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Atti Del Museo Civico Di Storia Naturale Di Trieste ISSN: 0335–1576 ATTI DEL MUSEO CIVICO DI STORIA NATURALE DI TRIESTE Supplemento al VOL. 53 – 2006 TRIESTE 2008 Atti Mus. Civ. Stor. Nat. Trieste Suppl. al 53 2008 3-12 novembre 2008 ISSN: 0335–1576 FOSSILI CRANIALI E MANDIBOLARI DI URSUS DA GROTTE DEL VENETO (ITALIA SETTENTRIONALE) NUOVE OSSERVAZIONI SUL QUADRO EVOLUTIVO DEGLI ORSI ITALIANI SANTI GIUSEPPE1 & ROSSI MARIO2 1 Dipartimento di Scienze della Terra, Università di Pavia, Via Ferrata 1, I–27100 Pavia (Italia): [email protected] 2 Museo Civico di Storia Naturale – Lungadige di Porta Vittoria 9, I–37127 Verona (Italia): [email protected] Abstract – Cranial and mandibular fossils of Ursus from caves of the Veneto region (North Italy). New observations on the evolutionary frame of the Italian bears. Caves of the Veneto region (North Italy) the Grotta del Ceré and Covoli di Velo (Verona province), are rich in bear’s fossils useful in the phyletic picture in particular. Considerations on a possible distinction of the two species (deningeri/spelaeus) and new morphological and morphometrical data mainly concerning the mandibular and dental (P4 e M1) remains, have been elaborated. The Ursus gr. deningeri–spelaeus from Grotta del Ceré should be Middle Pleistocene in age (proviously dated Upper Pleistocene), while fossils from Covoli di Velo are ascribed to the Late Pleistocene. Key word: Ursus gr. deningeri–spelaeus, mandibles, dentition, phylogenetic degree, Pleistocene, Northern Italy Riassunto – Sulla base di nuovi dati morfologici e morfometrici di resti di mandibole e denti (P4 e M1) attribuiti ad Ursus gr., deningeri–spelaeus raccolti nella Grotta del Cerè e ai Covoli di Velo (Veneto, Italia Settentrionale), sono proposte nuove considerazioni sul livello filetico raggiunto da questi orsi in questa parte dell’Italia. I fossili diU. deningeri (unicamente dalla Grotta del Cerè) avrebbero un’età riferita al Pleistocene medio, mentre quelli dai Covoli di Velo, al Pleistocene superiore. Inoltre, è fatto cenno al problema sulla possibile distinzione delle due specie di orso delle caverne: U. deningeri Von Reichenau, 1906 e U. spelaeus Rosenmüller, 1794. Parole chiave: Ursus gr. deningeri–spelaeus, mandibole, dentizione, grado filogenetico, Pleistocene, Italia Settentrionale 1. – Introduzione Le grotte del Veneto (Italia Settentrionale) (Fig. 1) hanno fornito una notevole quantità di resti di orso delle caverne (Ursus gr. deningeri–spelaeus) che sembrano coprire l’intero cammino evolutivo di questo plantigrado, dalle forme più antiche (U. deningeri Von Reichenau, 1906) a quelle più moderne (U. spelaeus Rosenmüller, 1794). Particolare interesse in questo senso rivestono le grotte del Cerè (Rossi, Santi, 2005) e dei Covoli di Velo (Zorzin et al., 2005) entrambe in provincia di Verona (Bon et al., 1991). Nella prima sono stati ritrovati diversi resti appartenenti alle tre specie tipiche del Quaternario europeo, U. deningeri, U. spelaeus ed U. arctos Linneo, 1758, mentre dalla seconda provengono abbondanti e ben conservati fossili di U. spelaeus del Pleistocene superiore, appartenenti quindi alle forme più evolute di questa specie. I reperti consentono di poter approfondire il quadro evolutivo disegnato dagli orsi “italiani”. 2. – Ursus deningeri Von Reichenau, 1906 (Fig. 2) A questa specie appartengono pochi resti mandibolari, alcuni denti isolati, tre crani, di cui uno fortemente deformato in corrispondenza delle orbite in seguito ai processi post mortem e diversi carpali e tarsali, quest’ultimi ancora in studio. 4 SANTI GIUSEPPE & ROSSI MARIO Fig. 1 – Posizione geografica delle principali grotte distribuite nel Veneto. Fig. 1 – Geographical position of the main caves located in the Veneto region (North Italy). Tutto il materiale proviene dalla Grotta del Cerè. Le mandibole sono caratterizzate da dimensioni molto piccole (Fig. 3), corrispondenti a quelle di esemplari medio–grandi di orso bruno. Rispetto alle forme “antiche” di questa specie (sensu Torres, 1988b), dal punto di vista morfologico questi reperti presentano però un maggior innalzamento del condilo e del processo angolare e soprattutto un bordo posteriore del ramo ascendente retto e non concavo, caratteri questi indicati da Torres (1988b) come tipici delle forme “moderne”, vicine alla transizione U. deningeri–U. spelaeus. La dentizione denota caratteri primitivi, particolarmente in P4 ed in M1. Il premolare è caratterizzato, infatti, dalla presenza del solo protoconide e di due cuspidi accessorie, appena accennate, una in corrispondenza del vertice anteriore della corona e la seconda (presente non i tutti i fossili) a ridosso del protoconide. Complessivamente i denti presentano notevoli somiglianze con quelli provenienti dalla Grotte de Fées ed appartenenti ad U. deningeri (Ballesio et al., 2003). Facendo riferimento ai morfotipi di Rabeder (1999), i reperti esaminati corrispondono al morfotipo B1. Il primo molare inferiore è caratterizzato da un’accentuata convergenza delle cuspidi, condizione questa tipica delle forme più primitive, ed un entoconide che OSSERVAZIONI SU FOSSILI DI URSUS DA GROTTE DEL VENETO, etc. 5 Fig. 2 – Ursus deningeri Von Reichenau, 1906. Grotta del Cerè (Verona). A. Cranio (Campione V 161), norma laterale sinistra, B. Mandibola destra (Campione V 4886), norma labiale. Entrambi i fossili sono custoditi presso il Museo Civico di Storia Naturale di Verona. Le parti contrassegnate con i numeri sono quelle che meglio identificano la specie. Per spiegazioni vedi testo. Fig. 2 – Ursus deningeri Von Reichenau, 1906. Grotta del Cerè (Verona). A. Skull (Specimen V 161), left lateral view, B. Right mandible (Specimen V 4886), labial view. Both fossils are stored into Museo Civico di Storia Naturale in Verona. Numbers sign the mandible parts useful for a identification of the species. Main explications in the text. mostra una cuspide principale preceduta da due cuspidi di dimensioni decrescenti, morfotipo considerato proprio della specie U. deningeri (Shütt, 1968; Bishop, 1982; Torres, 1988a; Argant, 1991, 1995; Grandal d’Anglade, López González, 2004). Dal punto di vista morfometrico (Figg. 4–5), i reperti rientrano completamente (P4) o parzialmente (M1) nel range tipico della specie, evidenziando complessivamente taglie medio–piccole. Nel caso del primo molare, la cui attribuzione è resa sicura dai caratteri morfologici presenti, il fatto che le dimensioni evidenziate dal grafico siano inferiori al range tipico di U. deningeri è molto probabilmente dovuto all’insufficienza di dati disponibili relativi ad altri depositi. 6 SANTI GIUSEPPE & ROSSI MARIO 85 U. arctos (AAVV) 80 U. spelaeus (AAVV) U. spelaeus (Cerè) 75 U. cfr. spelaeus (Cerè) U. deningeri (Cerè) 70 U. cfr. deningeri (Cerè) U. cfr. arctos (Cerè) 65 U. spelaeus (Velo) 60 55 50 45 40 Altezza della mandibola fra M2 e M3 35 65 70 75 80 85 90 95 100 105 110 115 Lunghezza della fila dentaria Fig. 3 – Relazione fra la “Lunghezza della fila dentaria” e l’“Altezza della mandibola fra 2M ed M3” (in mm) in orsi dalla Grotta del Cerè, Covoli di Velo e da grotte straniere. Fig. 3 – “Lenght of dental row” versus “Height of the mandible between M2 and M3” (in mm) in bears from Grotta del Cerè, Covoli di Velo and different foreigner caves also. 13 12 4 11 10 9 U. spelaeus (AAVV) U. deningeri (AAVV) 8 U. arctos (AAVV) U. arctos (Olocene) U. spelaeus (Cerè) Larghezza assoluta P 7 U. deningeri (Cerè) 6 U. arctos (Cerè) U. spelaeus (Velo) 5 10,5 11,5 12,5 13,5 14,5 15,5 16,5 17,5 18,5 Lunghezza assoluta P4 Fig. 4 – Relazione fra la “Lunghezza assoluta di P4” e la “Larghezza assoluta di P4 “(in mm) in orsi dalla Grotta del Cerè, Covoli di Velo e da grotte straniere. Fig. 4 – “Absolute lenght of P4” versus “Absolute breadth of P4” (in mm) in bears from Grotta del Cerè, Covoli di Velo and different foreigner caves also. OSSERVAZIONI SU FOSSILI DI URSUS DA GROTTE DEL VENETO, etc. 7 18 U. spelaeus (AAVV) 17 U. deningeri (AAVV) U. arctos (AAVV) 1 16 U. arctos (Olocene) U. spelaeus (Cerè) 15 U. deningeri (Cerè) U. arctos (Cerè) 14 U. spelaeus (Velo) 13 12 Larghezza assoluta M 11 10 9 20 22 24 26 28 30 32 Lunghezza assoluta M1 Fig. 5 – Relazione fra la “Lunghezza assoluta di M1” e la “Larghezza assoluta di M1” (in mm) in orsi dalla Grotta del Cerè, Covoli di Velo e da grotte straniere. Fig. 5 – “Aboslute lenght of M1” versus “Absolute breadth of M1” (in mm) in bears from Grotta del Cerè, Covoli di Velo and different foreigner caves also. 3. – Ursus spelaeus Rosenmüller, 1794 (Fig. 6) I resti relativi a questa specie provenienti dalla Grotta del Cerè hanno dimensioni piccole (Fig. 3), ma soprattutto presentano dentizioni con caratteri di arcaicità. L’unico quarto premolare, infatti, ha solo due cuspidi accessorie, seppure ben sviluppate, corrispondente al morfotipo C1 sensu Rabeder (1999). Il primo molare inferiore è caratterizzato da una convergenza delle cuspidi maggiore di quella che identifica i reperti provenienti dai giacimenti più recenti (Grandal d’Anglade, López González, 2004). Le mandibole provenienti dai Covoli di Velo presentano dimensioni (Fig. 3) e caratteri morfologici tipici delle forme più evolute. La dentizione, ancora in studio, fornisce invece, ad una prima analisi, elementi più contraddittori. Il quarto premolare, infatti, non presenta una particolare complicazione della superficie dentaria, simile a quella visibile nelle forme più evolute (Rabeder, 1999), mentre il primo molare ha una superficie triturante decisamente complessa ed un grado di convergenza delle cuspidi paragonabile a quella tipica delle forme più recenti (Grandal d’Anglade, López González, 2004). 4. – Discussione e conclusioni I fossili provenienti dalla Grotta del Cerè e dai Covoli di Velo sembrano rappresentare l’intera linea evolutiva dell’orso delle caverne: dalla prima, infatti, 8 SANTI GIUSEPPE & ROSSI MARIO Fig. 6 – Ursus spelaeus Rosenmüller, 1794. Covoli di Velo (Verona). A. Cranio (Campione V 9894), norma laterale destra, B. Mandibola sinistra (Campione 14.56928), norma linguale. Entrambi i fossili sono custoditi presso il Museo Civico di Storia Naturale di Verona. Fig. 6 – Ursus spelaeus Rosenmüller, 1794. Covoli di Velo (Verona). A. Skull (Specimen V 9894), right lateral view, B.
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