Rivista Italiana di Paleontologia e Stratigrafia volume 117 no. 1 pp. 173-187 April 2011 STEPHANORHINUS ETRUSCUS (FALCONER, 1868) FROM PIRRO NORD (APRICENA, FOGGIA, SOUTHERN ITALY) WITH NOTES ON THE OTHER LATE EARLY PLEISTOCENE RHINOCEROS REMAINS OF ITALY LUCA PANDOLFI 1 & CARMELO PETRONIO 1* Received: November 12, 2010; accepted: February 3rd, 2011 1 Dipartimento di Scienze della Terra, Sapienza, Università di Roma, p.le Aldo Moro 5, 00185 Roma, Italy. *Corresponding author. E-mail address: carmelo.petronio@uniroma1.it Key words: Stephanorhinus etruscus, Stephanorhinus hund- sheimensis, morphology, morphometry, Pirro Nord, late Early Pleis- tocene, Italy. Abstract. During the late Early Pleistocene (latest Villafranchi- an-earliest Galerian), a marked faunal turnover occurred in Italy, with a progressive disappearance of Villafranchian species and the gradual arrival of new species that later characterized the Middle Pleistocene. Two rhinoceros species are reported during this time: Stephanorhinus etruscus and S. hundsheimensis. The morphological and morphomet- rical analysis of the rhinoceros remains unearthed in the quarries of Pirro Nord (Pirro Faunal Unit, about 1.3-1.6 Ma) show a great affin- ity with S. etruscus, in particular with its second evolutionary stage. This stage is characterized by a general variation in body size and proportions. Other rhinoceros remains, found in Italian sites chrono- logically correlated with Pirro Faunal Unit (FU), are also ascribed to the second evolutionary stage of S. etruscus. In addition, in Italy, the presence of S. hundsheimensis is unknown before the Colle Curti FU (about 1 Ma). This is likely related to the scarcity of sites chronologi- cally referable to the Early-Middle Pleistocene transition or to a delay in the dispersal event of this species in Italy. Furthermore, the occur- rence of S. hundsheimensis in Europe and in Italy may be related to the climatic deterioration of the latest Early Pleistocene, as may be the variation in proportions and size of S. etruscus. Riassunto. Durante la fine del Pleistocene Inferiore (tardo Villafranchiano - inizio del Galeriano), in Italia si ha un marcato rin- novamento faunistico, con la progressiva scomparsa di specie villa- franchiane e il graduale arrivo di nuove specie che caratterizzeranno il Pleistocene Medio. Durante questo periodo di tempo, sono segnalate due specie di rinoceronte: Stephanorhinus etruscus e S. hundsheimen- sis. L’analisi morfologica e morfometrica dei resti di rinoceronte rin- venuti nelle cavità carsiche del sito di Pirro Nord (Unità Faunistica di Pirro, circa 1,3-1,6 Ma) ha permesso di riferire gli stessi al secondo sta- dio evolutivo della specie S. etruscus. Questo stadio è caratterizzato da una generale variazione della taglia corporea e delle proporzioni. Alla stessa specie vengono poi riferiti altri resti di rinoceronte rinvenuti in siti italiani cronologicamente correlabili con l’Unità Faunistica di Pirro. Inoltre, in Italia, la presenza di S. hundsheimensis sembrerebbe essere certa solo a partire dalla Unità Faunistica di Colle Curti (circa 1 Ma). Ciò è probabilmente da mettere in relazione con la scarsità di siti cronologicamente riferibili alla transizione Pleistocene Inferiore-Me- dio oppure è da correlare con un ritardo nell’evento dispersivo della specie in questione nella Penisola. Infine, la comparsa di S. hundshei- mensis in Europa ed in Italia potrebbe essere legata al deterioramento climatico verificatosi dalla fine del Pleistocene Inferiore, così come la variazione di taglia e proporzioni di S. etruscus. Introduction Rhinoceroses represent a common element of the Plio-Pleistocene faunas of Europe but they are, how- ever, a poorly known taxon. The identification of the different rhinoceros species as well as determining their temporal and spatial distribution is sometimes very dif- ficult. The taxonomy of fossil Rhinocerotidae is still debated and the major reason for this is, according to Heissig (1981) and Fortelius et al. (1993), “that the rhinoceroses are a highly stereotyped group with little morphological divergence”. In the past centuries, all remains of European rhinoceroses not ascribed to the genus Coelodonta were referred to the genus Rhinoceros and later to the genus Dicerorhinus. Guérin (1980) ascribed them to the genus Dicerorhinus and the new subgenus Brand- torhinus; later, this last was elevated to the genus rank (Guérin 1989). However, according to Groves (1983), the Pleistocene European rhinoceroses show little similarity with the genus Dicerorhinus. In fact, among 174 Pandolfi L. & Petronio C. the other characters, lost or non-functional incisors and a partially ossified nasal septum distinguish them from the genus Dicerorhinus. Despite this, some au- thors continue to refer them to the present-day genus (McKenna & Bell 1997; Guérin 2004). The nomenclature used in this work follows by Fortelius et al. (1993) and the European Pleistocene rhinoceroses, except for Coelodonta and Elasmothe- rium, are referred to the genus Stephanorhinus as pre- viously used by several authors (Heissig 1973, 1981, 1989; Cerdeño 1993, 1995, 1998; Prothero et al. 1986; inter alios). This choice is based on the morphological characters differentiating between Dicerorhinus and European fossil rhinoceroses, as already discussed by other authors (see Groves 1983; Fortelius et al. 1993; Lacombat 2005). During the time span between the Late Villa- franchian and earliest Galerian (late Early Pleistocene), two rhinoceros species are known from Europe: Stepha- norhinus etruscus (Falconer, 1868) and S. hundshei- mensis (Toula, 1902) but their stratigraphic range is not well-defined. In the Italian literature (Gliozzi et al. 1997; Fortelius et al. 1993; Mazza et al. 1993; Palombo et al. 2002; inter alios), the Etruscan rhinoceros was first present as early as the Montopoli Faunal Unit and lasted until the Tasso FU. The Hundsheim rhinoceros was instead found in the recent Galerian sites. In 1993, Mazza et al. reported small-sized rhinoceros popula- tions in the Farneta, Pirro and Colle Curti FUs (lat- est Villafranchian-earliest Galerian), and they referred these rhinoceros to S. cf. hundsheimensis. Small-sized rhinoceroses were subsequently reported to be S. aff. hundsheimensis, S. cfr. hundsheimensis and S. hundshei- mensis (Mazza et al. 1993; Gliozzi et al. 1997; Lacombat 2005), thus extending the stratigraphic range of this last species. In Europe, the occurrence of these two species is still discussed and their chronological ranges are rela- tively well-defined (Guérin 1980; Fortelius et al. 1993). At present, S. etruscus is reported from sites correlated with the Late Villafranchian and the Early Galerian, while the first occurrence of S. hundsheimensis is re- corded during the latest Villafranchian. Furthermore, Guérin (1980) described two evolutionary stages of S. etruscus: the first stage related with the specimens from the Early and Middle Villafranchian sites of Europe and the second one related with the specimens from the Late Villafranchian sites. According to the previous author, the second stage is characterized by a general variation in body size and proportions (Guérin 1980: 598-604). Also, two size variants are recognized for S. hundshei- mensis (Guérin 1980; Lacombat 2005); they are prob- ably related with geographic and/or climatic factors. The present study is aimed at the examination of rhinoceros remains found in Italy, in reference to a chronological interval related to the latest Villafranchi- an and earliest Galerian. It will attempt to clarify the presence of rhinoceros species during this time span and their implications. Materials and methods The morphological characteristics considered in this work are those listed as diagnostic by several authors, including Guérin (1980), Mazza (1988), Fortelius et al. (1993) and Lacombat (2005, 2006). The morphometric methodology is based on the work of Guérin (1980), Mazza (1988), Fortelius et al. (1993) and Lacombat (2005, 2006). The metric values obtained for Italian rhinoceroses are compared with those of different European sites. Simpson ratio diagrams are made, using data on extant Diceros bicornis (data from Guérin 1980) as a reference. This reference was chosen to better highlight differences in specimen proportions and avoid errors of attribution derived using fossil species. The rhinoceros remains found in Pirro Nord are preserved in the museums of Palaeontology of Rome (MPUR), Turin (MPUT) and Florence (MPUF). The small collection in the Museum of Palaeontology at Sa- pienza University of Rome contains a lower molar (MPUR s.n.), the fragmented mandible of a young rhinoceros (MPUR s.n.) and a fourth metacarpal bone (MPUR s.n.). These remains were found in the karst cavity present near P10 and later destroyed during the quarry activity. One D3/ (MPUT PU106664), one M1/ (MPUT PU123658), one P/2 (MPUT PU s.n.), a third metacarpal bone (MPUT PU106398) remains were recovered in the last few years in the karst cavities P10 and P13, located in the Dell’Erba quarry; these remains are preserved in the Museum of Palaeontology of Turin. The material at the Museum of Geology and Palaeontology of Florence was found in karst cavity (called P80) in limestone quar- ries in Apricena and was ascribed to Dicerorhinus etruscus (De Giuli et al. 1987), Stephanorhinus cf. hundsheimensis (Mazza et al. 1993), and later to S. cf. S. etruscus (Abbazzi et al. 1996) and S. cfr. hund- sheimensis (Gliozzi et al. 1997). The material was listed by Mazza et al. (1993). Description and comparisons Rhinoceros remains from Pirro Nord The remains preserved in the Museums of Rome and Turin are in a good state of preservation. The teeth (Fig. 1) have a number of characteris- tics in common with S. etruscus and S. hundsheimensis. However, the following morphological characteristics appear to be more indicative of S. etruscus: • the presence of small cuspids at the outlets of the U-shaped valleys between protocone and hy- pocone in D3/; • the absence of the lingual cingulum in the same tooth; • the presence of a V-shaped posterior valley and a weak mesial cingulum in P/2; • the great difference of height between the bot- toms of the valleys in the M/1; • the profile of the ectoloph in M1/, slightly con- vex, lacking a marked paracone fold. Stephanorhinus etruscus from Pirro Nord and other late Early Pleistocene rhinoceros remains of Italy 175 In the lower deciduous teeth, the difference in height between the bottoms of the valleys is large (0.67 in D/3 e 0.72 in D/4), as in S. etruscus. Also, the V- shaped posterior valley, the absence of a mesial cingu- lum and the strong depth of the vestibular syncline in D/4 of Pirro Nord is a typical character of S. etruscus (see also Lacombat 2006). Fig. 1 - Rhinoceros remains from Pirro Nord. (A) First lower molar, lingual view; (B) Second lower premolar, (1) buccal view, (2) occlusal view; (C) Fragmentary right mandible, occlusal view. Scale bar: 2 cm. Tab. 1 - Comparative dimensions of lower deciduous teeth, D/1 and D/2, from Pirro Nord and of S. etruscus (S. etru.) (data from Mazza 1988; Lacombat 2006) and S. hundsheimensis (S. hund.) (data from Kahlke 2001; Lacombat 2006). L max = maximal length; l max = maximal breadth. D/1 L max l max Pirro Nord 16 8 Senèze (S. etru.) 17.52 9.55 med 18.34 10.82 Upper Valdarno+Olivola (S. etru.) min 18.17 10.67 max 18.51 10.97 med 17.86 11.22 Vallonnet (S. hund.) min 16.6 10.48 max 20.46 12.44 Isernia (S. hund.) 19.2 13.8 med 18.09 11.5 Untermassfeld (S. hund.) min 16 10.4 max 21 15 D/2 L max l max Pirro Nord 29.5 15 med 29.91 15.66 Senèze (S. etru.) min 29.28 15.1 max 30.55 16.62 med 38.45 21.66 Upper Valdarno+Olivola (S. etru.) min 37.9 21.66 max 39 21.66 med 31.12 17.69 Vallonnet (S. hund.) min 28.5 15.46 max 32.87 18.95 med 31.6 17.64 Untermassfeld (S. hund.) min 30 16.5 max 33.1 19.8 D/4 L max l max Pirro Nord 36.2 22 med 40.36 Senèze (S. etru.) min 39.5 max 41.84 med 38.47 22.7 Upper Valdarno (S. etru.) min 37.82 max 39.11 med 41.17 24.5 Isernia (S. hund.) min 40 max 42.39 med 40.05 24.29 Vallonnet (S. hund.) min 38.1 23.42 max 41.99 25.46 med 42.13 23.93 Untermassfeld (S. hund.) min 39.8 22.8 max 44.5 25.8 Tab. 2 - Comparative dimensions of the lower deciduous tooth, D/4, from Pirro Nord and of S. etruscus (S. etru.) (data from Guérin & Heintz 1972; Mazza 1988) and S. hund- sheimensis (S. hund.) (data from Kahlke 2001; Lacombat 2006). L max = maximal length; l max = maximal breadth. 176 Pandolfi L. & Petronio C. The dimensions of the lower deciduous teeth are comparable with the minimum values of S. etruscus and are smaller than S. hundsheimensis (Tab. 1; Tab. 2). In fact, in this last species, the lower deciduous teeth are more developed than in S. etruscus (Lacombat 2006). The dimensions of the distal epiphysis of the third metacarpal bone (Fig. 2A) are also comparable with the minimum values of S. etruscus (Tab. 3). The fourth metacarpal bone (Fig. 2B) has a sub- triangular surface at its dorsal end. In medial view, two articular facets are present. The anterior one is stretched along the upper margin of the epiphysis, while the pos- terior facet is sub-elliptical and much wider than the anterior. These characters are present in S. etruscus (see also Mazza 1988; Lacombat 2005). The dimensions of the fourth metacarpal are comparable to the minimum values of Etruscan rhinoc- eros (see for example Guérin 1980) and are similar to that of Créspia (Tab. 4). In the Simpson diagram, the curve of the remains of Pirro Nord is close to S. etrus- cus of Senèze and La Puebla de Valverde. It is clearly different from S. hundsheimensis of Vallonnet, Isernia, Soleilhac and Untermaßfeld (Fig. 3). The rhinoceros remains preserved in the Muse- um of Geology and Palaeontology at the University of Fig. 2 - Rhinoceros remains from Pirro Nord. (A) Fragmentary third metacarpal bone, anterior view; (B) Fourth metacar- pal bone, anterior view. Scale bar: 2 cm. Fig. 3 - Ratio diagrams for the MC IV from Pirro Nord and Pi- etrafitta (data from Mazza et al. 1993) versus S. etruscus (S. etru.). (A) (data from Guérin & Heintz 1972; Mazza 1988) and S. hundsheimen- sis (S. hund.); (B) (data from Fortelius et al. 1993; Kahlke 2001; Lacombat 2005) (stand- ard Diceros bicornis). Abbre- viations in Tab. 4. Stephanorhinus etruscus from Pirro Nord and other late Early Pleistocene rhinoceros remains of Italy 177 Florence are well-described and depicted by Mazza et al. (1993). According to these authors, the rhinoceros from Pirro Nord shows similarities with both S. etrus- cus and S. hundsheimensis, despite the incomplete and fragmentary state of the remains. According to Mazza et al. (1993), all the teeth have morphology compara- ble to that of S. etruscus, while some postcranial ele- ments show differences from this last species. In par- ticular, these differences are observable in the scapulae, humeri, scaphoid and metapodial bones. Other post- cranial elements are morphologically comparable to S. etruscus from the Upper Valdarno. The morphology of the considered scapulae is similar to S. etruscus of Créspia (García-Fernandez et al. 2003b) and the dimensions are comparable to those of S. etruscus from different European sites and differ from those of S. hundsheimensis (Tab. 5). There are few morphological and metric differences between the humeri of S. etruscus and S. hundshemensis (Guérin 1980; Mazza 1988; Fortelius et al. 1993; Lacombat 2005). However, the morphological characteristics and morphometric values listed by Mazza et al. (1993) are comparable to those of S. etruscus (Mazza 1988; Kahlke 2001; Lacombat 2005). In the remains of the Museum of Florence, only one fragmentary scaphoid is present and its dimensions are comparable to that of S. etruscus. In particular, the proximal articular surface allows for it to be distin- guished from S. hundsheimensis (Lacombat 2005) (Fig. 4). The morphological characters described by Mazza et al. (1993) may fall within the intraspecific variation of the Etruscan rhinoceros. Only the proximal epiphysis of the third meta- carpal is preserved. The Simpson diagram of this last bone shows similarities with S. hundshemensis from Vallonnet and S. etruscus from La Puebla de Valverde and Senéze (Fig. 5). In addition, the proportions of the fourth metacarpal are comparable to those of S. etrus- cus (Fig. 6). The metatarsal bones do not greatly differ in proportion between the two species (Figs 7, 8) and the morphological characters, as listed by Mazza et al. (1993), could lie within the intraspecific variation of S. etruscus. Rhinoceros remains from other latest Villa- franchian-earliest Galerian sites of Italy Madonna della Strada (Scoppito, L’Aquila, cen- tral Italy) The sandy-lignitiferous deposits of Madonna della Strada are chronologically related to the latest Early Pleistocene (Magri et al. 2010). At this site, rhi- noceros remains were found together with Mammuthus meridionalis and were first ascribed to D. etruscus and later to S. aff. hundsheimensis and S. cf. hundsheimen- sis (Maccagno 1962; Caloi & Palombo 1995; Palombo et al. 2002). However, the morphological and metric characters of these remains make it possible to ascribe them to S. etruscus. P/2 (MPUR s.n.) is characterized by quite smooth enamel. The cingulum, represented by small cuspulae, occurs at the vestibular side of the tooth; this character can be seen also in the premolars of the S. etruscus specimens from the Upper Valdarno (Mazza 1988). The astragalus preserved in the Museum of San Giuliano (L’Aquila) (MSG 396), shows in anterior view, a convex, but quite small, trochlear distal side. The artic- ular surface is well defined in medial view, and the me- dial tuberculum is posterodistally oriented. In posterior view, the mesiodistal articular surface is sub-elliptical. Metatarsal IV (MPUR s.n.) has a sub-triangular medial- anterior articular facet, connected to the proximal side. The corresponding posterior facet is sub-circular and placed at a lower level than the anterior one. The morphology of these bones is consistent with an attribution to S. etruscus following the diag- nostic features pointed out by several authors (Guérin MC III DTDmax DAPD Pirro Nord PU106398 54.55 37.2 med 56.25 40.25 Upper Valdarno+Olivola (S. etru.) min 54 39 max 59 42 Capitone (S. etru.) 58 40 med 45.5 40 Puebla de Valverde (S. etru.) min 44 39 max 47 41 med 48.3 41 Senèze (S. etru.) min 48 40 max 49 42 med 58.14 43.5 Vallonnet (S. hund.) min 58.04 41.77 max 58.24 45.22 Soleilhac (S. hund.) 54.6 42.5 59.17 med 58.1 46.06 Untermassfeld (S. hund.) min 52 42 max 64.6 49.2 Mauer (S. hund.) 56.8 43.8 Mosbach (S. hund.) 53.7 42.1 med 63.1 45.7 Vergranne (S. hund.) min 61.5 45 max 65.5 46.5 med 53.4 41.15 Pietrafitta min 51.8 40.3 max 55 42 Tab. 3 - Comparative dimensions of MC III from Pirro Nord, Pie- trafitta (Mazza et al. 1993) and of S. etruscus (S. etru.) (data from Guérin & Heintz 1972; Mazza 1988) and S. hundshe- imensis (S. hund.) (data from Fortelius et al. 1993; Guérin 1983; Kahlke 2001; Lacombat 2005). DTDmax = maximal breadth of distal epiphysis; DAPD = depth of distal epi- physis. 178 Pandolfi L. & Petronio C. 1980; Fortelius et al. 1993; Lacombat 2005). Other re- mains discovered at this site, as recently described by Magri et al. (2010), consist of some fragmentary teeth and postcranial elements, probably of an old rhinoc- eros (MPUR s.n.). These remains were ascribed to S. cf. hundsheimensis, but no diagnostic character is present in the teeth analyzed by Magri et al. (2010). Also, the dimensions of the few and fragmentary postcranial ele- ments fall into size ranges of S. etruscus and S. hund- sheimensis (see Guérin & Heintz 1972; Guérin 1980). Morphological characters given by Magri et al. (2010) for the distal articular face of the cuboid and posterior articular faces of talus are present in S. etruscus from the Upper Valdarno (Mazza 1988). Colle Curti (Umbria, central Italy) and Casta- gnone (Monferrato, northern Italy) The early Galerian fauna is not well-know in Ita- ly, because of the limited fossil record of this time span. The Colle Curti local fauna was correlated with the base of Jaramillo subchron (about 1.1 Ma) and is more recent than the Pirro FU (Gliozzi et al. 1997). In this latter local fauna, Ficcarelli & Mazza (1990) ascribed three upper molars to S. etruscus, stating: “which can easily be attributed to Falconer’s Dicerorhinus etrus- cus” but that “this attribution is not conclusive, since recent observations on fossil rhinoceroses of the lat- est Early Pleistocene pointed out the occurrence of a new species, equipped with teeth very similar to those of Dicerorhinus etruscus”. Later, these remains were ascribed to S. cf. hundsheimensis (Mazza et al. 1993). However, the morphology of the upper molar, as fig- ured by Ficcarelli & Mazza (1990), and, generally, the morphology of the isolated upper molars, seems not to have diagnostic characteristics to distinguish be- tween S. etruscus and S. hundsheimensis (Guérin 1980; MC IV L max PTD PAPD TD diaph. APD diaph. DTDmax DAPD DTD artic. Pirro Nord s.n. 152 38 30 25 22 36 35 30 Pietrafitta 150 38 35.2 29 16 37 35.3 33 Pirro Nord* 32.6 19 36.1 37.8 34 med 167 41.3 35.8 31.1 22 42.7 37.9 37 Senèze (S. etru.) min 158 40 33 28 20 40 37 34 max 171 45 38 34 24 46 39 39 Crespia (S. etru.) 146.6 38 32.4 23.2 20 35 27 30 Puebla de Valverde (S. etru.) 162 42 32 30 22 38 36 35 44 33 med 176 43 37.75 30.5 21 46 42 Saint Vallier (S. etru.) min 41 36 30 max 45 40 31 med 166.75 39.75 37.6 34.66 18.83 40 36.5 33.75 Upper Valdarno+Olivola (S. etru.) min 157 37 32 32 17 33 34 32 max 176 43 41 39 20 46 39 36 med 169.41 45.37 41 33.5 23.5 43.58 40.83 38.41 Vergranne (S. hund.) min 167 43.5 38.5 29.5 22 40 39 37 max 172.5 47.5 48 37.5 24 45.5 43.5 41 41.8 44.3 Isernia (S. hund.) 42 44 167 45.5 44.5 33.7 22 42.2 42 37 med 174.12 43.26 41.62 36.93 22.36 46.22 39.6 38.02 Mosbach (S. hund.) min 164 42.8 38.5 34.6 20 42.1 37.2 34 max 181.3 44 44 39.8 23.1 50.2 41.2 41 Vallonnet (S. hund.) 153.2 39.52 37.55 33.22 20.44 45.59 39.2 38.02 Soleilhac (S. hund.) 181 41.92 37.06 17.82 47.67 44.54 165 37.02 37.17 19.67 41.82 38.45 36.37 Sussenborn (S. hund.) 43.7 46 med 174.01 42.01 43.68 33.53 20.78 44.78 41.88 40.53 Untermassfeld (S. hund.) min 162.9 37.8 41.2 31.6 19 41.5 40.3 35.9 max 186.3 45.5 48.4 34.8 21.9 48.4 44 44.9 Tab. 4 - Comparative dimensions of MC IV from Pirro Nord (* = data from Mazza et al. 1993), Pietrafitta (data from Mazza et al. 1993) and of S. etruscus (S. etru.) (data from Guérin & Heintz, 1972; Mazza 1988; García-Fernández et al. 2001) and S. hundsheimensis (S. hund.) (data from Fortelius et al. 1993; Guérin 1983; Kahlke 1969, 2001; Lacombat 2005). L max = maximal length of the bone; PTD = breadth of proximal epiphysis; PAPD = depth of proximal epiphysis; TD diaph. = breadth of diaphysis; APD diaph. = depth of diaphysis; DTDmax = maximal breadth of distal epiphysis; DAPD = depth of distal epiphysis; DTD artic. = breadth of distal articular surface. Stephanorhinus etruscus from Pirro Nord and other late Early Pleistocene rhinoceros remains of Italy 179 Fortelius et al. 1993; Lacombat 2006). The site of Castagnone was also correlated with the Colle Curti FU (Siori & Sala 2007). At this site, rhi- noceros remains consist of one M/3 and several frag- mentary teeth, ascribed “with some uncertainty” to S. cf. S. hundsheimensis by Siori & Sala (2007). Also in the site of Castagnone, the morphological and morpho- metrical characters do not allow for a distinction be- tween S. etruscus and S. hundsheimensis. In these cases, Scapula DTD TDCS LG BG Pirro Nord* 108 73 104 87 67 51 Pietrafitta 110 87 70 57 Upper Valdarno (S. etru.) 72 58 98 91 76 59 Créspia (S. etru.) 115 70 >60 Capitone (S. etru.) 111 86 74ca Senèze (S. etru.) med 134.91 109.91 84.42 65.63 min 128 104.95 78.11 60.16 max 150.88 117.8 91.5 71.3 Cagnes sur Mer (S. etru.) 123.64 107.49 76.26 62.12 Vergranne (S. hund.) 130 115.5 80 59 116ca 80 71 Untermassfeld (S. hund.) 120.8 108 81.5 75 108.8 73 Mosbach (S. hund.) 127 115 83 72.8 114 85 65 Tab. 5 - Comparative dimensions of scapulae from Pirro Nord (* = data from Mazza et al. 1993), Pietrafitta (data from Mazza et al. 1993) and of S. etruscus (S. etru.) (data from Guérin & Heintz 1972; Mazza 1988; García-Fernández et al. 2001) and S. hundsheimensis (S. hund.) (data from Fortelius et al. 1993; Guérin 1983; Kahlke 2001; Lacombat 2005). DTD = breadth of distal epiphysis; TDCS = breadth of collum scapulae; LG = length of the glenoid cavity; BG = breadth of the glenoid cavity. Fig. 4 - Bivariate diagram of scaphoids from Pirro Nord and Pietrafitta (data from Mazza et al. 1993), S. etruscus (S. etru.) (data from Mazza 1988) and S. hundsheimensis (S. hund.) (data from Kahl- ke 2001; Lacombat 2005). LPB = length of the proxi- mal articular surface; BPA = breadth of the proximal ar- ticular surface. all the rhinoceros remains should be ascribed to Stepha- norhinus sp. Cava Redicicoli (Rome, central Italy) The site of Cava Redicicoli, discovered by Blanc in the 1950s (Blanc et al. 1955), may be chronologically correlated to the Colle Curti FU. In this site, Caloi et al. (1979) listed a few rhinoceros remains, including a skull fragment, a fragmentary mandible and some teeth. These remains were ascribed to Dicerorhinus sp. (Rhi- noceros sp. in Blanc et al. 1955) and were later reported as S. hundsheimensis (Caloi & Palombo 1988; Di Ste- fano et al. 1998; Palombo et al. 2002). The remains from Cava Redicicoli have morphological characters similar to S. hundsheimensis, such as the very small dif- ference in height between the bottoms of the valleys in the lower molars (MPUR 1956 R45a, b), the relative- ly large and robust horizontal ramus of the mandible (MPUR 1956 R45), the profile of the occipital crest of the skull (MPUR 1956 s.n.), convex in posterior view and slightly concave in dorsal view (Pandolfi, unpub- lished data). The faunal list of the site includes typical Villafranchian taxa and taxa more frequently recorded in Galerian faunas or closely related to the Galerian ones. Due to the composition of the Redicicoli fauna, Caloi et al. (1979) and Di Stefano et al. (1998) suggest the presence of two different faunal assemblages, which are believed to have come from two different levels. The first level is correlated with the Pirro FU and the second one with the Isernia FU. According to Di Ste- fano et al. (1998) the rhinoceros remains come from this second level. Caloi & Palombo (1995, 1997), Pa- lombo et al. (2002) and Milli & Palombo (2005) suggest that the taxa are coeval and that the Redicicoli faunal assemblage should be ascribed to the Colle Curti FU only. Unfortunately, the fossiliferous deposit does not exist anymore and the problem is difficult to resolve. 180 Pandolfi L. & Petronio C. Imola basin (Imola, northern Italy) Rhinoceros remains discovered in the Imola ba- sin (referred to the Late Villafranchian-Early Galerian) were ascribed to Rhinoceros leptorhinus by Falconer (1868) and to Dicerorhinus cf. etruscus by Azzaroli & Berzi (1972). Later, they were reported as S. cf. hund- sheimensis (Palombo et al. 2002). The upper toothrow, preserved in the Museum of Imola and figured in Fal- coner (1868) and in Azzaroli & Berzi (1972), has a number of common characteristics with S. etruscus and S. hundsheimensis. However, some characters allow to the remains to be ascribed to Falconer’s species, such as the general profile of the vestibular wall in the mo- lars and premolars, the absence of the crista and the presence of a single crochet in P3/, the presence of a crista, a single crochet and a mesial cingulum in P4/, the absence of the antecrochet and the presence of a protocone constriction in M1/, a well developed single crochet and the absence of the antecrochet in M2/ (see also Lacombat 2006). In addition, the upper toothrow shows many similarities in morphology and dimension with S. etruscus from Capitone (Terni, Central Italy) (Fig. 9). Mugello basin (Mugello, central Italy) Specimens with S. hundsheimensis affinities were reported in the Mugello basin; in particular, a deformed and damaged skull from Grezzano (IGF 12728) pre- served in the Museum of Geology and Paleontology of the University of Florence (Mazza 1988). The charac- ters of the teeth and the skull from Grezzano are more similar to Toula’s species, even if the dorsal profile an- gle (n) is more quite similar to that of S. etruscus (n = about 150) than S. hundsheimensis (n >170) (see also Lacombat 2005). Also, in the skull of Grezzano the an- terior border of the orbital cavity overlies the anterior half of M2/ and the posterior border of the narial notch lies at the level of P4/, but these characters are simi- lar to that of the skull of S. etruscus from Capitone (in which the anterior border of the orbital cavity lies at Fig. 5 - Ratio diagrams for MC III from Pirro Nord (FI = pre- served in the Museum of Pal- aeontology of Florence, data from Mazza et al. 1993), Pi- etrafitta (data from Mazza et al. 1993), S. etruscus (S. etru.) (data from Guérin & Heintz 1972; Mazza 1988) and S. hundsheimensis (S. hund.) (data from Fortelius et al. 1993; Kahlke 2001; Lacom- bat 2005) (standard Diceros bicornis). Abbreviations in Tab. 4. Stephanorhinus etruscus from Pirro Nord and other late Early Pleistocene rhinoceros remains of Italy 181 the level of M2/ and the posterior border of the narial notch at the anterior half of P4/). Unfortunately, the remains were found in fluvio-lacustrine deposits with a very long chronological range and they have no direct stratigraphic control. From the same localities, typical Early Galerian mammal remains have been found as well (Masini et al. 1994; Abbazzi et al. 1995). Discussion and Conclusions The proportions, morphology and dimensions of the rhinoceros remains from latest Villafranchian sites in Italy are comparable to those of S. etruscus. Accord- ing to Guèrin (1980), during the Late Villafranchian, this species is inclined towards a general variation in body size and proportions. Guérin (1980) considers the populations of this time span as “second evolutionary stage”. The same proportions and size of the latter are evident in the Capitone sample and in the remains from Senéze, well as in the specimens from Pirro Nord. In Italy, S. etruscus is reported from several Villafranchian sites (Ambrosetti 1972; Ambrosetti & Cremaschi 1975; Mazza 1988; Petronio et al. 2002); according to vari- ous authors, its last occurrence is at the beginning of the Late Villafranchian (Tasso-Olivola FUs; Gliozzi et al. 1997). In the present study, however, the occurrence of the Etruscan rhinoceros is confirmed until the Villa- franchian-Galerian transition. During the latest Villafrachian and earliest Galerian, the presence of S. etruscus has been reported from several European sites, for example Cueva Victo- ria, Incarcal, Venta Micena, Huéscar-1, Cúllar de Baza 1, Atapuerca (Gran Dolina TD4-8 and Sima del Ele- fante TE14), La Sartanette, Blassac-La-Girondie and Peyrolles (Heintz et al. 1974; Cerdeño 1993; van der Made 1998; Rosas et al. 2001; Garcìa-Fernandez et al. 2003a; van der Made et al. 2003; Palombo & Valli 2003; Lacombat 2005) (Fig. 10). Stephanorhinus etruscus was also initially reported from the Spanish sites of Fuente Nueva-3 and Barranco León-5 (1.3-1.1 Ma), where lithic artefacts have been found (Turq et al. 1996; Mar- tínez Navarro et al. 1997; Gibert et al. 1999; Oms et al. 2000). The rhinoceros remains of these sites were later ascribed to S. cf. hundsheimensis (Martínez Navarro et al. 2003). In Europe, the first occurrence of S. hundshe- Fig. 6 - Ratio diagrams for the MC IV from Pirro Nord (FI = preserved in the Museum of Palaeontology of Flor- ence, data from Mazza et al. 1993), Pietrafitta (data from Mazza et al. 1993) versus S. etruscus (S. etru.). (A) (data from Guérin & Heintz 1972; Mazza 1988) and S. hundshe- imensis (S. hund.) (B) (data from Kahlke 2001; Lacombat 2005) (standard Diceros bi- cornis). Abbreviations in Tab. 4. 182 Pandolfi L. & Petronio C. imensis is reported during the latest Early Pleistocene and early Middle Pleistocene. It is present in sites such as Vallonnet, Sainzelles, Tour de Grimaldi, Ceyssaguet, Saint-Prest, Durfort, Untermaßfeld, Dorn-Dürkheim 3 and Trlica (Guérin 1980; Radulescu & Samson 1985; Codrea & Czier 1991; Codrea & Dimitrijević 1997; Franzen et al. 2000; Kahlke 2001; Palombo & Valli 2003; Guérin et al. 2003; Lacombat 2005) (Fig. 10). Re- cently, S. hundsheimensis was reported from the Span- ish site of Vallparadís (1-0.800 Ma) (Alba et al. 2008). Thus, during the latest Early Pleistocene – earli- est Middle Pleistocene, S. etruscus and S. hundsheimen- sis seem to be coeval in Europe. However, in agreement with van der Made (2000), the typical early Middle Pleistocene S. hund- sheimensis appears to be more primitive than the Early Pleistocene Stephanorhinus; this last is represented by an evolutionary stage of S. etruscus (second evolution- ary stage). S. hundsheimensis may have been an immigrant that replaced S. etruscus at the end of the Early Pleis- tocene. This is in line with the model of Fortelius et al. (1993), which does not derive S. hundsheimensis from S. etruscus, rather than with Guérin’s (1980) model. Stephanorhinus hundsheimensis probably reached Europe from Asia between 1.3-1.1 Ma and gradually replaced S. etruscus; at the same time, populations of the latter species were fragmented into isolated demes. In Italy, the occurrence of S. hundsheimensis in the Late Villafranchian sites (Pirro Nord, Madonna della Strada) is not considered reliable at present time. In the lignitiferous deposit of Pietrafitta (Perugia, Cen- tral Italy), chronologically earlier than Pirro Nord site (Farneta FU), several rhinoceros remains were found as early as 1900. At first, Ugolini (1921) ascribed some teeth to Rhinoceros etruscus and later, other rhinoceros remains were ascribed to Dicerorhinus etruscus (Am- brosetti et al. 1987) and S. cf. hundsheimensis (Mazza et al. 1993). Some morphological and morphometrical considerations of the rhinoceros remains from Pie- Fig. 7 - Ratio diagrams for MT IV from Pirro Nord (data from Mazza et al. 1993), Pietrafit- ta (data from Mazza et al. 1993). S. etruscus (S. etru.) (data from Guérin & Heintz 1972; Mazza 1988) and S. hundsheimensis (S. hund.) (data from Fortelius et al. 1993; Kahlke 2001; Lacom- bat 2005) (standard Diceros bicornis). Abbreviations in Tab. 4. Stephanorhinus etruscus from Pirro Nord and other late Early Pleistocene rhinoceros remains of Italy 183 trafitta, listed by Mazza et al. (1993), are reported as evolutionary trends of S. etruscus by Guérin (1980: 599-603). Furthermore, the small late Early Pleistocene rhinoceroses, very likely identical to that was assigned to S. cf. hundsheimensis by Mazza et al. (1993), are re- ported from Spain as S. etruscus (Cerdeño 1993; van der Made 1998). However, the remains from Pietrafitta are proportionally more similar to those of S. etruscus than S. hundsheimensis (for example see Figs 6-8). Be- sides this, some morphological and metrical differences in the Pietrafitta remains may fall within the intraspe- cific range of variation of the Etruscan rhinoceros; they may also be the result of adaptation to a particular hab- itat (see also García-Fernandez et al. 2001). However, a more detailed analysis is needed. Other rhinoceros remains discovered in Ital- ian sites chronologically related to the Villafranchian- Galerian transition (Mugello basin) have close affini- ties with S. etruscus. Also, remains attributable to S. hundsheimensis from the earliest Galerian sites (Colle Curti, Castagnone) are uncertain and scarce, and no more data is available on the origin of the remains of Cava Redicicoli. Thus, S. hundsheimensis seems to be present in Italy only after 1 Ma (Slivia FU). This spe- cies is certainly present during the Middle Galerian (Middle Pleistocene). It is found in Cesi (about 0.7 Ma) (Mazza 1996; Ficcarelli et al. 1997), in the lower levels of Ponte Galeria (about 0.8 Ma) (Petronio 1988; Petro- nio & Sardella 1999; Pandolfi, unpublished data) and in Isernia La Pineta (about 0.6-0.55 Ma) (Sala & Fortelius 1993; Coltorti et al. 2005). The dispersal event of S. hundsheimensis into Europe and Italy may be related to the climatic dete- rioration of the latest Early Pleistocene. The same may be true of the variations in proportions and size of S. etruscus. In fact, during this time, the passage to a phase characterized by long glaciations with 100.000-year cy- cles occurs. The difference between the temperatures Fig. 8 - Ratio diagram for MT II from Pirro Nord (data from Mazza et al. 1993), Pietrafit- ta (data from Mazza et al. 1993). S. etruscus (S. etru.) (data from Mazza 1988) and S. hundsheimensis (S. hund.) (data from Kahlke 2001; Lacombat 2005) (standard Diceros bicornis). Abbrevia- tions in Tab. 4. 184 Pandolfi L. & Petronio C. during glacial and interglacial phases becomes more pronounced (Leroy 2007). In Italy, more arid condi- tions, during the Pirro FU, are testified to by the oc- currences of Bison degiulii and a lightly build, medi- um-sized horse, Equus altidens. Also, typical African taxa, such as Hystrix refossa, Megantereon whitei and Theropithecus sp. disperse on the Peninsula (Petronio et al. in press). Concurrently, the disappearances of Leptobos and Eucladoceros were recorded. However, at Pirro Nord, woody areas and humid patches must have been still present at some point, as indicated by the presence of Apodemus flavicollis, as well as the presence of amphibians and insectivores. At other sites related to the Pirro FU, in which S. etruscus was dis- covered, lacustrine basins or ponds were present (for example in the Scoppito site; Maccagno 1962). Later, a very arid-steppic phase is recorded dur- ing the Colle Curti FU. In the Ranica site in northern Italy, a cold, steppic phase with Cervalces latifrons is well documented (Ravazzi et al. 2005). Nevertheless, trees remain present at the foot of the Alps even dur- ing this phase. In contrast, in central Italy, the pollen diagrams of Colle Curti and Cesi (Bertini 2000) show high percentages of herbaceous forms, a clear evidence of more open vegetation. Among the mammals, pres- ence of a cold microfauna represented by Prolagurus pannonicus and Predicrostonyx sp. is reported in the blue-clay layer of the Ponte Galeria area (Roma, cen- tral Italy) (Kotsakis et al. 1992). These finds testify a very cold phase, correlated with the Marine Isotopic Stage 24-22. During this time, S. etruscus disappeared from Italy and the occurrence of S. hundsheimensis is reported. Other Asian taxa, such as Praemegaceros verticornis and Bison schoetensacki, co-occur with S. hundsheimensis. The diffusion of S. hundsheimensis can be also related to a greater capacity of the species to adapt to the climatic conditions and diet (with highly flex- ible feeding, Kahlke & Kaiser in press) than S. etrus- cus. Moreover, the distribution of the latter seems to be more influenced by humidity (Guérin 1980). In Italy, the absence of remains definitely attrib- utable to S. hundsheimensis before 1 Ma may be due to the conformation of the Italian Peninsula. This may have played a decisive role in the delay of the disper- sal event when compared to the rest of continental Eu- rope. In fact, delayed dispersal events are common in Fig. 9 - Upper toothrows of S. etruscus from the Imola ba- sin (A) (after Azzaroli & Berzi, 1972; not in scale) and Capitone (B) (scale bar: 5 cm). Fig. 10 - Location map of selected late Early Pleistocene sites of Europe and Italy with remains of S. etruscus and S. hund- sheimensis. Sites with S. etruscus: 1 = Pirro Nord (1.3-1.6 Ma); 2 = Scoppito (1.3-1.1? Ma); 3 = Imola basin (latest Early Pleis- tocene); 6 = Lézignan-le-Cèbe (about 1.5 Ma); 8 = Ata- puerca Gran Dolina TD4-8 (about 0.800 Ma) and Sima del Elefante TE14 (1.4-1.1 Ma); 9 = Cueva Victoria (about 1 Ma); Sites with S. hundsheimensis: 4 = Vallonnet (about 1 Ma); 5 = Soleilhac (about 0.750 Ma); 7 = Fuente Nueva-3 and Bar- ranco León-5 (1.3-1.1 Ma); 10 = Dorn-Dürkheim 3 (about 0.800 Ma); 11 = Untermaßfeld (about 1 Ma); 12 = Trlica (latest Early Pleistocene). Stephanorhinus etruscus from Pirro Nord and other late Early Pleistocene rhinoceros remains of Italy 185 Italy during the Pleistocene, in particular for species that arrive from Asia (for example Coelodonta antiqui- tatis is not present before MIS 4 and Praemegaceros so- lilhacus is present during the Isernia FU, about 0.600- 0.550 Ma) (Petronio et al. in press). Some African or southwest Asian taxa seem instead to be present first in Italy and later in central Europe. This is the case of Dama clactoniana and perhaps Homo. 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