Microsoft Word - 00_indice_LM04.docx Il Quaternario IT ISSN 039-3356 Italian Journal of Quaternary Sciences 24 (1), 2011 - 103-112 STEPHANORHINUS KIRCHBERGENSIS (JÄGER, 1839) FROM THE MIDDLE PLEISTOCENE DEPOSIT OF RIANO (ROME, CENTRAL ITALY) Luca Pandolfi Dipartimento Scienze della Terra, Università di Roma Sapienza, Roma Correspondent author: ABSTRACT: Pandolfi L., Stephanorhinus kirchbergensis (Jäger, 1839) from the Middle Pleistocene deposit of Riano (Rome, central Italy). In this work the rhinoceros remains discovered in the site of Riano are analyzed. The remains are preserved in Museum of Paleontology of University “Sapienza” of Rome and they have been recovered in a diatomaceous deposit together with remains of Elephas antiquus, Cervus elaphus rianensis and Dama clactoniana. The faunal assemblage of Riano can be correlated with Torre in Pietra Faunal Unit on the base of fossil mammal fauna and stratigraphical data. Rhinoceros remains belong to only one individual and consist of various post- cranial elements. Morphological and morphometrical characters of the remains are comparable with Eurasian rhinoceros S. kirchbergen- sis. This species has been rarely recorded, although the distribution range has been relatively wide. S. kirchbergensis has been discov- ered in a large part of Europe and in the Early and Middle Pleistocene deposits of central Asia, where the species has probably origin. Riano is one of the few European localities where post-cranial remains of S. kirchbergensis are reported. RIASSUNTO: Pandolfi L., Stephanorhinus kirchbergensis (Jäger, 1839) dal deposito del Pleistocene Medio di Riano (Roma, Italia centrale). In questo lavoro vengono analizzati i resti di rinoceronte rinvenuti nel bacino diatomitico di Riano e conservati presso il Museo di Paleon- tologia dell’ Università “Sapienza” di Roma. La fauna a grandi mammiferi, rinvenuta negli stessi depositi da cui proviene il rinoceronte, è composta da Elephas antiquus, Cervus elaphus rianensis e Dama clactoniana; il sito è correlabile, per composizione faunistica e strati- grafia, con l’Unità Faunistica di Torre in Pietra. I resti di rinoceronte, composti da diverse parti dello scheletro post-craniale e appartenen- ti ad un unico individuo, presentano caratteristiche morfologiche e morfometriche comparabili con la specie euroasiatica S. kirchbergen- sis. Questa specie, nonostante occupasse un vasto areale, è relativamente rara e probabilmente ha origine in Asia centrale, da dove provengono le segnalazioni più antiche riferite al Pleistocene Inferiore. La presenza di S. kirchbergensis nel sito di Riano rappresenta una delle poche in Europa in cui vengono descritti resti dello scheletro post-craniale di questa specie, prevalentemente conosciuta sulla base di denti isolati. Key Words: Stephanorhinus kirchbergensis, Middle Pleistocene, Riano. Parole-chiave: Stephanorhinus kirchbergensis, Pleistocene Medio, Riano. 1. INTRODUCTION The site of Riano is situated 20 km north of Rome and is characterized by tufaceous-diatomitic Formation. This last is lying on the “yellow vacuolized tuff” of the Pleistocene tufaceous series of “Vulcano Sabatino” (AMBROSETTI et al., 1969, 1972; ACCORDI & MACCAGNO, 1962; BONADONNA & BIGAZZI, 1969). In the past century, from the diatomaceous levels, well-preserved and al- most complete vertebrate skeletons have been discov- ered. Among the others, the fossil fauna is represented by Elephas antiquus FALCONER & CAUTLEY, Cervus ela- phus rianensis LEONARDI & PETRONIO, Dama clactonia- na (FALCONER), several fishes, amphibians and reptiles (MACCAGNO, 1962; ACCORDI & MACCAGNO, 1962; MA- STRORILLI, 1965; LEONARDI & PETRONIO, 1974; CALOI et al., 1980). The fossil flora is well represented by several forest assemblages (FOLLIERI, 1958a; 1958b; 1962) that allow to well-reconstruct the climatic conditions near Riano during the Middle Pleistocene. The landscape around the site was characterized by a dense forest, with a remarkable biodiversity compared to coeval records of vegetation in central and northern Europe. The modern analog of this landscape may be found in areas with over 2000 mm mean annual precipitation, favouring the development of such a dense forest. In particular, three forest phases are recognised: a mixed oak forest at the basis, followed by a period with domi- nant Pterocarya KUNTH, and by a final phase with a pro- gressive increase of Abies MILLER and Fagus LINNAEUS pollen and the decrease of that of Pterocarya (AMBRO- SETTI & BONADONNA, 1967; FOLLIERI, 1962; FOLLIERI & MAGRI, 2001). AMBROSETTI et al. (1972) and BONADONNA & BI- GAZZI (1969) proposed, for the diatomaceous deposit of Riano, the Formation name ‘Rianian’ to cover the late Middle Pleistocene in the stratigraphical scheme for the Tyrrhenian margin in central Italy. On the basis of strati- graphical relationships and faunal characteristics, MA- LATESTA (1978) regarded these deposits as strictly representing the continental facies equivalent to the flu- vio-lacustrine-brackish-marine sediments that form the Aurelia Formation and, according to CALOI et al. (1998), this deposit can be correlated with Torre in Pietra Faun- al Unit (sensu GLIOZZI et al., 1997). L. Pandolfi 104 2. MATERIALS AND METHODS The nomenclature used in this work was set by FORTELIUS et al., (1993). Also, according to GROVES (1983) the Pleistocene European rhinoceroses show little similarity with the genus Dicerorhinus GLOGER and they are ascribed to genus Stephanorhinus KRETZOI, as previously used by several authors (HEISSING, 1973, 1981, 1989; CERDEÑO, 1993, 1995, 1998; PROTHERO et al., 1986; inter alios). Rhinoceros remains found in the site of Riano are preserved in the Museum of Paleontology of “Sapienza” University of Roma (MPUR). Some of the remains are unpublished while some of them are previously ascribed to Dicerorhinus cf. mercki KAUP (LEONARDI & PETRONIO, 1974; GUÉRIN, 1980). Later, the rhinoceros of Riano has been reported in literature as Dicerorhinus cf. hemitoe- chus FALCONER, Stephanorhinus sp., Stephanorhinus cf. hemitoechus (CALOI et al., 1980, CALOI & PALOMBO, 1994; PALOMBO et al., 2002). ). The remains analyzed have been found in the ’70ties of the past century in the Cave of diatomaceous earth of Valle dell’Inferno (Riano) (PE- TRONIO, pers. com.). In the same Cave, in the ’60ties of the past century, the skeletons of Cervus elaphus rianen- sis have been discovered. The rhinoceros remains belong to the same individual and some of them have been found articulated. Also, the rhinoceros lacks of the cranial portion and preserves several post-cranial elements among the os coxae, femurs, calcaneum, talus. The most of post-cranial elements are damaged or deformed by characteristics of the fossiliferous site, therefore some morphometrical characters can not be taken into account. Morphological characters considered are those listed as diagnostics by several authors, including GUÉRIN (1980), FORTELIUS et al. (1993), LACOMBAT (2005). When possible, the morphometric methods are based on the works of GUÉRIN (1980), MAZZA (1988), FORTELIUS et al. (1993), LACOMBAT (2005). 3. SPECIMENS STUDIED AND COMPARATIONS The rhinoceros remains from Riano are represented by several elements of post-cranial skeleton and frag- ments of bones. The principal elements of the skeleton are reported and described below. 3.1 Vertebrae The vertebral column is represented by several thoracic and lumbar vertebrae (MPUR 7/1-7/16) in a dif- ferent state of preservation (Fig. 1). Some of them are articulated with other vertebrae and ribs (MPUR 7/21- 7/25). Also, among the vertebrae is present the proximal portion of an atlas, very damaged. Due to the deformation and damage of the remains, no morphological characters or morphometric values can be taken into account. 3.2 Scapulae (MPUR 7/54) Two scapulae are present, one right and one left, articulated with the ribs. The scapulae are compressed latero-medially and are very damaged (Fig. 1). The left scapula preserved only the proximal portion, while, the right one preserved the proximal portion (depth of the proximal portion > 212 mm) and the spine. No diagnos- tic character is present in the preserved portions. 3.3 Humeri Two humeri are present; one of them preserved only the diaphysis (MPUR 7/69) and a portion of the proximal articular surface (MPUR s.n.). The other one preserved the diaphysis and a very damaged proximal epiphysis (MPUR 7/56) (Fig. 1). The dimensions of the diaphysis (breadth = about 65 mm; depth = > 65 mm) seem not to be helpful to distinguish between the differ- ent rhinoceros species. In fact, the dimensional ranges of the diaphysis of the European Pleistocene rhinoce- roses are quite similar (cfr. GUÉRIN, 1980). The proximal articular surface is almost intact and have a circular outline. The approximate dimensions (breadth = > 80 mm; depth = > 90 mm) allow to exclude that it can be ascribed to S. etruscus FALCONER (cfr. FORTELIUS et al., 1993). 3.4 Sacrum (MPUR 7/20) The sacrum (maximal breadth = about 167 mm; maximal length = > 165 mm) is articulated with some lumbar vertebrae (MPUR 7/17-7/19) (Fig. 1). No morpho- logical or morphometrical study has been done on the sa- crum of European Pleistocene rhinoceroses. Considering that, the comparison with other specimens is very difficult. 3.5 Os coxae Several fragmentary portions of the os coxae are present (MPUR 7/59-7/65). Also, the left os coxae is almost intact (MPUR 7/57) and the articular surface of the right os coxae is present (MPUR 7/58). The aceta- bulum is relatively broad, deep and sub-elliptical, com- pressed dorsally (Fig. 1). This shape seems not to be present in S. etruscus, S. hundsheimensis TOULA and S. hemitoechus (cfr. LACOMBAT, 2005); it is more similar to S. kirchbergensis JÄGER from Taubach (cfr. PORTIS, 1878). Also, posteriorly, the acetabulum is separated into two articular lips; a dorsal and a ventral one. The dorsal lip is inclined posterior-ventrally and is much wid- er and longer that the ventral one (Fig. 1). These cha- racters allow to distinguish this bone from those of S. etruscus, S. hundsheimensis and S. hemitoechus (cfr. LACOMBAT, 2005), and seem to be present in S. kir- chbergensis from Taubach (cfr. PORTIS, 1878). Also, unlike S. hemitoechus, in the os coxae from Riano the pit of acetabulum is well-evident (cfr. LACOMBAT, 2005). The dimensions of the acetabulum are much larg- er than S. etruscus from Upper Valdarno. The value of the maximal length of the acetabulum is much greater than S. hemitoechus and it is more similar to the max- imal values of S. hundsheimensis (Tab. 1). Also, the values of the diaphysis of the Ilium and the height of the acetabulum appear relatively large in the specimens of Riano (Tab. 1). 3.6 Femurs The two distal epiphysis of the femurs (MPUR 7/66; MPUR 7/68) and a damaged portion of the right proximal epiphysis are preserved (MPUR 7/70) (Fig. 2). The proximal articular surface is partially damaged in the medial portion and seems to be slightly compressed dorso-ventrally (Fig. 2). In proximal view, it seems to be much developed and much compressed latero-medially compared with S. hundsheimensis from Mauer (from Fortelius et al., 1993) and Ponte Molle. The articular sur- faces of the distal epiphysis of the femurs, in anterior- Stephanorhinus kirchbergensis (Jäger, 1839) from the Middle Pleistocene deposit of Riano … 105 Fig. 1 - Rhinoceros remains from Riano: 1) Scapula, lateral view; 2) Humerus, posterior view; 3) Os coxae, a-dorsal view, b-lateral view of articular surface; 4) Sacrum, ventral view; 5) Thoracic vertebrae, right lateral view. The bar is of about 2 cm. Resti di rinoceronte di Riano: 1) Scapola, visione laterale; 2) Omero, visione posteriore; 3) Bacino, a-visione dorsale, b-visione laterale della superficie articolare; 4) Sacro, visione ventrale; 5) Vertebre toraciche, visione laterale destra. La barra è di circa 2 cm. Fig. 2 - Rhinoceros remains from Riano: 1) Femur, proximal epiphysis, proximal view; 2) Femur, distal epiphysis, a-anterior-distal view, b-posterior view; 3) Patella, lateral view; 4) Tibia, proximal view; 5) Talus, anterior view; 6) Calcaneum, a-medial view, b-distal view; 7) Metapodial bone articulated with first phalanx, lateral view. The bar is of about 2 cm. Resti di rinoceronte di Riano: 1) Femore, epifisi prossimale, visione prossimale; 2) Femore, epifisi distale, a-visione anteriore, b-visione posteriore; 3) Rotula, visione laterale; 4) Tibia, visione prossimale; 5) Astragalo, visione anteriore; 6) Calcagno, a-visione mediale, b- visione distale; 7) Metapodiale in connessione anatomica con la prima falange, visione laterale. La barra è di circa 2 cm. L. Pandolfi 106 Tab. 1 - Comparative dimensions of os coxae from Riano and of S. etruscus (S. etru.) (data from MAZZA, 1988), S. hemitoechus (S. hem.) and S. hundsheimensis (S. hund.) from Isernia (data from LACOMBAT, 2005). La = outer length of the acetabulum; Lar = inner length of the acetabulum; lar = inner width of the acetabulum; Lo = length of the foramen obturatum; BsI = breadth of the shaft of the ilium; His = height of the shaft of the ilium; Ha = height of the acetabulum; LL = lateral length; ca = about. Dimensioni comparative del bacino di Riano e di S. etruscus (S. etru.) (dati da MAZZA, 1988), S. hemitoechus (S. hem.) e S. hundshei- mensis (S. hund.) di Isernia (dati da LACOMBAT, 2005). La = lunghezza esterna dell’acetabolo; Lar = lunghezza interna dell’acetabolo; lar = larghezza interna dell’acetabolo; Lo = lunghezza del foramen obturatum; BsI = ampiezza della diafisi dell’ilium; His = altezza della dia- fisi dell’ilium; Ha = altezza dell’acetabolo; LL = lunghezza laterale; ca = circa. Tab. 2 - Comparative dimensions of femurs from Riano and of S. etruscus (S. etru.), S. hemitoechus (S .hem.), S. hundsheimensis (S. hund.) and S. kirchbergensis (S. kirch.) (1 = data from ForTelius et al., 1993; 2 = data from LACOMBAT, 2005; 3 = data from KAHLKE, 1977). DTap = breadth of the proximal epiphysis; DAPap = depth of the proximal epiphysis; DTD = breadth of the distal epiphysis; DAPDM = medial depth of the distal epiphysis; DAPDL = lateral depth of the distal epiphysis; DTtr = breadth of the distal trochlea; DTcon = breadth of the posterior condyles; DTfc = breadth of the intercondyloid fossa. Dimensioni comparative dei femori di Riano e di S. etruscus (S. etru.), S. hemitoechus (S. hem.), S. hundsheimensis (S. hund.) e S. kir- chbergensis (S. kirch.) (1 = dati da FORTELIUS et al., 1993; 2 = dati da LACOMBAT, 2005; 3 = dati da KAHLKE, 1977). DTap = diametro tra- sversale dell’epifisi prossimale; DAPap = spessore dell’epifisi prossimale; DTD = diametro trasversale dell’epifisi distale; DAPDM = spessore mediale dell’epifisi distale; DAPDL = spessore laterale dell’epifisi distale; DTtr = diametro trasversale della troclea distale; DTcon = diametro trasversale dei condili posteriori; DTfc = diametro trasversale della fossa intercondilea. Stephanorhinus kirchbergensis (Jäger, 1839) from the Middle Pleistocene deposit of Riano … 107 distal view, are undamaged and well- developed (Fig. 2). The trochlea is clearly asymmetric and the medial lip is much larger and developed than the lateral one. The trochlear trough is broad and deep. In medial view, the surface of the bone is broad and the tuberosity is well-evident and massive. It is situated slightly higher than the lateral one. In lateral view, the surface of the bone is less broad than the medial one and the tuberosity is much rounded (Fig. 2). This morphology can be observable in the distal epiphysis of S. kir- chbergensis from Taubach (cfr. KAHLKE, 1977) and Mosbach (cfr. FORTELIUS et al., 1993) and it is different from those of the other species of the Stephanorhinus genre. These last have a different development and shape of the medial and lateral tube- rosities and of the articular surfaces (GUÉRIN, 1980; FORTELIUS et al., 1993; LACOMBAT, 2005). In posterior view, the condyles are well-developed and rounded; the intercondyloid fossa is strong, broad and deep (Fig. 2). The medial condyle is much broader than the lateral one; this last is postero-laterally inclined. Even these characters seem to be distinctive of S. kirchbergensis and allow to distinguish it from the other species of the Stephanorhinus genre (GUÉRIN, 1980; FORTELIUS et al., 1993; LACOMBAT, 2005). At last, the dimensions of the femurs of Riano are relatively large. They are comparable with the maximal values of S. hemitoechus and the values of S. kir- chbergensis (Tab. 2; Tab. 7). 3.7 Patellae (MPUR 7/72; MPUR 7/71) Two patellae are preserved, one right and one left, differently damaged (Fig. 2). In posterior view, two articular surfaces are present, a medial one, broad and concave, and a lateral one, less broad and concave than the former. The two articular surfaces are well-separated by a saddle. The di- mensions of the bones are relatively large if compared with those of S. etruscus, S. hundsheimensis and S. hemitoechus re- ported by FORTELIUS et al. (1993) and LA- COMBAT (2003) (Tab. 3); in particular, the DTmax is much greater. This last character is peculiar of S. kirchbergensis (GUÉRIN, 1980) (Tab. 7). 3.8 Tibia (MPUR 7/73) Only one tibia is present. This last is very damaged and preserved the proximal epiphysis and a portion of the diaphysis. The tibia is massive, the anterior tuberosity is damaged and deformed, the tibial spine is quite salient and the interspine fossa is relatively broad (Fig. 2). The lateral rim of the tibial spine is higher than the medial Tab. 3 - Comparative dimensions of patellae from Riano and of S. etruscus (S. etru.), S. hemitoechus (S. hem) S. hundsheimensis (S. hund.) and S. kirchbergensis (S. kirch.) (1 = data from FORTELIUS et al., 1993; 2 = data from LACOMBAT, 2005; 3 = data from KAHLKE, 1977). Lmax = maximal lenght; La = articular lenght; DT max = maxim- al breadth; DTa = articular breadth; DAPmax = maximal depth. Dimensioni comparative delle rotule di Riano e of S. etruscus (S. etru.), S. hemitoe- chus (S. hem) S. hundsheimensis (S. hund.) e S. kirchbergensis (S. kirch.) (1 = dati da FORTELIUS et al., 1993; 2 = dati da LACOMBAT, 2005; 3 = dati da KAHLKE, 1977). Lmax = lunghezza massima; La = lunghezza articolare; DT max = diametro trasver- sale massimo; DTa = diametro trasversale articolare; DAPmax = diametro antero- posteriore massimo. Tab. 4 - Comparative dimensions of talus from Riano and of S. etruscus (S. etru.), S. hemitoechus (S .hem.) S. hundsheimensis (S. hund.) and S. kirchbergensis (S. kirch.) (1 = data from FORTELIUS et al., 1993; 2 = data from LACOMBAT, 2005; 3 = data from KAHLKE, 1977). DTmax = maximal breadth; DAPl = lateral depth; Htrl = height of the lateral trochlea; Hl = lateral height. Dimensioni comparative dell’astragalo di Riano e di S. etruscus (S. etru.), S. hemito- echus (S. hem.), S. hundsheimensis (S. hund.) e S. kirchbergensis (S. kirch.) (1 = dati da FORTELIUS et al., 1993; 2 = dati da LACOMBAT, 2005; 3 = dati da KAHLKE, 1977). DTmax = diametro trasversale massimo; DAPl = diametro antero-posteriore laterale; Htrl = altezza della troclea laterale; Hl = altezza laterale. L. Pandolfi 108 one. The tibial fossa, in dorsal view, seems to be rela- tively wide; however, this last character may have been accentuated by the deformation. The medial articular surface of the proximal epiphysis is more antero- posteriorly elongated than the lateral one. The state of preservation of the tibia does not al- low to carry out an exhaustive morphometrical work. The relative dimension of the transversal diameter of the proximal epiphysis (DTp = > 129 mm) seems to be larg- er than S. etruscus and S. hundsheimensis (respectively DTp = 87÷110 mm and DTp = 111,5÷119 mm in FORTE- LIUS et al., 1993; respectively DTp = 102÷115 mm and DTp = 103÷123,5 mm in GUÉRIN, 1980) and is closer to maximal values of S. hemitoechus (DTp = 101÷137,5 mm in GUÉRIN, 1980) and to the values of S. kirchber- gensis reported by GUÉRIN (1980) (DTp = 136 mm) and KAHLKE (1977) (DTp = 122,5). 3.9 Talus (MPUR 7/76) The talus (Fig. 2) is relatively damaged; in anterior view, the trochlea lacks of the medial portion; in lateral view, the articular surface is relatively wide and is post- eriorly limited by a slight depression. In medial view, the distal tuberosity is well-developed and is situated in the anterior-distal margin of the medial face, unlike S. hemi- toechus (GUÉRIN, 1980; LACOMBAT, 2005). In posterior view, the proximo-lateral articular surface is well- developed, concave and sub-quadrangular in shape; it is elongated in the latero-distal side with a smaller arti- cular facet. The mesio-distal articular surface have sub- trapezoidal shape; it is well-separated from the proximo- lateral one and is relatively damaged. The maximal transverse diameter is greater than S. etruscus and S. hundsheimensis. The dimensions of the bone are comparable with the maximal values of S. hemitoechus and fall in the variation range of S. kirchber- gensis (Tab. 4; Tab. 7). 3.10 Calcaneum (MPUR 7/75) In lateral view, the calcaneum (Fig. 2) is slightly antero-posteriorly compressed; on the whole, the bone is massive and di- mensionally large. The tuber calcanei is relatively large and high; it has an antero- posterior development similar to the beak. This last character allow to distinguish this bone from those of S. etruscus and S. hundsheimensis (GUÉRIN, 1973; 1980; LA- COMBAT, 2005). The sustentaculum talii is about per- pendicular to the axis of the bone and it is slightly damaged in the medial portion. In antero-distal view, the articular surfaces are well-evident. The upper lateral articular sur- face has rounded shape; the articular sur- face on the sustentaculum talii has a clear outline and has sub-circular shape. These last articular surfaces are clearly well- separated between them. The lower articular surface is slightly damaged and more higher than long. The articular surface with the cu- boid is flat and antero-posteriorly elon- gated. All these morphological characters are comparable with S. kirchbergensis and allow to distinguish from S. hemitoechus. In this last species the articular surfaces with the talus are not well-separated between them and the lower articular surface is more longer than high (cfr. GUÉRIN, 1973; 1980; KAHLKE, 1977). Also, the dimensions of the calcaneum are relatively great and are comparable with the values of S. kirch- bergensis (Tab. 5; Tab. 7). 3.11 Cuboids (MPUR s.n.) The cuboids, one right and one left, are incomplete and differently damaged. The anterior face of the bone is sub-trapezoidal in shape and the lateral side is more high- er than the medial one. The lower margin of the anterior face is about straight, while the upper margin is clearly ob- lique. In proximal view, the articular surface seems to be flat in the anterior-lateral side and oblique in the medial side. No other morphological character is observable and the few characters present do not allow to distinguish among the species (cfr. FORTELIUS et al., 1993; LACOM- BAT, 2005). The dimensions of the cuboids from Riano are comparable with the values of S. kirchbergensis and the maximal values of S. hemitoechus (Tab. 6). 3.12 Great cuneiform (MPUR s.n.) Only one cuneiform is preserved. It is very damaged and deformed. In lateral view, two articular surfaces are visible. The proximo-medial articular surface has sub- triangular shape, while the distal one is elongated. Due to the deformation of the bone, no other morphological or morphometrical character is recognizable. Tab. 5 - Comparative dimensions of calcaneum from Riano and of S. etruscus (S. etru.), S. hemitoechus (S .hem.), S. hundsheimensis (S. hund.) and S. kirchbergen- sis (S. kirch.) (1 = data from FORTELIUS et al., 1993; 2 = data from LACOMBAT, 2005; 3 = data from KAHLKE, 1975; 1977). Hmax = maximal height; DTs = breadth of the tu- ber calcanei; DAPs = depth of the tuber calcanei; DTmp = minimum posterior breadth; DTst = breadth of the sustentaculum talii; DAPb = depth of the beak. Dimensioni comparative del calcagno di Riano e di S. etruscus (S. etru.), S. hemitoe- chus (S. hem.), S. hundsheimensis (S. hund.) e S. kirchbergensis (S. kirch.) (1 = dati da FORTELIUS et al., 1993; 2 = dati da LACOMBAT, 2005; 3 = dati da KAHLKE, 1975; 1977). Hmax = altezza massima; DTs = diametro trasversale del tuber calcanei; DAPs = diametro antero-posteriore del tuber calcanei; DTmp = diametro trasversale minimo posteriore; DTst = diametro trasversale del sustentaculum talii; DAPb = dia- metro antero posteriore del becco. Stephanorhinus kirchbergensis (Jäger, 1839) from the Middle Pleistocene deposit of Riano … 109 Tab. 6 - Comparative dimensions of cuboids from Riano and of S. etruscus (S. etru.), S. hemitoechus (S .hem.), S. hund- sheimensis (S. hund.) and S. kirchbergensis (S. kirch.) (0 = data from GUÉRIN, 1980; 1 = data from FORTELIUS et al., 1993; 2 = data from LACOMBAT, 2005). H ant. = anterior height; l = breadth. Dimensioni comparative dei cuboidi di Riano e di S. etruscus (S. etru.), S. hemitoechus (S. hem.), S. hundsheimensis (S. hund.) e S. kirchbergensis (S. kirch.) (0 = dati da GUÉRIN, 1980; 1 = dati da FORTELIUS et al., 1993; 2 = dati da LACOMBAT, 2005). H ant. = altezza anteriore; l = larghezza. 3.13 Metapodial bone (MPUR s.n.) Very fragmentary metapodial bone is preserved. It consists of a distal epiphysis articulated with the first and second phalanges (Fig. 2). No morphological or morphometrical character are recognizable. 4. DISCUSSION AND CONCLUSIONS Even if partially damaged, the rhinoceros re- mains from Riano preserve some characters, such as the articular surfaces, which allow an accurate analy- sis. In particular, the shape and size of the articular surfaces and tuberosities of the femurs, the characters of the os coxae and the morphology of the calcaneum of the rhinoceros from Riano are comparable with S. kirchbergensis (cfr. GUÉRIN, 1973; 1980; FORTELIUS et al., 1993). Moreover, the size of the remains fall into the vari- ation range know for S. kirchbergensis (cfr. KAHLKE, 1977; GUÉRIN, 1980; FORTELIUS et al., 1993). S. kirchbergensis seems to be a relatively rare species, although it is reported in most of Europe and Asia (GUÉRIN, 1980; TONG, 2001; LACOMBAT, 2005; BILLIA, 2008; BILLIA & PETRONIO, 2009). This rarity could be related with the ethology of the species; probably, it was present in the area with groups of few individuals. However, it can be assumed that the spe- cies living in ecological niche unfavourable for its fossi- lization (LOOSE, 1975). In this regards, the species seems to be typical of forest environments; this hypo- thesis is supported by the brachydont teeth of the spe- cies and by the remains of flora and fauna that have been discovered with this species (GUÉRIN, 1980). Re- cently, S. kirchbergensis has been found in sites of southern China, 2000 m above sea level, that corres- pond to temperate deciduous and coniferous forest (TONG & WU, 2009). The fossil flora of Riano, discov- ered with the rhinoceros remains, bear out the hypo- thesis that species could be typical of forest environ- ments. This fact could be explain the relatively ab- sence or rarity of the species in southern Italy, in the Iberian Peninsula and in North Africa, unlike to the coeval European species S. hemitoechus (GUÉRIN, 1980; CERDEÑO, 1990; BILLIA & PETRONIO, 2009). The phylogenetics of S. kirchbergensis is still un- resolved. This species is directly connected to Lartero- therium megarhinus (DE CHRISTOL) by WÜRST (1922), STAESCHE (1941) and GUÉRIN (1980), even if the evolu- tionary connections between the two species are not clear much. However, S. kirchbergensis seems to has origin in central Asia. In fact, it is reported in Early Pleistocene sites of China and Kazakhstan (KHISARO- VA, 1963; DONG et al. 2000; TONG & MOIGNE, 2000; TONG, 2001; BILLIA & PETRONIO, 2009; TONG & WU, 2009). Later S. kirchbergensis is present in the Middle Pleistocene deposits of Russia and Eastern Europe (CZYŻEWSKA, 1962; BORSUK-BIALYŃICKA & JAKUBOWSKI, 1972; BILLIA & SHPANSKY, 2005; BILLIA, 2008). The species may be evolved in central Asia from a common ancestor of the evolutionary lineage that led to S. hemi- toechus in Europe. Later, it would spread westward during the Middle Pleistocene. In this regards, an ex- haustive analysis of Mio-Pliocene Asian rhinoceros re- mains would confirm this hypothesis. In Italy, the oldest remains ascribed to S. kirchber- gensis are those discovered in the Middle Pleistocene sites of Valdemino (NOCCHI & SALA, 1997) and Ponte Molle (CAPASSO BARBATO et al., 1998; BILLIA & PETRO- NIO, 2009), both correlable with Isernia FU (sensu GLIOZZI et al., 1997). Also, the species is reported in few sites of late Middle Pleistocene of northern and central Italy and it seems to be absent in the second part of Late Pleistocene (PETRONIO et al., 2007). How- ever, the most part of the remains ascribed to this spe- cies consist of isolated teeth. The remains of Riano are, instead, one of the rare record in Europe of S. kir- chbergensis based on post-cranial elements belonging to a single individual. L. Pandolfi 110 Tab. 7 - Comparative dimensions of femurs, patellae, talus and calcaneum from Riano and of S. hemitoechus (S .hem.), S. hund- sheimensis (S. hund.) and S. kirchbergensis (S. kirch.) reported by GUÉRIN (1980). Dimensioni comparative dei femori, delle rotule, dell’astragalo e del calcagno di Riano e di S. hemitoechus (S .hem.), S. hundsheimensis (S. hund.) e S. kirchbergensis (S. kirch.) dati da GUÉRIN (1980). ACKNOWLEDGEMENTS Thanks to Prof. C. Petronio for critical review and support. Also thanks to two anonymous reviewers, Prof. H.W. Tong, Prof. M. Follieri, Prof. D. Magri, Dr. R. Manni and Sig. L. Riti. REFERENCES ACCORDI B. & MACCAGNO A. M. (1962) - Researches in the Pleistocene of Riano (Rome) - Geol. 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Ms. ricevuto il 13/05/2010 Testo definitivo ricevuto il 4/08/2010 Ms. received: May 13, 2010 Final text received: August 4, 2010