Manuscript Details

Manuscript number GEOBIO_2018_44_R1

Title Biochronological framework for the late Galerian and early-middle Aurelian Mammal Ages of peninsular Italy

Article type Research paper

Abstract Following a recent chronostratigraphic revision of 17 fossiliferous sites hosting assemblages constituting local faunas of the Aurelian Mammal Age (AMA) for peninsular Italy, we provide a re-structured biochronological framework and discuss the current validity and significance of the Middle Pleistocene Faunal Units (FU) for this region. In contrast with the previous model of a wide faunal renewal during Marine Isotope Stage (MIS) 9 (~330 ka), the First Occurrences (FO) of several species of the Torre in Pietra FU are significantly backdated and referred to the Fontana Ranuccio FU (530-400 ka). We show that the faunal renewal was more gradual and occurred earlier than previously assumed. Many taxa that are typical of the Late Pleistocene register their FO in the Fontana Ranuccio FU, latest Galerian, which is characterized by the almost total disappearance of Villafranchian taxa and by the persistence of typical Galerian taxa such as Dama clactoniana, Bison schoetensacki and Ursus deningeri, and by the FO of Stephanorhinus kirchbergensis, S. hemitoechus, Hippopotamus amphibius, Cervus elaphus eostepahnoceros, Ursus spelaeus, Canis lupus, and Vulpes vulpes. The next Torre in Pietra FU is characterized only by the FO of Megaloceros giganteus and of Mustela putorius. However, we observe that MIS 9 marks the actual moment when the faunal assemblages of this region are represented only by those taxa characterizing the late Middle Pleistocene and Late Pleistocene. For this reason, we propose to consider the Torre in Pietra (lower levels) local fauna still as a conventional boundary for the Galerian-Aurelian transition. Finally, we remark that the strong faunal renewal in MIS 13, with six FOs, coincides with the temperate climatic conditions due to the absence of marked glacial periods that could have favored the FOs and the subsequent spread of these taxa.

Keywords Galerian; Aurelian; Mammal Age; biochronology; chronostratigraphy; Middle Pleistocene; peninsular Italy

Corresponding Author fabrizio marra

Corresponding Author's Istituto Nazionale di Geofisica e Vulcanologia Institution

Order of Authors Carmelo Petronio, Giuseppe Di Stefano, Anastassios Kotsakis, Leonardo Salari, fabrizio marra, Brian Jicha

Suggested reviewers Danielle Schreve, Gennady Baryshnikov, Dimitris Kostopoulos, Frederic Lacombat, Lorenzo Rook, Adrian Lister

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There are no linked research data sets for this submission. The following reason is given: Data will be made available on request Dear Gilles, please, consider the revised version of the paper "Biochronological framework for the late Galerian and early-middle Aurelian Mammal Ages of peninsular Italy", by C. Petronio, G. Di Stefano, T. Kotsakis, L. Salari, F. Marra, B. Jicha that I'm re-submitting to Geobios for possible publication. Besides describing the modifications performed according to the reviewers comments, I would like to drove your attention on the slight modifications we provided to the chronostratigraphy of two sites treated in our paper, in order to have your approval for including them in the revised version of our manuscript. In light of very recent field work I performed for an ongoing research project (Boschian et al., in progress) aimed at providing further direct age constraints to the geologic sections hosting archeological materials (both lithic and faunal assemblages) in the investigated area, I had the chance to acquire new elements allowing at refining the chronostratigraphic setting in two sites (i.e. Malagrotta and km 19.3 of Via Aurelia) described in the present paper submitted to Geobios. While the proposed slight modifications do not affect any of the attributions to the Faunal Units of the fossils that are the subject of the paper, and therefore have no implication for our topic, I believe that the paper would benefit from this more detailed information and it is worth to include it in this revised version. Indeed, this modification concerns the possibility to distinguish better between the MIS 13 and MIS 11 deposits at the abovementioned investigated sections, and since the faunal assemblages occurring within these deposits are part of the same Faunal Unit (Fontana Ranuccio FU), their allocation to MIS 11 or MIS 13, yet implying a difference in age of ca. 100 ky, is not influent regarding the FU of attribution. Therefore, I hope you may approve the introduction of these modifications in the revised text, along with those performed to address the reviewers comments and suggestions. Regarding these latter, please see detailed explanations provided hereby.

We have provided all the modification requested in your editorial points.

Reviewer #1

We accepted all the modifications/corrections provided in the annotated pdf and we have re- formulated the sentences marked as unclear. Concerning the suggestion to provide "visual" information about previous biochronologic framework, we think that an additional figure may be redundant with the detailed report in section '2. Historical background and paleontological setting', and we have addressed this issue by marking in bold in Figure 3 the FO of the taxa which were previously included within MIS 9.

Reviewer #2

We have considered all the insightful comments and suggestions by this reviewer and we have provided modifications or argued our statements as follows.

We would like to specify that our contribution discusses the relocation in late Galerian, Middle Pleistocene, of the FO of some taxa previously referred to the early Aurelian, late Middle Pleistocene (and not to Late Pleistocene as inferred by the reviewer). We would like to clarify that our biochronological review is not based solely on literature data, but that a large part of the fossil remains have been directly revised and, when necessary, taxonomically re-determined, as they are kept in the Paleontological Museum of the Sapienza University of Rome. Therefore, we have added a new paragraph in the methods section in which we more clearly report this fact.

Regarding the observation that a characterizing point of a FO should be the simultaneous appearance of a species in geographically distanced regions, we agree and remark that we implicitly account for this concept, both in the introduction and in the discussion, by highlighting that following the proposed biochronologic revision the faunal renewal in Italy is more synchronous with the faunal renewal in Western Europe.

We have more extensively discussed the taxonomic status of the wolf of Castel di Guido; however, in our opinion, the rank of chrono-subspecies for the little wolf of Lunel Viel, C. lupus lunellensis, remains valid in agreement with many other authors (eg, Brugal and Boudadi-Maligne, 2011; Boudadi-Maligne, 2012; Palombo, 2014; Sardella et al., 2014; Sansalone et al., 2015);

We have extensively discussed the taxonomic status of the Cava Rinaldi bear in the evolutionary context U. deningeri - U. spelaeus, also incorporating most of the suggested references (e.g., Rabeder, 1999; Baryshnikov, 2006; Dabney et al., 2013);

We have eliminated any reference to differences in diet between U. deningeri and U. spelaeus;

We have well received the suggested reference (Baryshnikov, 2011) about the wildcat.

In light of these modifications through which we attempted at receiving all the comments and suggestions provided by the reviewers we are now confident that our paper may be judged fit for publication. Thank you very much for your kind attention, Best regards,

Fabrizio Marra Cited references: Baryshnikov, G., 2006. Morphometrical variability of cheek teeth in cave bears. Scientific Annals School of Geology AUTH, special vol. 98, 81-102. Baryshnikov, G.F., 2011. Pleistocene Felidae (Mammalia, Carnivora) from the Kudaro Paleolithic cave sites in the Caucasus. Proceedings of the Zoological Institute RAS 315, 197-226. Boudadi-Maligne, M., 2012. Une nouvelle sous-espèce de loup (Canis lupus maximus nov. subsp.) dans le Pléistocène supérieur d’Europe occidentale. A new subspecies of wolf (Canis lupus maximus nov. subsp.) from the upper Pleistocene of Western Europe. Comptes Rendus Palevol 11, 475-484. Brugal, J.P., Boudadi-Maligne, M., 2011. Quaternary small to large canids in Europe: Taxonomic status and biochronological contribution. Quaternary International 243, 171-182. Dabney, J., Knapp, M., Glocke, I., Gansauge, M.-T., Weihmann, A., Nickel, B., Valdiosera, C., García, N., Pääbo, S., Arsuaga, J.-L., 2013. Complete mitochondrial genome sequence of a Middle Pleistocene cave bear reconstructed from ultrashort DNA fragments. Proceedings of the National Academy of Sciences 110, 15758-15763. Palombo, M.R., 2014. Deconstructing mammal dispersals and faunal dynamics in SW Europe during the Quaternary. Quaternary Science Reviews 96, 50-71. Rabeder, G. 1999. Die Evolution des Höhlenbärengebisses. Mitteilungen der Kommission für Quartärforschung der Österreichischen Akademie der Wissenschaften 11, 1-102. Sansalone, G., Bertè, D.F., Maiorino, L., Pandolfi, L., 2015. Evolutionary trends and stasis in carnassial teeth of European Pleistocene wolf Canis lupus (Mammalia, Canidae). Quaternary Science Review 110, 36-48. Sardella, R., Bertè, D., Iurino, D.A., Cherin, M., Tagliacozzo, A., 2014. The wolf from Grotta Romanelli (Apulia, Italy) and its implications in the evolutionary history of Canis lupus in the Late Pleistocene of Southern Italy. Quaternary International 328, 179-195. REVIEWER 1 We accepted all the modifications/corrections provided in the annotated pdf and we have re- formulated the sentences marked as unclear. Concerning the suggestion to provide "visual" information about previous biochronologic framework we have addressed this issue by marking in bold in Figure 3 the FO of the taxa which were previously included within MIS 9.

REVIEWER 2

Authors, basing on the new stratigraphic data, reconsidered a time of appearance for a number of mammal taxa in the paleontological record of Italian peninsula as well as revised species compositions of faunal units on the border between the Galerian and Aurelian Mammal Age. They conclude that several species, such as Ursus spelaeus, Canis lupus, and Vulpes vulpes, appear to be characteristic for the Middle Pleistocene stage of evolutionary history, whereas preceding researchers referred them to the Late Pleistocene.

This assumption is very interesting and could be regarded as convincing one, if the authors would use not only the newest data on dating of localities and faunal units, but carried out a taxonomic revision of determinations provided earlier. Unfortunately, the authors did not revised earlier determinations and limited themselves only by published data.

We would like to clarify that our biochronological review is not based solely on literature data, but that a large part of the fossil remains have been directly revised and, when necessary, taxonomically re-determined, as they are kept in the Paleontological Museum of the Sapienza University of Rome. Therefore, we have added a new paragraph in the methods section in which we more clearly report this fact.

First Occurrences (FO), which is a special focus by authors, is determined by a time of origin of a species or by a time of its dispersal from adjacent territories. The history of dispersal of Homo to Asia and America as well as the data on mammal acclimatization show that the formation of even large distribution ranges may occur very rapidly. Therefore, the important confirmation for FO should be the simultaneous appearance of a species in geographically distanced regions. Authors poorly use this argument.

We agree with this important observation by the reviewer and we remark that we implicitly account for this concept, both in the introduction and in the discussion, by highlighting that following the proposed biochronologic revision the faunal renewal in Italy is more synchronous with the faunal renewal in Western Europe.

They note that earliest findings of Canis lupus came from the early Middle Pleistocene of Beringia and its dispersal into Europe occurred in the late Middle Pleistocene (p. 23). However the remains of large-sized C. lupus are recorded in Europe about 350 Ka, whereas earlier levels (400-600 Ka) were the place of finding of small-sized C. l. lunellensis or C. lunellensis (Palombo & Valli 2004). Therefore, it would be desirable to define more precisely the taxonomic status for the wolf from MIS13.

It seems to be necessary to provide morphological grounds for the extremely early appearance (about 516 Ka) of Ursus spelaeus. The paper has no references to the modern detailed studies on the morphological evolution (Rabeder 1999; Rabeder et al. 2010; Baryshnikov 2006; Baryshnikov & Puzachenko 2011, 2017, etc.) and genetics (Knapp et al. 2009; Dabney et al. 2013; Stiller et al. 2014) of cave bears. In Austria, cave bears of similar age are referred to U. deningeroides. In addition, the time of split of sister European clades, spelaeus and ingressus, is evaluated approximately as 170 Ka (Stiller et al. 2014), whereas the divergence between U. deningeri and U. spelaeus might occur nearly 300-400 Ka (Dabney et al. 2013).

The authors report that U. deningeri had less specialized dental morphology than U. spelaeus and regarded it to be more carnivorous. However the analysis of stable isotopes revealed the vegetarian diet for U. kudarensis from Caucasus (Bocherens et al. 2014), despite this mitochondrial clade had isolated earlier than U. deningeri (Dabney et al. 2013). I think differences in diet between U. deningeri and U. spelaeus were not considerable and depended on the local conditions.

We have eliminated any reference to differences in diet between U. deningeri and U. spelaeus;

It could be added to the debates on the evolution of Felis silvestris (p. 26) that this species has been found in the Middle and Late Pleistocene of Caucasus (Baryshnikov 2011).

We have well received the suggested reference (Baryshnikov, 2011) about the wildcat. 1 Biochronological framework for the late Galerian and early-middle Aurelian

2 Mammal Ages of peninsular Italy

3 4 Petronio C.1, Di Stefano G.1, Kotsakis T.2, Salari L.1, Marra F.3*, Jicha B.R.4 5 6 1 Dipartimento di Scienze della Terra, Sapienza, Università di Roma, Rome, Italy 7 8 2 Dipartimento di Scienze, sezione di Geologia, Università degli Studi “Roma Tre”, Rome, 9 Italy 10 11 3 Istituto Nazionale di Geofisica e Vulcanologia, Rome, Italy 12 13 4 Department of Geoscience, University of Wisconsin-Madison, USA 14 15 16*corresponding author: [email protected] 17 18 Abstract

19 Following a recent chronostratigraphic revision of 17 fossiliferous sites hosting assemblages

20 constituting local faunas of the Aurelian Mammal Age (AMA) for peninsular Italy, we provide a re-

21 structured biochronological framework and discuss the current validity and significance of the

22 Middle Pleistocene Faunal Units (FU) for this region. In contrast with the previous model of a wide

23 faunal renewal during Marine Isotope Stage (MIS) 9 (~330 ka), the First Occurrences (FO) of

24 several species of the Torre in Pietra FU are significantly backdated and referred to the Fontana

25 Ranuccio FU (530-400 ka). We show that the faunal renewal was more gradual and occurred earlier

26 than previously assumed. Many taxa that are typical of the Late Pleistocene register their FO in the

27 Fontana Ranuccio FU, latest Galerian, which is characterized by the almost total disappearance of

28 Villafranchian taxa and by the persistence of typical Galerian taxa such as Dama clactoniana, Bison

29 schoetensacki and Ursus deningeri, and by the FO of Stephanorhinus kirchbergensis, S.

30 hemitoechus, Hippopotamus amphibius, Cervus elaphus eostepahnoceros, Ursus spelaeus, Canis

31 lupus, and Vulpes vulpes. The next Torre in Pietra FU is characterized only by the FO of

32 Megaloceros giganteus and of Mustela putorius. However, we observe that MIS 9 marks the actual

1 33 moment when the faunal assemblages of this region are represented only by those taxa

34 characterizing the late Middle Pleistocene and Late Pleistocene. For this reason, we propose to

35 consider the Torre in Pietra (lower levels) local fauna still as a conventional boundary for the

36 Galerian-Aurelian transition. Finally, we remark that the strong faunal renewal in MIS 13, with six

37FOs, coincides with the temperate climatic conditions due to the absence of marked glacial periods

38 that could have favored the FOs and the subsequent spread of these taxa.

40 keywords: Galerian; Aurelian; Mammal Age; biochronology; chronostratigraphy; Middle

41 Pleistocene; peninsular Italy

2 42 1. Introduction

43 Vertebrate paleontologists subdivide geological time using biochronological scales based on

44 the succession of evolutionary stages of faunal assemblages and dispersal events. For the Plio-

45 Pleistocene on the Italian peninsula, the first proposed Mammal Age was the Villafranchian

46 followed by the Galerian, and finally by the Aurelian proposed by Gliozzi et al. (1997, and

47 references therein). The division into Faunal Units (= FU's) of the Villafranchian Mammal Age

48 dates back to Azzaroli (1977). This publication was followed by the proposals for subdivision in

49 FU’s of the Galerian Mammal Age (Azzaroli, 1983) and finally, at the same time with his

50 institution, those of the Aurelian Mammal Age (Gliozzi et al., 1997).

51 Gliozzi et al. (1997) used the data available at that time (including those concerning small

52 mammals) to establish a biochronological scale valid for the Italian Peninsula, shared by all

53 researchers (including malacologists and ostracologists) who worked on the Plio- Pleistocene

54 continental deposits. The proposed biochronological scheme received general approval and has

55 been used by all Italian scholars. Over the past twenty years, new geological research, geochemical

56 methods, fossil discoveries, and systematic reviews of material already collected has led to a large

57 amount of new data. As a result, there have been a series of proposals for modifications or

58 integrations of the biochronological chart (for the entire period of time or for a part of it) with the

59 unification of various FUs, introduction of further FUs, and establishment of a new Mammal Age

60 (e.g. Petronio and Sardella, 1999; Palombo et al., 2004; Palombo, 2007, 2009; Palombo and

61 Sardella, 2007; Masini and Sala, 2007, 2011; Petronio et al., 2007, 2011; Sardella and Palombo,

62 2007; Bellucci et al., 2015). In the same period, works based on small mammals were also

63 published (Kotsakis et al., 2003; Sala and Masini, 2007). Finally, scales have been proposed, based

64 on the concept of "palaeocummunity" (PCOM) (Raia et al., 2006) as a non-alternative but

65 integrative method of the biochronological scale based on First Occurences (FO’s) / Last

66 Occurences (LO’s) of taxa, or on the concept of Faunal Complexes (FC) (Palombo, 2014).

3 67 Since the scheme of Gliozzi et al. (1997) continues to be a point of reference for many

68 researchers and because it has been used for comparisons with the faunas of various European

69 countries (e.g., Alberdi et al., 1998, Barishnikov, 2002, Palombo and Valli, 2004, Moigne et al.,

70 2006; Palombo et al., 2006; Kostopoulos et al., 2007; Nomade et al., 2014), it will be used

71 provisionally in this work, waiting for a new collective effort to update/modify/change it using all

72 new data.

73 The appearance of the taxa representing the core of the present day mammal fauna in the

74 European continent is a crucial biostratigraphic event of the Middle Pleistocene (Gliozzi et al.,

75 1997; Kahlke et al., 2011; Palombo, 2014). Until recently, such event was regarded as occurring

76 between 350 and 320 ka in peninsular Italy at the onset of Marine Isotope Stage (= MIS) 9, and

77 characterized, among the others, by the FO’s of several carnivorous species, including Panthera

78 spelaea, Ursus spelaeus, Canis lupus and Vulpes vulpes along with other carnivore and herbivore

79 taxa, constituted the Aurelian Mammal Age (Gliozzi et al., 1997; Petronio et al., 2011). In contrast,

80 an earlier appeareance for some of these species was proposed in continental Europe and Britain,

81 where the FO of Ursus spelaeus and Canis lupus are reported within MIS 11 (Schreve and

82 Bridgland, 2002; Brugal and Boudadi-Maligne, 2011; Palombo, 2014,). A contrasting picture is

83 achieved instead in the Transcaucasian region, on the European border, where Vulpes vulpes is

84 reported since MIS 14/13, but Canis lupus appears only since MIS 7 and Ursus deningeri occurs

85 until MIS 9 (Barishnikov, 2002).

86 A reconciliation of this contrasting pattern seems to emerge from a new chronostratigraphic

87 assessment of the sites of central Italy (Latium region), where the faunal assemblages constituting

88 the Aurelian Mammal Age were recovered (Fig.ure 1). The age review involved several

89 sedimentary sequences of the Tyrrhenian area of Latium, particularly along the Via Aurelia, near

90 Rome (Marra et al., 2018), previously assigned to the Aurelia Formation (Conato et al., 1980; Milli

91 et al., 2004; Palombo et al., 2004), correlated with MIS 9 (Conato et al., 1980; Karner and Marra,

4 92 1998). In fact, several of these deposits were re-allocated to the Valle Giulia Formation (MIS 13)

93 and to the San Paolo Formation (MIS 11) (Table 1).

94 This revision was carried out using a methodology introduced and widely described in a

95 series of recent publications (Marra et al., 2014a; 2015; 2016a; 2016b; 2017a; 2018; Pandolfi and

96 Marra, 2015; Ceruleo et al., 2016; Villa et al., 2016; Pereira et al., 2017a, 2017b; Petronio et al.,

97 2017), which uses the integration of:

98the direct dating with the 40Ar/39Ar method of volcanic levels intercalated in sedimentary

99 deposits;

100correlation with the MISs through identification of the aggradational successions deposited

101 as a consequence of the glacio-eustatic oscillations during the glacial terminations;

102correlation of sedimentary sequences with depositional and erosional paleosurfaces

103 generated by the interaction between glacio-eustatism and regional uplift.

104 In this work we present also a new 40Ar/39Ar dating of a pyroclastic layer closing the

105 volcanic succession cropping out along the Via Aurelia, marking the chronostratigraphic transition

106 between the deposits of MIS 13 and MIS 11 in this area, providing a further constraint in

107 biochronologic reconstructions.

108 The only sites in the area around Rome where the Aurelia Formation (MIS 9) was

109 recognized are those of Torre in Pietra (lower levels) and of Polledrara di Cecanibbio (Table 1). All

110 the others are older than previously assumed (MIS 11-MIS 13), while some are slightly younger

111 (MIS 8.5). The results of this chronostratigraphic revision require a careful re-examination of the

112 faunal content of many sites in central Italy, given their importance for the origin and distinction of

113 the Faunal Units (FU's) between Late Galerian and Early and Middle Aurelian (Gliozzi et al., 1997,

114 Petronio et al., 2011, Marra et al., 2014a). The different chronostratigraphic interpretation of these

115 continental deposits included between the FU's of Fontana Ranuccio, Torre in Pietra and Vitinia,

116 involves a necessary systematic and above all biochronological discussion on many mammal taxa

117 found in the area, and offers the opportunity to compare the Italian framework with the European 5 118 one, providing points for discussion on the evolutionary processes and migration paths of some

119 species, such as the cave bear, the wolf, the fox, the deer and the fallow deer.

120

121 2. Historical background and Paleontological setting

122 According to Gliozzi et al. (1997), the typical Galerian mammals, such as Equus altidens,

123 Equus gr. bressanus-suessenbornensis and the cervids of genus Megaceroides (= Praemegaceros),

124 already occur in the Late Villafranchian and the faunal renewal that produced the Galerian

125 assemblages was a gradual phenomenon. The Colle Curti local fauna is characterized by the FO of

126 Megaceroides verticornis (= Praemegaceros verticornis) and it was conventionally used to mark

127 the beginning of the Galerian Mammal Age (Gliozzi et al., 1997). Beyond Colle Curti FU the next

128 FU's of the Galerian Mammal Age are: Slivia FU, Ponte Galeria FU, Isernia FU and Fontana

129 Ranuccio FU (Gliozzi et al., 1997; Petronio and Sardella, 1999).

130 As mentioned above, the late Galerian is represented by Fontana Ranuccio FU and it was

131 defined as a period characterized by the total disappearance of typical Villafranchian taxa (with the

132 probable exception of Homotherium), the appearance of Hippopotamus amphibius and of the red

133 deer subspecies Cervus elaphus eostephanoceros, whereas Dama clactoniana, Equus altidens and

134 E. suessenbornensis are still present (Gliozzi et al., 1997). Recently, this FU was constrained within

135 a time interval spanning from MIS 13 to MIS 11 (Marra et al., 2014a). Furthermore, the occurrence

136 of Homotherium in the late Middle Pleistocene was confirmed (Bona and Sardella, 2014), and the

137 FO during MIS 13 of Stephanorhinus kirchbergensis and Stephanorhinus hemitoechus was

138 established (Pandolfi and Marra, 2015). Conversely, the LO’s of E. altidens and E.

139 suessenbornensis were re-collocated into the Isernia FU (Alberdi and Palombo, 2013). The most

140 representative mammal assemblages of the Fontana Ranuccio FU in the Rome area are Cava Nera

141 Molinario, Fontignano 2, Via della Pisana and Cava Rinaldi (Gliozzi et al., 1997; Marra et al.,

142 2014a).

6 143 Gliozzi et al. (1997) proposed the beginning of the Aurelian Mammal Age approximately in

144 correspondence with MIS 10/9, characterized by the appearance of the taxa representing the core of

145 the present day mammal fauna (early Aurelian or Torre in Pietra FU, MIS 10-9). In particular, the

146 faunal renewal in Italy was characterized by the disappearance of the cervid species Megaceros

147 savini (= Megaloceros savini), Megaceroides verticornis (= Praemegaceros verticornis), and the

148 archaic forms of red deer (Cervus elaphus acoronatus and C. elaphus eostephanoceros), which

149 were replaced by Megaloceros giganteus, along with the spread of the quasi endemic red deer

150 subspecies, Cervus elaphus rianensis, and by the FO of several carnivorous species, including

151 Ursus spelaeus,, Canis lupus and Panthera spelaea. Afterward, Petronio et al. (2011) added to the

152 above bioevents the FO of Mustela putorius, Vulpes vulpes and Felis silvestris in Central Italy.

153 The middle Aurelian faunal assemblages constituting the Vitinia FU were characterized by

154 the appearance of a subspecies of modern fallow deer Dama dama tiberina and of a small sized

155 equid with slender limbs, Equus hydruntinus (see Gliozzi et al., 1997; Di Stefano and Petronio,

156 1997; Petronio et al., 2011). Initially correlated with MIS 7 (Gliozzi et al., 1997; Palombo et al.,

157 2002; Petronio et al., 2011), its chronology was recently expanded to MIS 8.5 (Marra et al., 2014a).

158 In this time interval the first archaic Neanderthal-like humans also occur in Italy (Marra et al., 2015,

159 2017a).

160 As for the late Aurelian, which begins with the last interglacial (MIS 5e) and ends with the

161 end of the last glaciation, Gliozzi et al. (1997) did not propose any FU. The main reason was that,

162 from then onwards, because of a series of climatic events and the different microclimatic and

163 environmental conditions, no FUs can be considered as representative of the whole Italian territory.

164 Later, however, Petronio et al. (2007) proposed for this time interval the FUs of Melpignano and

165 Ingarano, representative of the traditional “warm” and “cold” mammal assemblages, respectively,

166 of Late Pleistocene.

167 Palombo et al. (2004) had proposed to merge the Vitinia FU with that of Torre in Pietra one,

168 and this due to the proximity in time of the dating of the two sites and the first appearance of the 7 169 archaic form of modern deer, Dama dama tiberina, at Sedia del Diavolo upper gravel, attributed by

170 these authors to MIS 9. However, Marra et al., (2014a, 2017a) have shown that the Sedia del

171 Diavolo upper gravel layer is geochronologically constrained within the younger MIS 8.5, and that

172 no occurrence of D. dama tiberina is reported before this period, remarking the biochronologic

173 distinction of the two FUs.

174

175 3. Material amd methods

176 3.1 Chronostratigraphic revisions

177 The recognition of the aggradational succession of the Valle Giulia (VG) and San Paolo

178 (SP) Formations at the sites cropping out in the vicinity of Via Aurelia has been achieved by Marra

179 et al. (2018) through the identification of two main tephrostratigraphic markers: the Tufo Giallo di

180 Prima Porta (TGPP) and the Sabatinian Tufo Rosso a Scorie Nere (TRSN) pyroclastic-flow

181 deposits, dated at 516±1 ka (Marra et al., 2017b) and at 449±2 ka (Karner et al., 2001), respectively

182 (Fig.ure 2(a)). For the present study we have added a further geochronology marker providing a

183 40Ar/39Ar age of 381±2 ka for the volcanoclastic deposits underlying a travertine horizon closing the

184 stratigraphic succession along via Aurelia, at km 16.6, supporting further the conclusion in Marra et

185 al. (2018) about the lack of any deposit atributable to the Aurelia Formation (MIS 9, ca. 330-300

186 ka) in the investigated area between Castel di Guido and Malagrotta (Fig.ure 2). Moreover, this new

187 younger geochronologic constraint allowed us at reconsidering the chronostratigraphic framework

188 of the volcanoclastic layers and of the deeply reworked massive pyroclastic-flow deposits

189 previously attributed to the Grottarossa Pyroclastic Sequence (GRPS, 510±4 ka) and to the TGPP

190 eruption cycles occurring at km 19.3 of Via Aurelia (Marra et al., 2018), suggesting a common

191 genesis for them within MIS 11. Direct 40Ar/39Ar dating of samples from Malagrotta and Via

192 Aurelia km 19.3 in progress within a research project by Boschian et al. will help to clarify this

193 issue.

194 8 195 3.2 Palaeontology

196 A review of the literature reporting descriptions, images, ostaeological data, as well as

197 regional syntheses on the major vertebrate deposits and fauna assemblages attributed to the

198 Aurelian Mammal Age (AMA) was performed. This review aims at evaluating the distribution of

199 different taxa. In many cases, i.e. Cava Rinaldi, Via Aurelia km 19.3, Collina Barbattini (in part),

200 Castel di Guido (in part), Malagrotta, Via Flaminia km 8.2, Riano, Sedia del Diavolo, Prati Fiscali,

201 Monte Sacro, Batteria Nomentana, Vitinia and Cerveteri-Migliorie San Paolo, the fossil remains

202 were re-examined by the authors (particularly CP, GDS, TK and LS), because they were stocked in

203 the Paleontology Museum Department, at Sapienza University in Rome.

204 The reviewed faunal lists for all the sections hosting the faunal assemblages previously

205 included in the FU's of the AMA are reported in Marra et al., (2018, table 2).

206

207 4. Results

208 4.1 40Ar/39Ar data

209 Sanidine phenocrysts were isolated from sample MG4 using standard magnetic and density

210 separation techniques, and were co-irradiated with the 1.1864 Ma Alder Creek sanidine standard

211 (Jicha et al., 2016; Rivera et al., 2013) at the Oregon State University TRIGA reactor in the

212 Cadmium-Lined In-Core Irradiation Tube. Single crystal fusion analyses were performed at the

213 WiscAr laboratory at the University of Wisconsin-Madison using a 60W CO2 laser and a Noblesse

214 multi-collector mass spectrometer following Jicha et al. (2016). Results are reported in Table 2.

215 Two crystal populations are present in sample MG4. An oldest population of eight crystals yield a

216 weighted mean age of 405±1 ka, and the youngest two crystals are 381±2 ka.

217 4.2 Reconstruction of the chrono-stratigraphic setting

218 In the investigated area, the stratigraphic setting reflects the occurrence of two consecutive

219 sea-level falls during the glacial periods corresponding to MIS 14 and MIS 12 (Fig.ure 2a-c). The

220 TGPP is emplaced above a marked paleomorphology and rests at the bottom of the fluvial incisions 9 221 eroded during the sea-level fall of MIS 13.2 (Marra et al., 2017b, Fig. 2 c). It is always associated

222 with the partially reworked deposit of the Grottarossa Pyroclastic Sequence (GRPS), which

223 emplaced 510±4 ka at the beginning of the successive sea-level rise culminating during MIS 13.1,

224 and responsible for the deposition ofwhen most of the sedimentary deposits of the VG Formation

225 cropping out along Via Aurelia were emplaced. The time of deposition of this aggradational

226 succession is bracketed by two more pyroclastic-fall deposits of the Fall A eruption cycle, that in

227 Cava Rinaldi have been dated at 498±2 and 496±9 ka (Fig.ure 2(a); Marra et al., 2014b; 2017b).

228 The pyroclastic-flow deposit of the Sabatinian TRSN also displays variable elevation along

229 Via Aurelia as a consequence of its emplacement during the late stages of the regressive phase

230 leading to the glacial maximum of MIS 12 (Fig.ure 2(c)). It is associated with an up to 4 m thick

231 volcanoclastic deposit, partly deriving from reworking of the TRSN and re-deposition within the

232 fluvial incisions and by an upper portion constituted of primary to partially reworked in sub-aerial

233 environment fallout deposits. The sedimentary deposits of the SP Formation in this sector

234 unconformably overly those of the Valle Giulia Formation and are scarce and represented by

235 diatomitic and travertinaceous clay horizons, emplaced within the higher portions of the fluvial

236 incisions during the late aggradational phase of MIS 11 (e.g., Castel di Guido, Riano, Fig.ure 2(a-

237 and b)). They are commonly associated with the fallout deposits of the first eruption period of the

238 Vico volcano (Vico , Cioni et al., 1987; Perini et al., 2004), dated 412±2 ka through 406±5 ka

239 (Marra et al., 2014b; 2018). The age of 381±2 ka provided here for the terminal portion of the

240 volcanic succession cropping out at km 16.6 of Via Aurelia (Fig.ure 2(a)) confirms the lack of any

241 sedimentary deposit of the Aurelia Formation above those of MIS 11 in this area. The dated sample

242 MG4 was collected in a massive pyroclastic deposit composed of loose crystals and lapilli,

243 originated by accumulation of fallout deposits on top of the morphological heights forming the

244 landscape during the MIS 11 sea-level highstand, as also inferred from its age. This sample yielded

245 two crystal populations, with a larger and oldest population of eight crystals yielding a weighted

246 mean age of 405±1 ka, and with two youngest crystals yielding 381±1 ka. The oldest age matches 10 247 that of the fallout deposit occurring at he top of the MIS 11 sedimentary deposits in Riano (Fig.ure

2482(b)), confirming the time span of 412-405 ka for the major eruptive activity in the late stages of

249 MIS 11, while the age of 381±2 ka suggests a slight reworking and mixing with the deposit of the

250 scanty volcanic activity characterizing the Late Southern Sabatini center in the time span 389-379

251 ka (San Abbondio Ashfall Succession, Marra et al., 2014b).

252 Therefore, in contrast with previous widespread attribution to the Aurelia (AU) Formation,

253 age of sample MG4 confirms conclusions by Marra et al. (2018) condensed in the cross-section of

254 Figure 2(a), showing that no sedimentary succession cropping out along the Via Aurelia between

255 Castel di Guido and Malagrotta correlates with MIS 9. However, the young age provided by sample

256 MG4 imposes to reconsider the K/Ar age of 350 ka reported in Cassoli et al. (1982) for a

257 volcanoclastic deposit at the top of the sedimentary succession cropping out in Malagrotta,

258 suggesting that at least part of it correlates with MIS 11.

259 Based on this renewed chrnostratigraphic pictureIn particular, the faunal assemblages

260 previously included in the Torre in Pietra FU occur:

261 above a paleosurface corresponding to a fluvial incision excavated during MIS 12 and sealed

262 by the Vico - fallout deposits (412±2 ka) and by the upper portion of the MIS 11

263 aggradational succession (ca. 400 ka, SP Formation) in Castel di Guido;

264 above a paleosurface interpreted to corresponding to with a fluvial incision excavated during

265 MIS 13.2 and partially filled by the pyroclastic-flow deposit of GRPS (510±4 ka) and,

266 successively, by the fluvial lacustrine deposits of the VG Formation (MIS 13) in

267 Via Aurelia at km 19.3, and at km 19 through 18.7 (Collina Barbattini) (Marra et al., 2018);

268 however, based on age of 381±2 ka yielded by sample MG4, we suggest that the

269 volcaniclastic deposits associated with such paleosurface may correlate those at km 16.6 of

270 Via Aurelia, and their lower elevation may be due to successive tectonic dislocation;

271 above and/or embedded in the upper portion of the pyroclastic-flow deposit of TGPP (516±1

272 ka), at the base of the deposits of the VG Formation in Malagrotta and in Cava Rinaldi. 11 273 Anzidei et al. (1993) proposed a chronology younger than 430 ka for the faunal assemblages

274 of Via Aurelia, basing on a now supersed K-Ar dating of the ignimbrite known as Tufo Rosso a

275 Scorie Nere (successively re-dated at 449±2 ka, Karner et al., 2001), whose reworked scoria were

276 erroneously identified within the deposit in which the fossils were recovered. However, Anzidei et

277 al. (1993) clearly report that all the archaeological material recovered at these four sites between km

278 17.9 and 19.3 in Via Aurelia was found within a deposit overlying the palaeosurface described as a

279 palaeomorphology originated by turbulent flux, excavated in the underlying fluvial-lacustrine

280 succession. Following the chronostratigraphic revision by Marra et al . (2018), the pyroclastic-flow

281 deposit of the GRPS (510±4 ka) is emplaced above this palaeosurface and the fluvial-lacustrine

282 succession, occurring below and above it, is the MIS 13 aggradational succession (VG Formation)

283 (Fig. 4). Therefore, fossils and lithic materials are backdated to ca. 510 ka. Remarkably, the GRPS

284 pyroclastic-flow deposit has a juvenile fraction composed of blackish, poorly vesicular scoria clasts

285 (Karner et al., 2001), a fact that may be at the base of the previous incorrect chronostratigraphic

286 interpretation, which was based on mere observational criteria and lacked of any objective element

287 (e.g., chemistry, mineralogical assemblage, etc.), leading to a ca. 200 ky chronologic misplacement

288 for the lithic industry and the faunal assemblage recovered at this site.

289 It is worth noting that the outcrop where the Malagrotta faunal assemblasge attributed to the

290 Torre in Pietra FU by Caloi and Palombo (1980) occurred (MG2 in Fig. 2) is a different one with

291 respect to that described by Cassoli et al. (1982) (MG1 in Fig. 2) where an acheulian lithic

292 assemblage was found. The pyroclastic-flow deposit cropping out in MG2 displays primary

293 features, apart from being reworked in its uppermost portion, allowing corelation with the TGPP

294 eruption unit; in contrast, the basal horizon of the sedimentary succession cropping out in MG1

295 includes a massive, deeply reworked volcanic layer, laterally passing to a sand deposit with sparse

296 gravel. Also in light of the age of 381±2 ka yielded by sample MG4, we suggest that previous

297 correlation with TGPP of the basal volcanic layer in MG1 proposed in Marra et al. (2018) requires

298 further verification, and an age within MIS 11 for the entire sedimentary succession cropping out at 12 299 this location cannot be excluded, a fact that will be clarified by dating of samples of the volcanic

300 deposits occurring in Malagrotta in progress within a study bi Boschian et al.

301 In Malagrotta, Capasso Barbato and Minieri (1987) have classified as Vulpes vulpes the

302 remains recovered in the travertine layers at the top of the sedimentary deposits. Unfortunately, no

303 precise stratigraphic correlation with respect to the stratigraphy of Malagrotta described by Palma di

304 Cesnola (2001) was provided, and Marra et al. (2018) have tentatively correlated the deposit with

305 the uppermost layer reported in this latter stratigraphic section of the site, attributing it to MIS 11.

306Age provided by the MG4 sample supports this correlation.

307 However, they have also remarked that other travertine layers occur in lower stratigraphic

308 position within the MIS 13 deposits of the VG Formation, leaving the exact age assessment of the

309 fox remains an open question. For the present study we have compared the stratigraphy reported by

310 Capasso Barbato and Minieri (1987) with that observed at different outcrops in the area of

311 Malagrotta. Consistent with description by these authors, a 1-2 m-thick horizon of travertine

312 deposits occurs in the upper portion of the Valle Giulia Formation cropping out throughout

313 Malagrotta, at elevation around 55 m a.s.l. (Figure 2a). This layer corresponds to an indurated

314 stratum (as inferred by its protruding feature despite no corresponding lithology is reported)

315 occurring in the middle of the stratigraphic succession of Malagrotta figured by Palma di Cesnola

316 (2001). In contrast, as already remarked by Marra et al. (2018), the whole upper portion of the

317 Malagrotta stratigraphy reported in Palma di Cesnola (2001) does not occur in any of the geologic

318 sections presently exposed in this area. However, part of it, including a ca. 4 m thick volcanoclastic

319 layer (identified as the "leucititic tuff" reported by Palma di Censola, 2001) and an overlaying, ca. 1

320 m-thick travertine layer, is observed at km 16.6 in Via Aurelia (Figure 2a). 40Ar/39Ar dating of

321 sample MG4 collected in the volcaniclastic layer at this location yielded age of 381±2 ka,

322 confirming correlation with late MIS 11 (Figure 2a). We conclude that the stratigraphy in Palma di

323 Cesnola (2001) is a composite cross-section, representing an ideal, merged stratigraphic setting of

324 the larger area of Malagrotta, comprehensive of both aggradational succession of MIS 13 and MIS 13 325 11, whereas the succession really cropping out at the archaeological site is only that described in

326 Capasso Barbato and Minieri (1987), which matches very well that described at the nearby site

327 named Malagrotta in Marra et al. (2017b), and reported in Marra et al. (2018). These sedimentary

328 deposits correspond only to the lower, oldest aggradational succession of the VG Formation;

329 therefore, the travertine layers hosting V. vulpes remains are part of the MIS 13 deposits dating 516-

330 500 ka.

331 The only deposits correlated with the AU Formation and MIS 9 in the whole region are

332 those cropping out in Torre in Pietra (lower levels) and in Polledrara di Cecanibbio.

333 In the first locality, the aggradational succession of the AU Formation has been bracketed by

334 two 40Ar/39Ar ages of 354±5 and 337±6 ka (Villa et al., 2016). The local fauna occurs within the

335 basal portion of the deposit, constituted by fluvial gravel and sand, where the oldest sample for

336 radiometric dating was collected.

337 In Polledrara di Cecanibbio, only a ca. 50 cm thick horizon of cross-laminated sand with

338 reworked volcanic material that yielded a 40Ar/39Ar age of 359±6 ka (Pereira et al., 2017a)

339 represents the sedimentary record correlated with the AU Formation and MIS 10/9 (Fig.ure 2(a)).

340 However, for the most part the local fauna occurs above this sedimentary layer, where it was

341 accumulated by fluvial transport, and it was later partially re-mobilized and embedded within a sub-

342 primary, volcanic mud-flow deposit (lahar), which has been dated at 325±2 ka and correlated with

343 the Tufo di Bracciano eruption (Pereira et al., 2017a). Moreover, three elephant skeletons have been

344 found in anatomical connection at the top of the mudflow, in which the pachyderms evidently were

345 bogged down by trampling above the only partially indurated mudflow, after its deposition.

346 In Riano, the eponymous locality of the subspecies Cervus elaphus rianensis which was

347 considered representative of the Aurelian Mammal Age (Gliozzi et al., 1997; Palombo, 2004;

348 Petronio et al. 2011), a pyroclastic-fall deposit intercalated in the upper portion of the lacustrine

349 succession in which the cervid remains were recovered, yielded age of 406±5 ka, evidencing

350 correlation with MIS 11 (Fig.ure 2(b)). 14 351 Similarly, the sedimentary deposits cropping out at km 8.0-8.2 of Via Flaminia, hosting C.

352 elaphus ssp. and previously attributed to MIS 7 (Palombo, 2004), have been identified as the basal

353 portion of the VG Formation, deposited during the early sea-level rise of MIS 13.3, as evidenced by

354 tephrostratigraphic constraints provided by Tufo del Palatino (533±2 ka, Marra et al., 2017b) and by

355 TRSN (449±2 ka) (Fig.ure 2(b)).

356 The chronostratigraphic framework of the sites located along the Aniene River Valley north

357 of Rome, and correlated with the Via Mascagni (VM) Succession and the Vitinia (VI) Formation,

358 deposited during sea-level rises associated with MIS 8.5 and MIS 7, respectively, has been

359 described in detail in Marra et al. (2017a). We direct the readers to the geologic section in Figure 10

360 of Marra et al. (2017a) for the chronostratigraphic constraints providing correlation with MIS 8.5

361 for Sedia del Diavolo and Monte delle Gioie, and with MIS 7 and the Vitinia geologic type-section

362 (Karner and Marra, 1998) for Casal de' Pazzi and Saccopastore.

363 Moreover, Marra et al. (2018) have correlated also the deposit of Prati Fiscal with the

364 sedimentary succession cropping out in Monte delle Gioie, framing their age within MIS 8.5.

365 The occurrence of Tufo Giallo di Sacrofano (Karner et al., 2001) at the top of the

366 sedimentary deposits of the Via Mascagni Succession allows to constrain the age of the MIS 8.5

367 sites between 285±2 ka and 295±5 ka (Marra et al., 2017a). The taphonomy of these sites, along

368 with an up-to-date technological analysis of the lithic industry recovered at Monte delle Gioie and

369 Sedia del Diavolo has been recently provided by Soriano and Villa (2017). Remarkably, these lithic

370 industries, along with the human remains recovered at Sedia del Diavolo and Ponte Mammolo, have

371 been recognized as the earliest evidence of Neanderthals in Italy (Marra et al., 2017a).

372 An age spanning 270-250 ka has been proposed for Casal de' Pazzi, based on correlation

373 with the early aggradational stages during MIS 7.5 combined with the post-quem age of 304±9 ka

374 provided by reworked pyroclastic material intercalated within the sedimentary deposit (Marra et al.,

375 2018).

15 376 An age spanning 245 - 220 ka, possibly corresponding to the two consecutive sea-level rises

377 during MIS 7.5 and MIS 7.3, has been suggested for the sedimentary deposits of Saccopastore

378 hosting two Neanderthal skulls, consistent with the identification of Dama dama tiberina remains

379 among the fossil remains from this locality hosted at Museo Pigorini in Rome (Marra et al., 2017a).

380 Finally, the upper levels of Torre in Pietra have been dated at 270-240 ka by Villa et al.

381 (2016), through the combined upper age constraint of a tephra yielding 208±2 ka, and the

382 correlation with the earliest sea-level rise during the complex pattern of eustatic oscillations at the

383 onset of Glacial Termination III (Marra et al., 2016c), providing attribution to MIS 7.5.

384 Remarkably, all the findings of D. dama tiberina and E. hydruntinus so far reported in the

385 literature occur at the sections correlated with MIS 8.5 and MIS 7 (Sedia del Diavolo, Batteria

386 Nomentana, Vitinia, Torre in Pietra upper levels, Saccopastore) (Marra et al., 2017a, 2018),

387 whereas these taxa are not present in the two MIS 9 locations of Torre in Pietra lower levels and

388 Polledrara di Cecanibbio. Moreover, in the only site so far referred to MIS 5, Fosso del Cupo

389 (Ceruleo et al., 2016), the presence of D. dama dama has been reported.

390 4.3 Biocronological review

391 The results of the illustrated chronostratigraphic review have made it necessary to review

392 and, in many cases, update the taxonomic and / or biochronological position of several taxa

393 previously considered typical of the Aurelian Mammal Age (Gliozzi et al., 1997; Kotsakis et al.,

394 2003; Petronio et al., 2011). The posed problems concern for small mammals the systematic review

395 of some fossils belonging to two taxa reported in the deposits of the latest Galerian and / or early

396 Aurelian. For large mammals instead, moving backwards in time of their FO.

397 1. Crocuidura cf. C. suaveolens (Pallas, 1811) reported by Anzidei et al. (1993) in the Via Aurelia

398 18.9 km site.

399 2. Arvicola terrestris (Linnaeus. 1758) (= A.amphibius (Linnaeus, 1758)) reported by Anzidei et al.

400 (1993) in two sites of Via Aurelia (Collina Barbattini, around km18.7, and at km 18.9); Caloi and

401 Palombo (1978) report Arvicola terrestris-amphibius group (Arvicola sp. in Marra et al., 2018). 16 402 3. Ursus spelaeus (Rosenmüller, 1794) found in Cava Rinaldi (Ponte Galeria, Rome) (Capasso

403 Barbato and Minieri, 1987), in Torre in Pietra, lower and upper levels (Caloi and Palombo, 1978)

404 and perhaps in Collina Barbattini (Ursus cf. U. spelaeus, Perrone, 2016);

405 4. Canis lupus Linnaeus, 1758, whose remains were found at Castel di Guido (Radmilli et al., 1979;

406 Capasso Barbato and Minieri, 1987; Sala and Barbi, 1996), at Polledrara di Cecanibbio (Anzidei et

407 al., 2012) and Torre in Pietra, lower and upper levels (Caloi and Palombo, 1978); reported also from

408 Malagrotta by Caloi and Palombo (1980) (Canis sp. in Marra et al., 2018);

409 5. Vulpes vulpes (Linnaeus, 1758), the remains of which have been found in the upper travertines of

410 Malagrotta (Capasso Barbato and Minieri, 1987), the Polledrara of Cecanibbio (Anzidei et al.,

411 2012) and Torre in Pietra, lower and upper levels (Caloi and Palombo, 1978);

412 6. Panthera spelaea (Goldfuss, 1810), whose remains were found in the sites of Castel di Guido

413 (Radmilli et al., 1979; Capasso Barbato and Minieri, 1987; Sala and Barbi, 1996), Torre in Pietra,

414 lower and upper levels (Caloi and Palombo, 1978) and perhaps at Collina Barbattini (Panthera cf.

415 P. spelaea, Perrone, 2016);

416 7. The archaic subspecies of D. dama (Linnaeus, 1758) and C. elaphus Linnaeus, 1758 (see Di

417 Stefano and Petronio, 1993, 1997, 2002; Marra et al., 2014a).

418

419 Crocidura suaveolens (Pallas, 1811)

420 The presence of this species is tentativelydubitatively reported from the site at Via Aurelia

421 km 18.9 (Anzidei et al., 1993, as Crocidura cf. C. suaveolens). In a recent phylogeographic study,

422 Castiglia et al. (2017) indicate a time interval between 60 and 149 ka as cloning period for the

423 populations of the Italian peninsula. This data fits well with the data from studies on Italian fossil

424 cCrocidurini (Kotsakis et al., 2003). As a result, the Via Aurelia km 18.9 soricid should be revised.

425 Unfortunately, a review of the fossil was not possible.

426

427 Arvicola Lacépède, 1799 17 428 Small vertebrates of early Toringian age, a Mammal age that in Italy includes the FU’s of

429 Isernia, Fontana Ranuccio, Torre in Pietra and Vitinia (Gliozzi et al., 1997; Kotsakis et al., 2003;

430 Sala and Masini, 2007), are rather rare in the Latium area and almost all are represented by very few

431 (even one) specimens: Casal Selce (Ponte Galeria, Rome), Monte Li Pozzi (Cerveteri, Rome),

432 Cretone (Sabina, Rome), Via Aurelia km 18.7 (Collina Barbattini, Rome) and km 18.9, Polledrara

433 di Cecanibbio (Rome), Torre in Pietra upper levels (Rome) and Vitinia (Rome). In all these

434 fossiliferous sites the only genus of rodent constantlyalways present is the water vole, Arvicola.

435 The specific and nomenclatorialtaxonomic status of the extant species of the genus Arvicola

436 has undergone significant changes over the last twenty years. On the basis of the Priority rule of the

437 ICZN Arvicola amphibius is now used to indicate the European water vole instead of the previously

438 used Arvicola terrestri (Musser and Carleton, 2005 with references). Masini et al. (2003) suggested

439 that the extant and fossil water voles of the Late Pleistocene of the peninsular Italy, could represent

440 a different evolutionary lineage. This hypothesis, highlighted in many biological publications, was

441 confirmed (analysis of mtDNA) by Castiglione et al. (2016) who revived the name Arvicola italicus

442 (Savi, 1838) at speciesfic level.

443 The appearance of the genus Arvicola in Central Europe was assumed at the beginning of

444 the Toringian age of small mammals (Fejfar and Heinrich, 1983) (for the Iberian pre-Jaramillo age

445 species Arvicola jacobaeus Cuenca-Bescós, Agustí, Melero Rubio and Rofes, 2010 see discussion

446 in Lozano-Fernández et al., 2013). In addition, the transition from the primitive species Arvicola

447 mosbachensis (Schmidtgen, 1911) (= Arvicola cantianus (Hinton, 1910)) (see discussion about the

448 use of the two species names in Maul et al., 2000) to the extant form A. amphibius marks the end of

449 the Early Toringian and the beginning of the Late Toringian. The distinction between these two

450 species is based on Heinrich's SDQ index (Schmeltzbanddifferenziation-quotient = Enamel

451 differentiation ratio) (Heinrich, 1978). SDQ (used also for distinction between the genera Mimomys

452 and Arvicola, see Koenigswald and Kolfschoten, 1996) is calculated by measuring the enamel

453 thickness on both sides of the salient angles of m1 and dividing the thickness of the trailing edge x 18 454 100 by the thickness of the leading edge. According to Maul et al. (1998), the comparison of the

455 SDQ values of the Arvicola populations of Central Europe and those of southern Italy, given the age

456 of the various fossiliferous sites, highlights a considerable heterochrony.

457 In Italy the genus Arvicola occurs for the first time in the local fauna of Isernia La Pineta

458 (Coltorti et al., 1982, 2005), typical of the Isernia FU, with the species A. mosbachensis (= A.

459 cantianus). In Latium, belongs to the same FU the local fauna of Casal Selce is characterized by the

460 presence of A. mosbachensis (cited as Ponte Galeria 3 in Kotsakis and Barisone, 2008 who give a

461 preliminary list of species; the fauna is in study by Sardella et al.). At a younger stage (Monte Li

462 Pozzi, Fontana Ranuccio FU) belongs the assemblage of Monte Li Pozzi in which the only rodent is

463 A. mosbachensis (see Mancini et al., 2006). A third reportstatement of A. mosbachensis comes from

464 Cretone (Di Canzio et al., 2003) in an assemblage referred to the Torre in Pietra FU.

465 Anzidei et al. (1993) report water vole's remains assigned to A. terrestris (= A.amphibius )

466 from two sites along Via Aurelia, the first one at km 18.9 and the second one at km 18.7 in Collina

467 Barbattini. Maul et al. (1998) have shown that populations similar to the extant ones appear in

468 peninsular Italy only during the Late Pleistocene.

469 López-García et al. (2017) in a publication on the small mammals of the Grotta Maggiore di

470 San Bernardino (Vicenza, Veneto) indicate the presence of A. mosbachensis in the layers attributed

471 to MIS 7 and the appearance of the extant species A. amphibius (= ? A. italicus) in the layer

472 attributed to the transition MIS 6 / MIS 5.

473 We have not been able to examine the material mentioned by Anzidei et al. (1993) but we

474 can reasonably assume that, given the new age of the fossiliferous sites (Marra et al., 2018) and the

475 composition of the faunal assemblages, these remains belong to A. mosbachensis.

476 Arvicola remains collected at Polledrara di Cecanibbio (Anzidei et al., 2004) did not allow

477 classification at a speciesfic level. In this site we observe the LO of the rhizodont arvicolid Pliomys

478 episcopalis Mehély, 1914 in Italy.

19 479 The remains collected in the upper levels of Torre in Pietra and Vitinia (Caloi et al. 1998),

480 and initially attributed to A. gr. amphibius-terrestris and Arvicola sp. respectively – (Caloi and

481 Palombo, 1978; Caloi et al., 1983; Kotsakis et al., 2003), and later reported as Arvicola sp. in

482 Kotsakis & Barisone (2008) should probably belong also to A. mosbachensis.

483 Obviously, a review of the remains of the late Middle Pleistocene water voles assigned to A.

484 terrestris is necessary, while attribution of such remains to A. mosbachensis can be considered as a

485 working hypotheses. Based on all the considerations above, the attribution of some fossil

486 assemblages to the Late Toringian (Kotsakis et al. 2003) should be re-examined. For the moment, it

487 is suggested to assigne these remains to Arvicola sp.

488

489 Ursus spelaeus Rosenmüller 1794

490 The faunal assemblage of Cava Rinaldi (Ponte Galeria, Rome) (Ambrosetti, 1965) is

491 characterized by the presence of a lower canine of Ursus sp. together with remains of Castor fiber

492 Linnaeus, 1758, Bos primigenius Bojanus, 1827 (prevalent), Elephas antiquus (= Palaeoloxodon

493 antiquus (Falconer 1857)) and Cervus cf. C. elaphus.

494 Capasso Barbato and Minieri (1987) classified a second upper molar coming from the same

495 site as Ursus spelaeus. The molar was found in the basal part of the tuff overlying the Ponte Galeria

496 Formation (Capasso Barbato and Minieri, 1987) which in the updated chronostratigraphy of Cava

497 Rinaldi (Marra et al., 2018) corresponds to the Tufo Giallo di Prima Porta (TGPP), occurring at the

498 base of the sedimentary deposits of the Valle Giulia Formation in this location, dated at 516±1 ka

499 (MIS 13; Marra et al., 2017b).

500 Cranial morphological and morphometric differences within Ursus deningeri von

501 Reichenau, 1906 and U. spelaeus were recently proposed by Santos et al. (2017), but they are useful

502 in the case of complete or sub-entire skulls only. The specific distinction between U. spelaeus the

503 cave bear and U.rsus deningeri von Reichenau, 1906 based only on the morphological

504 characteristics of the teeth is not simple and often unsafe (Mazza and Rustioni, 1992, 1994), 20 505 considering the morphological similarity between two taxa which gradually pass during the middle

506 Galerian from a more carnivorous diet (U. deningeri) to a more omnivorous one (U. spelaeus).

507 However, the second upper molar from Cava Rinaldi shows numerous accessory tubercles, usually

508 absent in Ursus arctos Linnaeus, 1758, and a strong cingulum at the base of the protocone, usually

509 absent in U. deningeri (see Bonifay, 1971; Torres, 1984; Capasso Barbato and Minieri, 1987;

510 Capasso Barbato et al., 1990)Considering the wide morphological and dimensional overlap field it

511 is therefore easier to distinguish the two species only in cases of molars testifying the most evident

512 carnivorous and / or omnivorous diet. The latter morphology is evident in U. spelaeus in teeth with

513 a very wide masticatory surface and with a greater number of tubercles. Despite the difficulty of

514 classification only through these morphologies, to a careful examination, the presence of numerous

515 tubercles and the typical dimensions of the cave bear (length-width ratio) (Capasso Barbato et al.,

516 1990), allow us to accept the attribution of the molar of Cava Rinaldi to U. spelaeus. Furthermore,

517 tThe absolute length and width data of the Cava Rinaldi bear molar and the values related to the

518 dispersion between the length of the molar and the width / length ratio both fall into the full range

519 of U. spelaeus (see Capasso Barbato and Minieri, 1987). Dimensional comparison with the

520 abundant cave bear remains molars of Grotta del Cerè (Veneto) Italy, continental Europe, Russia

521 and Caucasus (Capasso Barbato and Minieri, 1987; Capasso Barbato et al., 1990; Rossi and Santi,

522 2001; Baryshnikov et al., 2003; Baryshnikov, 2006) further reinforce thise attribution.

523 This datum obviously updates the biochronological position of this taxon which, according

524 to Gliozzi et al. (1997) and Petronio et al. (2011), previously characterized the first part of the

525 Aurelian, given that its FO was recorded in some local faunas attributed to Torre in Pietra FU (MIS

526 10-9). The shift back in time of the FO of U. spelaeus within MIS 13 complicates the possible

527 phylogenetic picture of this species that is considered here morphologically similar to U. deningeri

528 and probably derived from this species (Petronio et al., 2003; Baryshnikov, 2006; Argant, 2009),

529 although many authors (Mazza and Rustioni, 1992; 1994; Rossi and Santi, 2001) do not agree with

530 this phylogenetic framework. U. deningeri makes its first appearanceFO in Italy during the Slivia 21 531 FU (Early Galerian) (Gliozzi et al., 1997; Petronio et al., 2011), while its last occurenceLO in

532 Latium is reported in the Fontana Ranuccio local fauna (Bidittu et al., 1979; Segre Naldini et al.,

533 2009; Petronio et al., 2011), dated 408±10 ka (Pereira et al., 2017b). The phylogenetic sequence

534 that links the Villafranchian chronospecies Ursus etruscus Cuvier, 1823 to the early and middle

535 Galerian U. deningeri and U. spelaeus, would thus be articulated in a different way by the

536 backdating of the FO of the last taxon within MIS 13. The relationship between U. deningeri and U.

537 spelaeus, also through intermediate forms, such as U. spelaeus deningeroides Mottle, 1947, is well

538 established both anatomically (e.g., Rabeder, 1999; Baryshnikov, 2006; Argant, 2009) and

539 genetically (e.g., Orlando et al., 2002; Hofreiter et al., 2002; Dabney et al., 2013). However, there is

540 no agreement on the divergence chronology, as different methodologies provide different

541 divergence ages, so much so that some authors (e.g., Grandal-d'Anglade and López-González,

542 2004) argue that they could even be grouped into one species. A coexistence between the two

543 species should It could be hypothesized. fFor the Italian peninsula, that U. spelaeus would be is the

544 result of a new dispersion of a bear populationsmore specialized towards omnivory, derived from

545 with a process of speciation from U. deningeri, likely occurred elsewhere (central orin the eastern

546 European? or western Asian? territories). Moreover, U. deningeri is still reported in MIS 9 in the

547 Caucasian territories (Baryshnikov, 2002, 2006) while U. spelaeus is reported in the late Middle

548 Pleistocene (MIS 11) in Germany, in Bilzingsleben 2, and in Great Britain, in Swanscombe

549 (Schreve and Bridgland, 2002).

550 In Italy, U. deningeri seems to become extinct after MIS 11, and U. spelaeus spreads in the

551 Italian peninsula with fairly conservative forms compared to the European populations (Rossi and

552 Santi, 2013), likely due to a consistent with the partial isolation typical of many mammal taxa

553 penetrated with difficulty in the peninsula through the Alpine passes, and therefore partially and

554 temporarily isolated.

555 Together with U. spelaeus, the modern form of brown bear, U. arctos Linnaeus, 1758 , a

556 species probably derived from Asian archaic forms, spreaded out in Great Britain (Purfleet) and in 22 557 Europe during MIS 9 (Schreve and Bridgland, 2002) and become common only in the Late

558 Pleistocene. The FO of this species in Italy it is probably in Bucine (Tuscany), generally referred to

559 the early-middle Aurelian (Mazza, 1997; Palombo et al., 2002; Petronio et al., 2011) and, in an

560 insular environment, in Sicily (Bonfiglio et al., 2001) around MIS 6/7 (Palaeoloxodon mnaidriensis

561 Faunal Complex). In fact, in a work of regional synthesis, the late Galerian bear fossils from

562 Fontana Ranuccio was recorded as U. arctos by Azzaroli (1983), followed inter alios by Gliozzi et

563 al. (1997), Sardella et al. (2006), Palombo et al. (2009), Palombo (2014)., but theseInstead, in all the

564 works that described the Fontana Ranuccio local fauna, the bear finds were always attributed to U.

565 deningeri (see e.g., Biddittu et al., 1979; Ascenzi et al., 1993; Palombo et al., 2002; Segre Naldini et

566 al., 2009).

567

568 Canis lupus Linnaeus, 1758

569 The probable origin of Canis lupus is placed in the early Middle Pleistocene of Beringia (Far

570 East Asia – North western America) and its dispersal into Europe occurred in the late Middle

571 Pleistocene (Brugal and Boudadi-Malign, 2011; Flower and Schreve, 2014; Salari et al., 2017). The

572 phylogenetic sequence that connect the Villafranchian Canis etruscus Major, 1877 with the

573 Galerian Canis mosbachensis Soergel, 1925 and the modern C. lupus is widely accepted in the

574 literature, although their relationships and taxonomy are debated (e.g., Sotnikova, 2001; Brugal and

575 Boudadi-Maligne, 2011; Petrucci et al., 2013; Flower and Schreve, 2014; Sardella et al., 2014).

576 Previously, the FO of C. lupus in Italy was reported in the C. lupus was reported in the past

577 for the first time in Italy during MIS 10/9, Torre in Pietra FU (Gliozzi et al., 1997; Petronio et al.,

578 2011). Among the fossil deposits of Via Aurelia, C. lupus is present at Castel di Guido (Radmilli et

579 al., 1979; Capasso Barbato and Minieri, 1987; Sala and Barbi, 1996; Boschian and Saccà, 2010), at

580 Polledrara di Cecanibbio (Anzidei et al., 2012), and at Torre in Pietra, lower and upper levels (Caloi

581 and Palombo, 1978). Furthermore, Caloi and Palombo (1980) have attributed to thise species a

582 small fragment of a carnassial of modest size recovered in the field of Malagrotta (now attributed to 23 583 MIS 13) which, however, in our opinion, is unclassifiable at a speciesfic level. Morphology and

584 dimensions of the lower carnassial from Castel di Guido (Sala and Barbi, 1996) fall within the

585 variability range of the extant Apennine wolf, C. lupus italicus Altobello 1821 (Salari et al., 2017),

586 while the upper canine, the third lower premolar ant the metacarpus V are close to those of the

587 small-sized wolf from Lunel Viel, C. lupus lunellensis Bonifay, 1971 (Capasso Barbato and

588 Minieri, 1987; Boudadi-Maligne, 2012; Salari et al., 2017).

589 According to the new chronostratigraphic data (Marra et al., 2018) for Castel di Guido, the

590 FO of this species must be placed within MIS 11 (Fontana Ranuccio FU). This first presence

591 coincides with other Western European data, such as Lunel Viel (France), MIS 11/10, and

592 Atapuerca TD 10 (Spain), MIS 11 (Cuenca Bescós and García, 2007, Brugal and Boudadi-Maligne,

593 2011; Salari et al., 2017). In contrast, in the Caucasian area, on the other hand, C. mosbachensis is

594 present at least until the MIS 9, while the modern wolf appears only in the MIS 7 (Baryshnikov,

595 2002).

596

597 Vulpes vulpes (Linnaeus, 1758)

598 The genus Vulpes appears in Italy for the first time in the middle Villafranchian (Costa S.

599 Giacomo FU) (Petronio et al., 2006, 2011) with the species Vulpes alopecoides (Major, 1877),

600 whose fossil remains were found near Anagni (Latium) (Biddittu et al., 1984; Bellucci et al., 2014).

601 In the late Villafranchian, the species is reported in the deposits of the Upper Valdarno, in the site of

602 Monte Riccio (Tarquinia, Latium) (UF Tasso) and in the Gargano peninsula (Apulia) (Pirro UF)

603 and is still present in some north-eastern Italian sites, including Cava Nord and Viatelle del Monte

604 Tondo, Soave (Veneto), assigned to Middle Pleistocene (Bon et al. 1991; Petronio et al. 2006, 2011;

605 Petrucci et al., 2013).

606 Phylogenetic relationships between this species and Vulpes praeglacialis (Kormos, 1932)

607 are unclear, as well as the relationship between them,with V. vulpes, and Alopex lagopus (Linnaeus,

608 1758) are unclear (see Petrucci et al., 2013 with references). 24 609 Remains of Vulpes cf. V. vulpes are reported in the site of Visogliano (TS) (Abbazzi et al.,

610 2000; Masini and Sala, 2007), in a Middle Pleistocene fauna considered by Sardella et al. (2006) as

611 transitional to Fontana Ranuccio FU. Other remains attributed with certainty to this species come

612 from the site of Bristie 1 (Venezia Giulia), referring correlated to the final part of Galerian or the

613 beginning of Aurelian (Lugli and Sala, 2000; Petronio et al., 2006). It has been recently highlighted

614 how Fontana Ranuccio FU includes MIS 13 and MIS 11 (Marra et al., 2014a), so the faunas found

615 in the sites mentioned above can have any age between 530 and 410 ka. Consistently, Falguères et

616 al. (2010) have estimated through ESR dating an age ranging from 480 to 440 ka for the lower

617 levels of Visogliano, corresponding to the late MIS 12 (Figs. 2c and 3). The presence of remains of

618 V. vulpes reported in the travertines of the Malagrotta site (Capasso Barbato and Minieri, 1987),

619 which are correlated with MIS 11, provides an opportunity to establish the FO of this species in

620 Italy at the MIS 12-MIS 11 transition.

621 The presence of remains of V. vulpes reported in the travertines of the Malagrotta site

622 (Capasso Barbato and Minieri, 1987), provides an opportunity to establish an absolute age for the

623 FO of this species in Italy. The comparative review of the literature on faunas and outcrops made in

624 the present work, allows to establish the stratigraphic position inside the VG Formation deposits

625 and to establish the FO for this species in Italy in MIS 13, between 510 and 500 ka.

626 In any case, it should be noted that the presence of the fox in Italy is still more recent than in

627 other regions of Western Europe. For example, in France the FO of V. vulpes is reported in the

628 mammalian standard zones MNQ 22 (Mammifères Néogènes et Quaternaires 22) (Palombo and

629 Valli, 2004) correlated with Isernia FU (Palombo and Valli, 2004, Nomade et al., 2014), slightly

630 older than the first reports in Caucasus (Azykh Cave, MIS 14/13) (Baryshnikov, 2002), instead

631 substantially coeval with that of Malagrotta.

632

633 Panthera spelaea (Goldfuss, 1810) and Felis silvestris Schreber, 1775

25 634 Panthera spelaea, remains from Castel di Guido (Radmilli et al., 1979; Capasso Barbato and

635 Minieri, 1987; Sala and Barbi, 1996) as well as those referred to Panthera cf. P. spelaea from

636 Collina Barbattini (Perrone, 2016), and assigned by Marra et al. (2018) to MIS 13 and MIS 11, have

637 to be reviewed considering their possible attribution allocation to Panthera fossilis (von Reichenau,

638 1906).

639 This lion-like pantherine felid occurs in Western Siberia since the late Early Pleistocene

640 (Sotnikova and Foronova, 2014) and in Europe during Middle Pleistocene, from MIS 19-17 to MIS

641 9-7 (Sabol, 2011; Marciszak et al., 2014; Sotnikova and Foronova, 2014). In Italy, P. fossilis occurs

642 in the local fauna of Isernia La Pineta, referred to the homonymous FU (MIS 15) (Gliozzi et al.,

643 1997; Petronio et al., 2011; Marra et al., 2014a), and in a few localities attributed generically to the

644 Middle Pleistocene (Petronio et al., 2011; Bona and Sardella, 2012). Instead, P. spelaea remains are

645 relatively common in the Late Pleistocene mammal assemblages (Capasso Barbato and Gliozzi,

646 1994; Bona, 2006; Petronio et al., 2011).

647 Also among felids, the probable presence of Felis silvestris Schreber, 1775 in the lower

648 levels of the Visogliano site (north-eastern Italy) (Abbazzi et al., 2000; Sardella et al., 2006; Masini

649 and Sala, 2007; Falguères et al., 2010) would move the FO of this species back to MIS 13/12, in the

650 Fontana Ranuccio FU.

651 Among the sites reviewed by Marra et al. (2018), F. silvestris occurs only in the local fauna

652 of Polledrara di Cecanibbio, Torre in Pietra FU, substantially coeval with that of Kudaro 1, layer 5c,

653 in the Caucasus (Baryshnikov, 2011). Nevertheless, the taxonomic position of Middle Pleistocene

654 wildcats and the origin of F. silvestris in the Italian Peninsula is debated (Ficcarelli and Torre, 1974;

655 Yamaguchi et al., 2004).

656 The fossil record suggests that the transition from the possible ancestor, the Villafranchian

657 Felis lunensis Martelli, 1906, to the modern F. silvestris may have occurred during Middle

658 Pleistocene (Kurtén, 1965), maybe by MIS 11 (García et al., 1997 with references). However,

659 fossils safely determined as wildcat are recorded only from the Late Pleistocene onwards in 26 660 southern and northern Africa and the Near East (Kowalski and Rzebik-Kowalska, 1991; García et

661 al., 1997; Yamaguchi et al., 2004, and references therein).

662

663 Dama sp.

664 The chronostratigraphic re-examination of the sites correlated with MIS 9, MIS 7 and MIS 5

665 in the area around Rome confirms the absence of Dama dama tiberina Di Stefano & Petronio, 1997

666 in the deposits older than MIS 8.5 (Via Mascagni succession), while this archaic form of modern

667 fallow deer widely co-occurs with E. hydruntinus Regalia, 1907 in almost all the sites referred to

668 MIS 8.5 and MIS 7, such as Sedia del Diavolo, Batteria Nomentana, Vitinia upper levels, Torre in

669 Pietra upper levels and Saccopastore (Marra et al. 2017a, 2018). In the only site referred to MIS 5,

670 Fosso del Cupo, the presence of D. dama dama (Linnaeus, 1758) has been reported (Ceruleo et al.,

671 2016; Marra et al., 2018). In particular, previous doubtful attribution to D. dama tiberina of the

672 more or less fragmentary remains of deer from km 18.7 - 18.9 of Via Aurelia (MIS 13) by Anzidei

673 et al. (1993), was revised as Dama sp. by Marra et al. (2018). Similarly, a fragment of the blade of a

674 fallow deer in Rignano Flaminio (MIS 11) was attributed to Dama sp. by Petronio et al. (2017).

675 As known (Di Stefano and Petronio, 1997), the genus Dama occurred with the archaic

676 species Dama clactoniana (Falconer, 1868) during the MIS 15 in the Isernia FU and was still

677 present in several deposits assigned to MIS 11 (e.g., Fontana Ranuccio local fauna). From MIS 8.5-

678 7 (Vitinia FU) the first occurrence of D. dama tiberina is reported (Di Stefano and Petronio, 1997,

679 2002). This subspecies is slightly smaller than the previous species and is characterized by a

680 morphology of the antlers with daggers tines pointing backwards that differs from D. clactoniana

681 and resembles that of the Late Pleistocene subspecies D. dama dama. This archaic form of D. dama,

682 as mentioned, is present in numerous sites around Rome (Vitinia, Ponte Milvio, Batteria

683 Nomentana, Sedia del Diavolo, Vigna San Carlo, Saccopastore), all correlated with Vitinia FU

684 (MIS 8.5-7) (Di Stefano and Petronio, 1997; Marra et al., 2015, 2018), in Western Europe (Di

27 685 Stefano and Petronio, 1997), and probably also in southern Italy (Abbazzi et al., 2001; Kotsakis and

686 Barisone, 2008).

687 The remains indicated asreferred to Dama cf. D. dama in Anzidei et al. (1993), do not seem

688 to show peculiar diagnostic features that allow recognition, except for the smaller size compared to

689 the characteristic fallow deer of Clacton, which, however, could refer to the sexual dimorphism that

690 characterizes all the cervids. However, considering the chronostratigraphic revision in Marra et al.

691 (2018) for the two deposits of Via Aurelia (Anzidei et al., 1993) and for Rignano Flaminio

692 (Petronio et al., 2017), correlated with MIS 13-11, a more certain systematic determination needs to

693 be addressed on these fallow deer remains and / or new discoveries.

694 From a strictly paleobiological point of view, however, the attribution of the remains from

695 these localities to the two fallow deer species diffused in the late Middle Pleistocene (as different

696 species, the coexistence of D. clactoniana and D. dama tiberina can not be excluded) it would be

697 possible, as it would seem to happen, for example, in Sedia del Diavolo upper gravel (MIS 8.5).

698 Further studies and discoveries may confirm these observations.

699

700 Cervus elaphus ssp.

701 Cervus elaphus acoronatus Beninde, 1937 is the most archaic form of the red deer and is

702 characterized by simple antlers with 5 daggerstines: brow tine, bez tine, trez tine and terminal fork

703 transversal to the sagittal axis of the body (Di Stefano and Petronio, 1992, 1993, 2002; Di Stefano

704 et al., 1994, 2015). This subspecies, probably derived from Cervus grayi Zdansky, 1925 of the

705 Asian Villafranchian (similar to the living Cervus nippon Temminck, 1838), appears in Italy in the

706 early-middle Galerian (Slivia FU, MIS 21) and no certain presences are reported after the Isernia

707 FU (MIS 15). In fact, the morphology of the remains of cervids of Visogliano (MIS 13/12)

708 attributed to C. elaphus acoronatus by Abbazzi et al. (2000) should be re-evaluated.

709 In the Late Galerian, Fontana Ranuccio FU, the FO of Cervus elaphus eostephanoceros Di

710 Stefano & Petronio, 1993 (Petronio et al., 2011) is reported., a mMore evolved subspecies than the 28 711 previous C. elaphus acoronatus, this subspecies is characterized by antlers in which the trez tine

712 moves towards the terminal part and from the appearance of supernumerary daggers tines forming a

713 primitive terminal crown. This subspecies (Di Stefano and Petronio, 1993) had a

714 diffusiondistribution that covered the whole Italian peninsula and part of Western Europe;

715 moreover, C. elaphus eostephanoceros it does not seem to be reported in sites of different age than

716 MIS 11. The remains of the red deer of Guado San Nicola (MIS 11) attributed to C. elaphus

717 acoronatus by Peretto et al. (2015) can also be assigned to the same chronosubspecies.

718 Cervus elaphus rianensis Leonardi & Petronio, 1974, an endemic subspecies with simpler

719 antlers and more archaic characters than the previous subspecies, more directly related to C. elaphus

720 acoronatus (Leonardi and Petronio, 1974; Petronio et al., 2011), was considered a common

721 occurrence in the early and middle Aurelian of Latium (Gliozzi et al., 1997, Palombo et al., 2002,

722 Petronio et al., 2011), which in the Late Pleistocene was suceceded in the Late Pleistocene by the

723 extant subspecies of C. elaphus (see Petronio et al., 2007, 2011).

724 Considering the chronostratigraphic revision of the deposits of central Italy in which all

725 these taxa were recovered (Marra et al. 2018), some systematic reconsiderations should be made. In

726 fact, the reports of C. elaphus eostephanoceros in Cava Nera Molinario (MIS 13), in Rignano

727 Flaminio (MIS 11) and Fontana Ranuccio (MIS 11), almost contemporary to C. elaphus rianensis

728 in the sites of Riano (MIS 11) and Castel di Guido (MIS 11) pose the problem of the possible

729 presence of two subspecies in the same area and in the same period.

730 This compresence, however, must be underlined, is only apparently contemporary. We must

731 remember, in fact, the great and rapid adaptability of cervids to different environments (e.g., Nussey

732 et al., 2005; Torres-Porras et al., 2009), due to their high phenotypic plasticity that facilitates the

733 development of local subspecies with peculiar characteristics, starting from isolated populations

734 (Lister, 2004). This phenomenon is widely present and studied in the extant populations of C.

735 elaphus and allows this species, in the course of a few generations (obviously not detectable in

736 paleontological times), to differentiate into distinct populations, morphologically and 29 737 geneticallysubspecies. Therefore, considering the particular paleoecological and paleogeographic

738 conditions of the Italian peninsula. while the presence of the subspecies C. elaphus eostephanoceros

739 in a large part of Europecontinues during MIS 13-11, throughout the European continentcould

740 justify, in the particular paleoecological and paleogeographical conditions of the Italian peninsula,

741 the simultaneous appearance in MIS11 the rapid devolopment of the morphological characteristics

742 of C elaphus rianensis, an endemic form of central Italy, can be considered justified. The

743 conservative morphological structures of this form are also common in deer of insular faunas and /

744 or, in any case, with partial degrees of endemism (e.g., Cervus elaphus tyrrenicus Azzaroli, 1961,

745 Late Pleistocene, Capri island; - Capasso Barbato and Gliozzi, 1998). C. elaphus rianensis, together

746 with other subspecies of the late Middle Pleistocene (for example Cervus elaphus aretinus Azzaroli,

747 1961 of the Val di Chiana, Angelelli, 1981) would therefore be attributable to the endemic

748 subspecies rank, typical only of some territories of the Italian peninsula, with reminiscent

749 morphological characteristics, also in the structure of the antlers, of those of C. elaphus acoronatus.

750 This phenomenon seems to have been realized in the final part of the Galerian (Riano local fauna)

751 and, beyond the previous considerations, from the paleontological point of view, it is supported by

752 the fact that the two subspecies C. elaphus eostephanoceros and C. elaphus rianensis have never

753 been found in the same fossiliferous site, nor morphotypes with intermediate characteristics have

754 ever been found. In fact, from the morphological and, in part, biometric point of view, the two

755 forms are easily identifiable (Di Stefano and Petronio, 1997), and each one shows substantially

756 homogeneous characteristics within every single population.

757

758 5. Discussion

759 The revision of From the data relating to the biochronological history of Galerian and

760 Aurelian (Gliozzi et al., 1997) combined with and the new chronostratigraphic data (Marra et al.,

761 2018), emerges the problem concerning the current validity and the new meaning of Fontana

762 Ranuccio, Torre in Pietra, and Vitinia FUs. 30 763 As can be seen from the problems related to the biochronological modifications mentioned

764 above and from Figure 3, the Fontana Ranuccio FU is richer in sites and apparently more significant

765 from a paleobiological point of view compared to the Torre in Pietra and Vitinia FUs. The faunal

766 renewal of the Italian peninsula seems in fact to have occurred at the transition between the Isernia

767 FU (MIS 15) and Fontana Ranuccio FU (MIS 13-11) instead at the transition between Galerian and

768 Aurelian Mammal Ages, as traditionally held (Gliozzi et al., 1997; Petronio et al., 2011) (Fig.ure 3).

769 This renewal, which mainly concerns the genera Ursus, Canis and Vulpes, however, is more

770 synchronous with the faunal renewal in Western Europe.

771 Descending from the backdating of the many taxa listed in Figure 3, we must therefore

772 question whether the renewal of mammal assemblages that largely falls within the MIS 13-11

773 interval of the Fontana Ranuccio FU can still justify the Galerian-Aurelian transition as it is

774 currently focused on the Torre in Pietra FU, which records the FOs of Megaloceros giganteus and

775 Mustela putorius, only.

776 In Western Eurasia, M. giganteus occurred about 400 ka BP, during MIS 11 (Lister et al.,

777 2005; Vislobokova, 2012). Among the oldest remains of the species in Italy, in addition to Torre in

778 Pietra lower levels, we can mention those found in Latium at the sites of Campoverde and Ponte

779 Mammolo (Vitinia FU) (Petronio et al., 2011; Marra et al., 2017a), which are correlated with MIS

780 8.5 based on chronostraigraphic and biostratigraphic constraints (Vianello et al., 1995; Marra et al.,

781 2017a). In the Late Pleistocene M. giganteus is present in several sites of the peninsula, in particular

782 in northern Italy, until its local extinction during the Lateglacial, Dryas III (Petronio et al., 2007,

783 2014).

784 More complex to reconstruct is the palaeontological history of the European polecat, M.

785 putorius, also complicated by the Pleistocene presence in Eastern and Central Europe of the twin

786 species Mustela eversmanni, the steppe polecat, during the late Middle and Late Pleistocene. The

787 oldest reports refer to remains of dubious taxonomic attribution and / or uncertain stratigraphic

788 position, as in Miesenheim I or in Grotta del Cerè (Kolfshoten, 1996; Koenigswald and Heinrich, 31 789 1999; Rossi and Santi, 2001). Polecat remains attributed with reasonable certainty to M. putorius in

790 Western Europe date back to the early-middle part of the Middle Pleistocene, as in Mosbach 2 and

791 Atapuerca SH (Koenigswald and Heinrich, 1999; García and Arsuaga, 2001), respectively

792 correlated with Isernia FU and Fontana Ranuccio FU, while in France the species is registered only

793 in some sites related to MNQ24 (Argant, 2000; Palombo and Valli, 2004), correlable with the early

794 Aurelian. The most ancient indication of the species in Italy would be that of Cretone (Di Canzio et

795 al., 2003), in a faunistic assemblage referring to Torre in Pietra FU (Petronio et al., 2011). However,

796 given the rarity and fragmentary nature of the fossil remains of mustelids, precocious penetration

797 into the peninsula cannot be excluded a priori even for this carnivore.

798 Therefore, considering the still undisproved FO of M. giganteus in MIS 9 in Italy, we can

799 still consider the local fauna of Torre in Pietra lower levels representative of the early Aurelian

800 (Torre in Pietra FU) and thus propose this local fauna as a conventional boundary for the Galerian -

801 Aurelian transition, as was supported by Gliozzi et al. (1997) about the local fauna of Colle Curti,

802 characterized only by the FO of Praemegaceros verticornis, as a conventional boundary of the

803 Villafranchian-Galerian transition.

804 Moreover, the faunal transitions cannot be considered as a point on the timeline, but have

805 always a dynamic character. If we keep this notion in mind, we can observe that MIS 9 marks the

806 actual moment since when the faunistic assemblages are reprersented only by those taxa

807 characterizing the late Middle Pleistocene and Late Pleistocene (taxa with occurrences in green in

808 Fig.ure 3), and with with Bos primigenius and Palaeoloxodon antiquus prevailing, accompanied by

809 cervids and rarer equids and rhinocerotids. On the other hand, some typical Galerian taxa (e.g.,

810 Dama clactoniana, Bison schoetensacki and Ursus deningeri; taxa with occurrences in red in

811 Fig.ure 3) persist in the faunal assemblages of the latest Galerian (Fontana Ranuccio FU).

812 We also note that, although the Torre in Pietra FU is impoverished and the bioevents that

813 characterize Torre in Pietra FU and Vitinia FU are rather close in time, they are nevertheless

814 distinctive (Di Stefano et al., 1998; Petronio et al. 2011) and well recognizable, at least locally, 32 815 given that mammalian assemblages are contained in two distinct formations (Marra et al., 2014a,

816 2015, 2017a, 2018; Pandolfi and Marra, 2015).

817 As for the Cervus elaphus subspecies, C. elaphus eostephanoceros has a large geographical

818 distribution and a biochronological range interval limited to MIS 13 and MIS 11, Fontana Ranuccio

819 FU. Therefore, this taxon can still be considered a significant biochronological marker. In contrast,

820 C. elaphus rianensis is a local form present only in some MIS 11 sites of Latium, such as Riano

821 and, possibly, Castel di Guido, whereas other remains referred in the past literature to this taxon

822 occurring at location dated within MIS 9 and MIS 7, or within MIS 13 (e.g., Via Flaminia km 8.2),

823 are more likely morpho-types or sub-species to be still identified.

824 Finally, with regard to the strong faunal renewal in MIS 13, with 6 5 FOs, we can note the

825 coincidence with the persisting of temperate climatic conditions due to the absence of particularly

826 marked glacial periods that could have favored the above FOs and especially the subsequent spread

827 of these taxa (note the thermal threshold represented by the dashed blue line in Fig.ure 3). In fact,

828 the lowstand of MIS 14.2, which marks the glacial between the MIS 15 and the 13, is the least

829 marked of the last 600,000 years, allowing for over 100,000 years of mild climate. In the MIS 11

830 (second part of the Fontana Ranuccio UF), preceded and followed by two notable glacial peaks

831 (MIS 12 and MIS 10), there is the FO of Canis lupus and of endemic forms of deer such as C.

832 elaphus rianensis. The subsequent faunal renewal during the MIS 9 and MIS 7 is also characterized

833 by the persistence of temperate climatic conditions, with two mild glacial (MIS 8.4, MIS 8.2)

834 interspersed with a warm peak (MIS 8.5).

835

836 6. Conclusions

837 The recent chronostratigraphic revision of several sedimentary successions of the Latium

838 Tyrrhenian area, near Rome, previously referred to the Aurelia Formation (MIS 9), has shown that

839 some of these deposits are in fact to be attributed to Valle Giulia Formation (MIS 13) and to the San

33 840 Paolo Formation (MIS 11). Consequently, the faunistic assemblages found in these sediments,

841 previously attributed to Torre in Pietra FU, must be referred to Fontana Ranuccio FU.

842 The presence of Ursus spelaeus in MIS 13, and of Vulpes vulpes at the MIS 12-MIS 11

843 transition, in MIS 13 and of Canis lupus, and Cervus elaphus rianensis in MIS 11, involves a

844 systematic and above all biochronological reworking revision of the range of these mammal taxa

845 and the redefinition of Fontana Ranuccio and Torre in Pietra FU's, and consequently, also of the

846 Galerian - Aurelian transition.

847 The characteristic faunal renewal of the Aurelian Mammal Age was more gradual and above

848 all occurred earlier than previously assumed. Many taxa that will be typical of the Late Pleistocene

849 register their FO in the Fontana Ranuccio FU, latest Galerian. However, in analogy with what was

850 claimed by Gliozzi et al. (1997) regarding the transition from Villafranchian to Galerian, justified

851 only by the FO of Praemegaceros verticornis, it can be considered coherent to keep the current

852 biochronological boundary valid and to assigne to the local fauna of Torre in Pietra lower levels,

853 with the FO of Megaloceros giganteus, the conventional role to mark the beginning of the Aurelian

854 Mammal Age during MIS 9. On a more global perspective, it should be remarked that this faunal

855 trransition in the European continent and the Caucasian region appears as an even more gradual

856 event, spanning MIS 13 thoroght to MIS 9, and eyewitnessing a complete renewal by MIS 8.5.

857 Incidentally, the latter is also the time during which Homo neanderthalensis spread through

858 peninsular Italy and Europe (Marra et al., 2017a, and references therein).

859 The late Galerian, Fontana Ranuccio FU (MIS 13 - MIS 11), is characterized by the total

860 disappearance of Villafranchian taxa, with the exception of Homotherium, from the persistence of

861 typical Galerian taxa such as Dama clactoniana, Bison schoetensacki and Ursus deningeri, and

862 from the FO of the rhinos Stephanorhinus kirchbergensis and S. hemitoechus, of various other

863 herbivores such as Hippopotamus amphibius and Cervus elaphus eostepahnoceros and of various

864 carnivores such as Ursus spelaeus, Canis lupus and Vulpes vulpes.

34 865 The next FU marks the beginning of the Aurelian and, as mentioned above, is characterized

866 by the FO of Megaloceros giganteus and probably also of the European polecat, Mustela putorius.,

867 and a characteristicThe Aurelian assemblages are dominated by Bos primigenius, and

868 Palaeoloxodon antiquus and by cervids. Although chronologically very close to the previous one, in

869 Vitinia FU we note the LO of D. clactoniana and occurs along with the FO of the small equid

870 Equus hydruntinus, of the modern fallow deer with the primitive subspecies Dama dama tiberina

871 and the first archaic Neanderthals. It is therefore proposed for the moment to maintain the

872 distinction between the two FUs.

873 Finally, regarding the problematic occurrences of Crocidura and Arvicola it is suggested to

874 classify the remains of Torre in Pietra upper levels and those of Aurelia km 18.7 and km 18.9 as

875 Crocidura sp. and Arvicola sp.

876

877

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47 1342 Figure captions

1343

1344 Figure 1 - Location map of the fossilferous sites of the Aurelian Mammal Age and other sites

1345 described in this work.

1346

1347 Figure 2 - a) Composite cross-section showing the chrnostratigraphic setting at the sites along Via

1348 Aurelia (modified from Marra et al., 2018). Position of sample MG4 dated for this work and the

1349 occurrences of the taxa previously considered exclusive of MIS 9 are shown.

1350 b) Geologic sections of Riano and Via Flaminia km 8.2 showing the occurrences of the Late

1351 Galerian and Aurelian taxa whose biostratigraphy is discussed in this paper.

1352 c) 40Ar/39Ar ages (vertical colored bars) of the volcanic deposits providing correlation between the

1353 sedimentary deposits and the aggradational successions (colored boxes) deposited in response to

1354 sea-level rise during the glacial terminations, and with the stages of the Oxygen Isotope curve (from

1355 Lisiecki and Raymo, 2005).

1356 Legend- TTPB: Tufi Terrosi con Pomici Bianche; GRPS: Grottarossa Pyroclastic Sequence; TGPP:

1357 Tufo Giallo di prima Porta; TP: Tufo del Palatino; TGS: Tufo Giallo di Sacrofano; TRSN: Tufo

1358 Rosso a Scorie Nere; VGF: Valle Giulia Formation; SPF: San Paolo Formation.

1359

1360 Figure 3 - Biochronological scheme showing the range and the FO (black arrows) of the principal

1361 taxa constituting the faunal assemblages of the late Galerian and early-middle Aurelian Mamal

1362 Ages of peninsular Italty. FO of the taxa in bold were previously reported within MIS 9.

1363 Comparison with the Oxygen Isotope timescale (Lisiecky and Raimo, 2005) is also shown. See text

1364 for comments and explanations.

1365

1366 Table 1 - Revised chronology for the sites of the Aurelian Mammal Age.

1367

48 1368 Table 2 - 40Ar/39Ar results.

PREVIOUS ATTRIBUTION REVISED AGE AND ATTRIBUTION SITE [1] [2] FORMATION MIS AGE (ka) Via Flaminia km 8.2 T MIS 7 n.d. Valle Giulia Fm 13 533±2 [3] Cava Rinaldi upper level O MIS 9 TIP FU Valle Giulia Fm 13 516±1 [4] Malagrotta (MG2) R MIS 9 TIP FU Valle Giulia Fm 13 516±1 [3] R Via Aurelia km 19.3 MIS 9 TIP FU San Paolo Fm-? 11? ~410 ka?* E Collina Barbattini MIS 9 TIP FU San Paolo Fm-? 11? ~410 ka?* Castel di Guido I MIS 9 TIP FU San Paolo Fm 11 412±2 [3] Riano N MIS 9 TIP FU San Paolo Fm 11 406±5 [3] Torre in Pietra lower level s MIS 9 TIP FU Aurelia Fm 9 355-335 [5] Polledrara di Cecanibbio P MIS 9 TIP FU Aurelia Fm 9 325±2 [6] Prati Fiscali I MIS 9 VIT FU Via Mascagni Sc 8.5 285±2 [3] E Sedia del Diavolo upper level MIS 9 VIT FU Via Mascagni Sc 8.5 285±2 [4] T [4] Monte delle Gioie R MIS 9 VIT FU Via Mascagni Sc 8.5 285±2 Cerveteri-Migliorie di S. Paolo A MIS 7 VIT FU Vitinia Fm 8.5/7 290-200 [3] Torre in Pietra upper levels MIS 7 VIT FU Vitinia Fm 7 270-240 [5] Casal de' Pazzi F MIS 7 VIT FU Vitinia Fm 7 270-250 [7] Vitinia upper level U MIS 7 VIT FU Vitinia Fm 7 253±8 [4] Saccopastore n.a. MIS 5 MEL FU Vitinia Fm 7 245-220 [7,8]

Table 1 - Revised chronology for the sites of the Aurelian Mammal Age. TIP: Torre in Pietra; VIT: Vitinia; MEL: Melpignano; FU: Faunal Unit; MIS: Marine Isotope Stage; Fm: Formation; Sc: succession; n.d.: not defined. [1] Palombo, 2004; Palombo et al., 2004; [2] Palombo et al., 2002; Petronio et al., 2011; [3] Marra et al. 2018; [4] Marra et al., 2014a; [5] Villa et al., 2016; [6] Pereira et al., 2017a; [7] Marra et al., 2017a; [8] Marra et al., 2015; *This work. TABLE 2 - 40Ar/39Ar results Single crystal fusions Sample: MG4 J-value: 0.0004007 ± 0.0000010 (2σ) Material: sanidine 40 39 37 36 Laser Ar ± 2σ40 Ar ±2σ39 Ar ± 2σ37 Ar ± 2σ36 Included in 40 39 40 File power (%) (cps) (cps) (cps) (cps) (cps) (cps) (cps) (cps) Ar*/ ArK ± 2σ % Ar* Age (ka) ± 2σ (ka) K/Ca wtd mean A NAG6042 30 483306 ± 131 796029 ± 923 8210 ± 42 155.89 ± 3.04 0.548959 ± 0.003471 90.42 402.6 ± 2.5 41.691 * NAG6044 30 470573 ± 124 696231 ± 882 8629 ± 41 289.87 ± 4.20 0.552028 ± 0.003696 81.67 404.9 ± 2.7 34.693 * NAG6047 30 186054 ± 74 329913 ± 447 6259 ± 35 16.93 ± 1.80 0.549598 ± 0.004410 97.45 403.1 ± 3.2 22.666 * NAG6049 30 224632 ± 88 374201 ± 492 5230 ± 34 57.81 ± 1.95 0.554749 ± 0.004053 92.41 406.9 ± 3.0 30.763 * NAG6052 30 215867 ± 79 342113 ± 443 4666 ± 33 90.32 ± 2.39 0.552700 ± 0.004082 87.59 405.3 ± 3.0 31.527 * NAG6054 30 464554 ± 119 274974 ± 323 5703 ± 33 1049.88 ± 9.90 0.550618 ± 0.010898 32.59 403.8 ± 8.0 20.732 * NAG6057 30 174853 ± 82 300260 ± 390 6148 ± 41 65.48 ± 2.17 0.518314 ± 0.004351 89.00 380.1 ± 3.2 21.000 YES NAG6059 30 322885 ± 97 577436 ± 737 7577 ± 37 76.21 ± 2.33 0.520272 ± 0.004100 93.04 381.6 ± 3.0 32.769 YES NAG6062 30 423730 ± 128 681430 ± 866 8531 ± 38 157.05 ± 3.14 0.553472 ± 0.003768 89.01 405.9 ± 2.8 34.346 * NAG6064 30 336781 ± 114 555935 ± 667 7413 ± 41 102.10 ± 2.66 0.551482 ± 0.003676 91.03 404.5 ± 2.7 32.246 * weighted mean age A (2 of 10): 380.9 ± 2.1

*(8 of 10): 404.6 ± 1.1

The values in this table have been corrected for instrument background, source mass bias, detector efficiency, and decay of 37Ar and 39Ar Instrument: Noblesse 5-collector mass spectrometer

Standard: Alder Creek rhyolite sanidine Standard age (Ma): 1.1864 ± 0.0003 Jicha et al. (2016)

Atmospheric argon ratios 40Ar/36Ar 298.56 ± 0.31 Lee et al. (2006) 38Ar/36Ar 0.1885 ± 0.0003 Lee et al. (2006) Interfering isotope production ratios 40 39 ( Ar/ Ar)K 0.00054 ± 0.00014 Jicha & Brown (2014) 38 39 ( Ar/ Ar)K 0.01210 ± 0.00002 Jicha & Brown (2014) 39 37 ( Ar/ Ar)Ca 0.000695 ± 0.00001 Renne et al. (2013) 38 37 ( Ar/ Ar)Ca 0.0000196 ± 0.0000008 Renne et al. (2013) 36 37 ( Ar/ Ar)Ca 0.000265 ± 0,00002 Renne et al. (2013) Decay constants -10 -1 40Ar (0.580 ± 0.014) x 10 a -10 -1 B- (4.884 ± 0.099) x 10 a 39Ar (2.58 ± 0.03) x 10-3 a-1 37Ar (8.23 ± 0.042) x 10-4 h-1 36Cl (2.303 ± 0.046) x 10-6 a-1