi di Preistoria e Protostoria, 4 - Preistoria e Protostoria della Puglia: 153 - 158.","COPPOLA D. 2005. - Grotta Laceduzza (San Michele Salentino): nuovi rinvenimenti e ipotesi per l'interpretazione di alcuni motivi decorativi nell'eta dei metalli. Spelaion, Martina Franca 149 - 158. http: // hdl. handle. net / 11586 / 51321","COPPOLA D. 2012 - Il riparo di Agnano nel paleolitico superiore: la sepoltura di Ostuni 1 ed i suoi simboli. Terra. Universita di Roma Tor Vergata, Roma, 368 p.","LINNAEUS C. 1758. - Systema naturae per regna tria naturae, secundum Classes, Ordines, Genera, Species, cum characteribus, differentiis, synonymis, locis. Tomus I. Holmie: impensis Direct. Laurentii Salvii, Stockolm: 824 p. https: // www. biodiversitylibrary. org / page / 726886","MECOZZI B., COPPOLA D., IURINO D. A., SARDELLA R. & DE MARINIS A. M. 2019. - The Late Pleistocene European badger Meles meles from Grotta Laceduzza (Brindisi, Apulia, Southern Italy): the analysis of the morphological and biometric variability. The Science of Nature 106: 13. https: // doi. org / 10.1007 / s 00114 - 019 - 1604 - 2","BOTTI U. 1874. - La Zinzolosa. Tipografia di G. Barbera, Tipografia di G. Barbera, Firenze 139 p.","BLANC A. C. 1962. - Relazione sulle osservazzioni e ricerche stratigrafiche eseguite nella Grotta Zinzulusa (Castro Marina, Lecce) e sull'esito di alcuni sopraluoghi nel Salento. 22 Dicembre 1958 - 5 Gennaio 1959. Quaternaria 5: 330 - 334.","BOTTI U. 1890. - La grotta ossifera di Cardamone in Terra d'Otranto. Bollettino della Societa Geologica Italiana 9: 659 - 716.","RUSTIONI M., FERRETTI M. P., MAZZA P., PAVIA M. & VAROLA A. 2003. - The vertebrate fauna from Cardamone (Apulia, southern Italy): an example of Mediterranean mammoth fauna. Deinsea 9 (1): 395 - 404.","CARDINI L. 1962 b. - Sui giacimenti paleolitici di recente scoperti nella penisola salentina. Atti dell'Istituto Italiano di Paleontologia Umana 6: 144 - 148.","CANCELLIERI E. 2017. - Il sito di Fondo Focone-Trincea B (Ugento, Lecce) nel contesto dell'Epigravettiano antico dell'Italia peninsulare orientale, Firenze. Studi di Preistoria e Protostoria 4. Instituto italiano di preistoria e Protostoria della Puglia, Firenze: 79 - 86.","COPPOLA D. 1992. - Nota preliminare sui rinvenimenti nella grotta di S. Maria di Agnano (Ostuni, Brindisi): i seppellimenti paleolitici ed il luogo di culto. Rivista di Scienze Preistoriche 44: 211 - 227.","VACCA E., NOVOTNY V. & PESCE- DELFINO V. 1992. - Note anthropologiche preliminari sui resti della gestante paleolitica di Santa Maria di Agnano (Ostuni, Brindisi). Rivista Sciences, Preistoria 44: 1 - 2.","VACCA E. & COPPOLA D. 1993. - The Upper Paleolithic burials at the cave of Santa Maria di Agnano (Ostuni, Brindisi): preliminary report. Rivista di Antropologia 71: 275 - 284.","BAILLS H. 2015. - Les industries lithiques des occupations Epigravettiens des SMA-esterno (Fouilles 2007 - 2010) (Santa Maria di Agnano, Ostuni, Italie). Bulletin du Musee Anthropologie prehistorique de Monaco 55: 105 - 122.","COPPOLA D., PINTO N. D., PELLEGRINO M., BAILLS H., STASOLLA V., DEWAILLY M., RUSSO J. & BARTOLOMEO J. 2017. - Santa Maria di Agnano (Ostuni, Puglia). Chronique des activites archeologiques de l'Ecole francaise de Rome 1 - 18.","CHAKROUN A., BAILLS H. & COPPOLA D. 2018. - The Site of Santa Maria di Agnano (Brindissi, Italy). Global Jorunal of Archaeology Anthropology 4 (2): 1 - 3.","CARDOSO J. L. & EISENMANN V. 1989. - Equus caballus antunesi, nouvelle sous-espece quaternaire du Portugal. Palaeovertebrata 19: 47 - 72.","FORSTEN A. 1991. - Size decrease in Pleistocene-Holocene true or caballoid horses of Europe. Mammalia 55: 407 - 419. https: // doi. org / 10.1515 / mamm. 1991.55.3.407","VAN ASPEREN E. N. 2012. - Late Middle Pleistocene horse fossils from northwestern Europe as biostratigraphic indicators. Journal of Archaeological Science 39: 1974 - 1983. https: // doi. org / 10.1016 / j. jas. 2012.02.025","AZZAROLI A. 1983. - Quaternary mammals and the \" end-Villafranchian \" dispersal event-a turning point in the history of Eurasia. Palaeogeography, Palaeoclimatology, Palaeoecology 44 (1 - 2): 117 - 139.","FORSTEN A. 1988. - Middle Pleistocene replacement of stenonid horses by caballoid horses - ecological implications. Palaeogeography, Palaeoclimatology, Palaeoecology 65: 23 - 33. https: // doi. org / 10.1016 / 0031 - 0182 (88) 90109 - 5","CRAMER B. 2002. - Morphometrische Untersuchungen and quartaren Pferden in Mitteleuropa. Unpublished PhD thesis, University Tubingen, 250 p.","VON REICHENAU W. 1915. - Beitrage zur naheren Kenntnis fossiler Pferde aus Deutschem Pleistozan, inbesondere uber die Entwicklung und die Abkaustadien des Gebisses vom Hochterrassenpferd. Equus mosbachensis v. R. Abhandlungen Geologischen Landsanstalt Darmstadt 7: 1 - 155.","NOBIS G. 1971. - Vom Wildpferd zum Hauspferd, Studien zur Phylogenie Pleistozaner Equiden Eurasiens und das Domestikationsproblem unserer Hauspferde, Fundamenta Reihe B, Band 6. Koln: Bohlau Verlag.","FREUDENBERG W. 1911. - Beitrage zur Gliederung des Quartars von Weinheim a. d. Bergstrasse, Maure b. Heidelberg, Jockgrim i. d. Pfalz u. a. und seine Bedeutung fur den Bau der oberrheinischen Tiefebene. Notizblatt des Vereins fur Erdkunde und der Grossh. Geologischen Landesanstalt IV 32: 76 - 149.","BERZI A. 1972. - An Early Middle Pleistocene fauna at Monte Oliveto (S. Gimignano, Siena, Italy). Palaeontographia Italica 68: 29 - 33.","CALOI L. & PALOMBO M. R. 1987. - Osservazioni sugli equidi italiani del Pleistocene medio inferiore. Geologica Romana 26: 187 - 221.","STRANI F., DEMIGUEL D., BONA F., SARDELLA R., BIDDITTU I., BRUNI L., DE CASTRO A., GUADAGNOLI F. & BELLUCCI L. 2018. - Ungulate dietary adaptations and palaeoecology of the Middle Pleistocene site of Fontana Ranuccio (Anagni, Central Italy). Palaeogeography, Palaeoclimatology, Palaeoecology 496: 238 - 247. https: // doi. org / 10.1016 / j. palaeo. 2018.01.041","STRANI F., PUSHKINA D., BOCHERENS H., BELLUCCI L., SARDELLA R. & DEMIGUEL D. 2019. - Dietary adaptations of Early and Middle Pleistocene equids from the Anagni basin (Frosinone, central Italy). Frontiers in Ecology and Evolution 7: 176. https: // doi. org / 10.3389 / fevo. 2019.00176","RUSTIONI M. 1998. - Il cavallo e l'idruntino del Salento (Puglia, Italia meridionale) nel quadro degli equidi del Pleistocene superiore dell'Italia. I Quaderni Museo Comunale Paleontologia di Maglie 5: 93 - 121.","SARTI L., BOSCATO P. & MARTO L. M. 1998. - Il Musteriano finale di Grotta del Cavallo nel Salento: studio preliminare. Origini: Preistoria e protostoria delle civilta antiche 22: 45 - 110","BOSCATO P. 1994. - Grotta Paglicci: la fauna a grandi mammiferi degli strati 22 - 24 (Gravettiano antico-Aurignaziano). Rivista di Scienze Preistoriche 46: 145 - 176.","CONTI N., PETRONIO C. & SALARI L. 2010. - The Late Pleistocene Equids from \" Tana delle Iene \" (Ceglie Messapica, Brindisi, Southern Italy). Bollettino della Societa Paleontologica Italiana 49 (2): 227 - 236.","EISENMANN V. 1991 b. - Les chevaux quaternaires europeens (Mammalia, Perissodactyla). Taille, typologie, biostratigraphie et taxonomie. Geobios 24 (6): 747 - 759. https: // doi. org / 10.1016 / S 0016 - 6995 (06) 80303 - 7","BOULBES N. 2010. - Le cheval de Romain-la-Roche, Equus achenheimensis (Mammalia, Perissodactyla) Contribution a la biochronologie des equides caballins au Pleistocene moyen. Revue de Paleobiologie 29 (2): 747 - 770.","GLIOZZI E., ABBAZZI L., ARGENTI A., AZZAROLI A., CALOI L., CAPASSO BARBATO L., DI STEFANO G., ESU D., FICCARELLI G., GIROTTI O., KOTSAKIS T., MASINI F., MAZZA P., MEZZABOTTA C., PALOMBO M. R., PETRONIO C., ROOK L., SALA B., SARDELLA R., ZANALDA E. & TORRE D. 1997. - Biochronology of selected Mammals. Molluscs and Ostracodes from the Middle Pliocene to the Late Pleistocene in Italy. The state of the art. Rivista. Italiana. Paleontologia. S. 103: 369 - 388. https: // doi. org / 10.13130 / 2039 - 4942 / 5299","FICCARELLI G., ABBAZZI L., ALBIANELLI A., BERTINI A., COLTORTI M., MAGNATTI M., MASINI F., MAZZA P., MEZZABOTTA C., NAPO- LEONE G., ROOK L., RUSTIONI M. & TORRE D. 1997. - Cesi, an early Middle Pleistocene site in the Colfiorito Basin (Umbro- Marchean Apennine), central Italy. Journal of Quaternary Science 12 (6): 507 - 518. https: // doi. org / dfmjkf","PALOMBO M. R. & ALBERDI M. T. 2017. - Light and shadows in the evolution of South European stenonoid horses. Fossil Imprint 73 (1 - 2): 115 - 140. https: // doi. org / 10.2478 / if- 2017 - 0006","BIDDITTU I., CASSOLI P. F., RADICATI DI BROZZOLO F., SEGRE A. G., SEGRE- NALDINI E. & VILLA I. 1979. - Anagni, a K-Ar dated Lower and Middle Pleistocene site, Central Italy: preliminary report. Quaternaria 21: 53 - 71.","GUADELLI J. L. 2007. - Etude paleontologique des grands mammiferes, in RAYNAL J. P. (ed.), Sainte-Anne I, Sinzelles, Polignac, Haute-Loire. Le Paleolithique moyen de l'unite J 1. Archeo-Logis / CDERAD, Laussonne: 163 - 188.","UZUNIDIS A. 2017. - Grands herbivores de la fin du Pleistocene moyen au debut du Pleistocene superieur dans le sud de la France. Implications anthropologiques pour la lignee neandertalienne. These de doctorat, Universite d'Aix-Marseille, Marseille, 156 p.","BOULBES N. & VAN ASPEREN E. N. 2019. - Biostratigraphy and Palaeoecology of European Equus. Frontiers in Ecology and Evolution 7: 301. https: // doi. org / 10.3389 / fevo. 2019.00301","STRANI F. 2020. - Impact of Early and Middle Pleistocene major climatic events on the palaeoecology of Southern European ungulates. Historical Biology: 1 - 16. https: // doi. org / 10.1080 / 0 8912963.2020. 1782898","LEONARDI M., BOSCHIN F., GIAMPOUDAKIS K., BEYER R. M., KRAPP M., BENDREY R., SOMMER R., BOSCATO P., MAN- ICA A., NUOGUES- BRAVO D. & ORLANDO L. 2018. - Late Quaternary horses in Eurasia in the face of climate and vegetation change. Science Advances 4 (7): eaar 5589. https: // doi. org / 10.1126 / sciadv. aar 5589","BON M. & BOSCATO P. 1993. - Analisi paleontologica e paleoecologica di macro e micromammiferi dei livelli romanelliani e mesolitici della Grotta delle Mura (Monopoli, Bari). Quaternaria Nova 3: 53 - 104.","DE GROSSI MAZZORIN J. & MONTEFINESE R. 2017. - L'estinzione del cavallo selvatico in Italia e la nuova introduzione allo stato domestico. Congress \" Fisica Nucleare e Preistoria \" Sala Convegni - CEDAD - Universita del Salento.","MAYR E. 1956. - Geographical character gradients and climatic adaptation. Evolution 10: 105 - 108. https: // doi. org / 10.2307 / 2406103","JAMES F. C. 1970. - Geographic size variation in birds and its relationship to climate. Ecology 51: 365 - 390. https: // doi. org / 10.2307 / 1935374","LINDSTEDT S. L. & BOYCE M. S. 1985. - Seasonality, fasting endurance, and body size in mammals. American Naturalist 125: 873 - 787. https: // doi. org / 10.1086 / 284385","BLACKBURN T. M., GASTON K. J. & LODER N. 1999. - Geographic gradients in body size: a clarification of Bergmann's rule. Diversity and Distributions. 5: 165 - 174. https: // doi. org / 10.1046 / j. 1472 - 4642.1999.00046. x","EISENMANN V. 2014. - Main Caballine Crania Morphs. https: // vera-eisenmann. com / main-caballine-crania-morphs","EISENMANN V., CREGUT- BONNOURE E. & MOIGNE A. M. 1985. - Equus mosbachensis et les grands chevaux de la Caune de l'Arago et de Lunel-Viel: craniologie comparee. Bulletin du Museum national d'histoire naturelle, 4 e serie, Section C, Sciences de la terre, Paleontologie, Geologie, Mineralogie 7: 157 - 173. https: // www. biodiversitylibrary. org / page / 55708570","GUERIN C. 1999. - La Fage-aven I (commune de Noailles), middle Pleistocene, and Jaurens (commune de Nespouls), upper Pleistocene. A comparison of two paleontological sites in Correze, France, in GAUDZINSKI S. & TURNER E. (eds), The Role of Early Humans in the Accumulation of European Lower and Middle Palaeolithic Bone Assemblages. Monographie des Romisch-Germanischen Zentralmuseums, vol. 42: 21 - 39.","PHILIPPE M., MOURER C. & EVIN J. 1980. - Les gisements paleontologiques quaternaires des Causses de Martel et de Gramat (Correze et Lot): faunes et chronologie. Publications du Musee des Confluences 18 (1): 57 - 67.","BONIFAY M. F. 1980. - Le Cheval du Pleistocene moyen des grottes de Lunel-Viel (Herault). Gallia prehistoire 23 (2): 233 - 281. https: // doi. org / 10.3406 / galip. 1980.1652","MAUL L. C., REKOVETS L., HEINRICH W. D., KELLER T. & STORCH G. 2000. - Arvicola mosbachensis (Schmidtgen 1911) of Mosbach 2: a basic sample for the early evolution of the genus and a reference for further biostratigraphical studies. Senckenbergiana lethaea 80 (1): 129 - 147."]}

Geodiversitas, In press. - no. 1763, accepted on 10 July 2020