SESS 3_14022011_edi as (3).pub Il Quaternario Italian Journal of Quaternary Sciences 24, (Abstract AIQUA, Roma 02/2011), Congresso AIQUA Il Quaternario Italiano: conoscenze e prospettive Roma 24 e 25 febbraio 2011 THE EARLY AND MIDDLE PLEISTOCENE SABERTOOTHED CATS IN EUROPE: TAXONOMY, BIOCHRONOLOGY AND PALEOECOLOGY. Raffaele Sardella & Dawid Adam Iurino Dipartimento di Scienze della Terra, Università “Sapienza”, Roma Corresponding author: R. Sardella ABSTRACT: Sardella R. & Iurino D., The Early and Middle Pleistocene sabertoothed cats in Europe: taxonomy, biochro- nology and paleoecology. (IT ISSN 0394-3356, 2011) Two different groups of sabertoothed cats (Machairodontinae, Felidae, Mammalia) characterized the Early Pleistocene terrestrial ecosystems in Europe: Homotherium and Meganteron. The taxonomy of these felids is matter of debate, and their importance in biochronology and paleocology is analyzed. In Europe Megantereon become exctinct at the end of Early Pleistocene, while Homotherium seems to survive until the Late Pleistocene. RIASSUNTO: Sardella R. & Iurino D., A. I felidi macairodonti nel Pleistocene Inferiore e Medio in Europa: questioni tas- sonomiche, biocronologia, paleoecologia. (IT ISSN 0394-3356, 2011) Gli ecosistemi continentali europei del Pleistocene Inferiore sono caratterizzati dalla presenza delle cosiddette “tigri dai denti a sciabola” (Machairodontinae, Felidae, Mammalia), rappresentate da due taxa che rispecchiano due diversi tipi di adattamento: Homotherium and Meganteron. La sistematica di questi felidi è molto dibattuta e sarà discussa insieme all’importanza biocronologica e paleoecologica di questi carnivori. In Europa Megantereon si estingue alla fine del Plei- stocene Inefriroe, mentre Homotherium diviene molto raro e sopravvive sino al Pleistocene Superiore. Key words: biochronology, mammals, dispersal events, paleoecology, terrestrial ecosystems Parole chiave: biocronologia, mammiferi, eventi di dispersione, paleoecologia, ecosistemi continentali Among the top predators sabertoothed cats (Machairodontinae, Felidae) were dominant in the Early Pleistocene terrestrial ecosystems of Europe. Two genera, corresponding to two different line- ages and morpho-functional adaptations, charac- terized the Villafranchian faunal assemblages: Homotherium and Megantereon. The taxonomy of these felids is based mainly on craniodental fea- tures and, despite an increasing number of locali- ties provided a larger amount of fossils, including also postcranial bones, the specific attribution of such material seems to be a quite complicated matter. Homotherium had unique craniodental as well as postcranial adaptations distinguishing it (and all the forms referable to the Homotherinii tribe) from other Machairodontinae. Homotherium evolved from derived forms of the Machairodus lineage and forms referable to the Late Miocene paraphyletic species complex Am- phimachirodus can be considered its immediate ancestors (WERDELIN L. & SARDELLA R., 2006). The timing of the Amphimachairodus-Homotherium transition is still unclear. The features of Homotherium suggest a unique ecomorphology and behaviour. It was a lion sized felid with crenu- lated teeth; upper canines were elongated, flat and serrated. Incisors were powerful, stronger than in any living felid. The limb proportions of Homotherium were very different than those of any living felid. The forelimbs were elongated, while the hindlimbs were shorter and quite stout. A number of different species have been proposed for the European Homotherium (H. hungaricus, H. sainzelli, H. crenatidens, H. latidens, H. moravi- cum) mainly based on the size and on the upper canine morphology. At present, many authors re- fers all the European specimens to Homotherium latidens, Nonetheless, the earliest European specimens show peculiar craniodental characters (longer and narrower upper canines, stronger anterior part of the mandible, with convex labial surface) that can be considered ‘‘archaic’’ features. Thus, the early Villafranchian specimens should be referred to the species Homotherium nestianus that has been re- corded at Roca Neyra (France), Pievefosciana (Garfagnana, Tuscany) and Sammezzano (Upper Valdarno, Tuscany) (SARDELLA R., 1994 and refer- ences therein). Homotherium latidens characterized the late Villa- franchian faunal assemblages and became very rare in the Middle Pleistocene; it seems to survive in Northwestern Europe until the Late Pleistocene (REUMER J. F. W. et al., 2003). Finally, the presence of a preparastyle in the upper carnassial character- izes some galerian large sized Homotherium specimens. In the past some of these specimens were referred to Homotherium moravicum (Stranska Skala, Moravia; Hundsheim, Austria) (SARDELLA R., 1994), but in many following studies this species has been referred to H. crenatidens 170 - 171 170 Ms. received: Testo ricevuto il and/or H. latidens. Fossils sharing the “H. moravi- cum morphology” in the upper carnassials have been found also in some latest Early Pleistocene sites in northern Italy (Domegliara Selvavecchia, and other unpublished localities). It is possible that the occurrence of a separate species has be re- considered. The genus Megantereon belongs to the Smilodon- tini tribe and includes Plio-Pleistocene jaguar- sized, short limbed, dirk-toothed felids with a stoutly built postcranial skeleton, in which the ro- bust distal limb bones indicate an ambushing hunt- ing method. The earliest occurrence of Megantereon in Europe in recorded in the Middle Pliocene of Les Etouaires, France (possibly 3.5 Ma, but not older than 2.5), the latest occurrence of the taxon is at Untermassfeld (Thuringia, Germany) (SARDELLA R., 1994 and references therein, HEMMER H., 2001). Taxonomy and evolution of this felid have been discussed in several papers issued during recent decades and different interpretations have been provided. All the authors agree to refer Pliocene European specimens to M. cultridens. Different interpretations are proposed for Early Pleistocene European forms. Turner considered the differences recorded into the Eurasian and African sample of Megantereon as due to sexual dimorphism, com- paring the data to those of the living leopard. Other authors claimed a convergent evolution of the European samples of M. cultridens with the African M. whitei, due to the paleoclimate and the paleoen- vironmental conditions occurring during the earliest part of the Pleistocene. They named this form M. cultridens adroveri (HEMMER H., 2001 and refer- ences therein). SARDELLA R. (1998) suggested the use of open taxonomy including the Early Pleisto- cene European specimens in Megantereon ex gr. cultridens (advanced form). Recently, the attribution to Megantereon whitei was supported by further data and adopted by many authors (PALMQVIST P. et al., 2007). The dispersal of M. whitei into Europe probably occurred concurrently with other African species following the Levantine Corridor. In Europe this taxon was found in Venta Micena (Spain), Monte Argentario, Pirro Nord (Italy) and Greece (Apollonia) (SARDELLA R. et al., 2008 and refer- ences therein). REFERENCES HEMMER H. (2001) - Die Feliden aus dem Epivillafran- chium von Untermaßfeld. In: Kahlke, R.-D. (Ed.), Das Pleistozän von Untermaßfeld bei Meiningen (Thüringen), Teil 3. Monographien des Römisch- Germanischen Zentralmuseums Mainz 40, 699-782. PALMQVIST P., TORREGROSA V., PÉREZ-CLAROS J.A., MARTÍNEZ-NAVARRO B. & TURNER A. (2007) - A re- evaluation of the diversity of Megantereon (Mammalia, Carnivora, Machairodontinae) and the problem of spe- cies identification in extinct carnivores. J. Vertebr. Pale- ontol. 27, 160–175. REUMER J. F. W., ROOK L., VAN DER BORG,K., POST K., MOL D. & DE VOS J. (2003) - Late Pleistocene survival of the saber-toothed cat Homotherium in Northwestern Europe. J. Vertebr. Paleontol., 23(1), 260-262. SARDELLA R. (1994) - Sistematica e distribuzione strati- grafica dei Macairodontini dal Miocene Superiore al Pleistocene, PhD dissertation, Bologna, Firenze, Mode- na and Roma “La Sapienza” Universities, Rome 1-137 pp. SARDELLA R. (1998) - The Plio-Pleistocene dirk toothed cat Megantereon ex gr. cultridens (Mammalia, Felidae, Macharodontinae) with comments on taxonomy, origin and evolution. N. Jahrb. Geol. Paläontol. Abh. 207, 1– 36. SARDELLA R., PETRUCCI M. & ROOK L. (2008) - The Afri- can species Megantereon whitei from the Early Pleisto- cene of Monte Argentario (South Tuscany, Central It- aly). - C. R. Palevol., 7, 601-606. WERDELIN L. & SARDELLA R. (2006) - The “Homotherium” from Langebaanweg, South Africa and the origin of Homotherium. Palaeontographica, Abt. A, 277 (1-6), 123-130  Fig. 1, Megantereon whitei skulls from Early Pleistocene Italian localities. G1 from Pirro Nord, Apulia (up) and ARG31 from Monte Argentario, Tuscany (lower, stored at the Istituto Italiano di Paleontologia Umana, Roma). Scale bar: 1 cm. Megantereon whitei. Crani provenienti da siti italiani del Pleistocene Inferiore. (G1) Pirro Nord, Puglia (sopra) e ARG31 dal Monte Argentario, Toscana (sotto, conserva- to presso l’Istituto Italiano di Paleontologia Umana, Ro- ma). Riferimento metrico: 1 cm.  January 15, 2011 15 gennaio 2011 The Early and Middle Pleistocene sabertoothed ... << /ASCII85EncodePages false /AllowTransparency false /AutoPositionEPSFiles true /AutoRotatePages /None /Binding /Left /CalGrayProfile (Dot Gain 20%) /CalRGBProfile (sRGB IEC61966-2.1) /CalCMYKProfile (U.S. Web Coated \050SWOP\051 v2) /sRGBProfile (sRGB IEC61966-2.1) /CannotEmbedFontPolicy /Error /CompatibilityLevel 1.4 /CompressObjects /Tags /CompressPages true /ConvertImagesToIndexed true /PassThroughJPEGImages true /CreateJobTicket false /DefaultRenderingIntent /Default /DetectBlends true /DetectCurves 0.0000 /ColorConversionStrategy /CMYK /DoThumbnails false /EmbedAllFonts true /EmbedOpenType false /ParseICCProfilesInComments true /EmbedJobOptions true /DSCReportingLevel 0 /EmitDSCWarnings false /EndPage -1 /ImageMemory 1048576 /LockDistillerParams false /MaxSubsetPct 100 /Optimize true /OPM 1 /ParseDSCComments true /ParseDSCCommentsForDocInfo true /PreserveCopyPage true /PreserveDICMYKValues true /PreserveEPSInfo true /PreserveFlatness true /PreserveHalftoneInfo false /PreserveOPIComments true /PreserveOverprintSettings true /StartPage 1 /SubsetFonts true /TransferFunctionInfo /Apply /UCRandBGInfo /Preserve /UsePrologue false /ColorSettingsFile () /AlwaysEmbed [ true ] /NeverEmbed [ true ] /AntiAliasColorImages false /CropColorImages true /ColorImageMinResolution 300 /ColorImageMinResolutionPolicy /OK /DownsampleColorImages true /ColorImageDownsampleType /Bicubic /ColorImageResolution 300 /ColorImageDepth -1 /ColorImageMinDownsampleDepth 1 /ColorImageDownsampleThreshold 1.50000 /EncodeColorImages true /ColorImageFilter /DCTEncode /AutoFilterColorImages true /ColorImageAutoFilterStrategy /JPEG /ColorACSImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /ColorImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /JPEG2000ColorACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000ColorImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasGrayImages false /CropGrayImages true /GrayImageMinResolution 300 /GrayImageMinResolutionPolicy /OK /DownsampleGrayImages true /GrayImageDownsampleType /Bicubic /GrayImageResolution 300 /GrayImageDepth -1 /GrayImageMinDownsampleDepth 2 /GrayImageDownsampleThreshold 1.50000 /EncodeGrayImages true /GrayImageFilter /DCTEncode /AutoFilterGrayImages true /GrayImageAutoFilterStrategy /JPEG /GrayACSImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /GrayImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /JPEG2000GrayACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000GrayImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasMonoImages false /CropMonoImages true /MonoImageMinResolution 1200 /MonoImageMinResolutionPolicy /OK /DownsampleMonoImages true /MonoImageDownsampleType /Bicubic /MonoImageResolution 1200 /MonoImageDepth -1 /MonoImageDownsampleThreshold 1.50000 /EncodeMonoImages true /MonoImageFilter /CCITTFaxEncode /MonoImageDict << /K -1 >> /AllowPSXObjects false /CheckCompliance [ /None ] /PDFX1aCheck false /PDFX3Check false /PDFXCompliantPDFOnly false /PDFXNoTrimBoxError true /PDFXTrimBoxToMediaBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXSetBleedBoxToMediaBox true /PDFXBleedBoxToTrimBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXOutputIntentProfile () /PDFXOutputConditionIdentifier () /PDFXOutputCondition () /PDFXRegistryName () /PDFXTrapped /False /CreateJDFFile false /Description << /ARA /BGR /CHS /CHT /CZE /DAN /DEU /ESP /ETI /FRA /GRE /HEB /HRV (Za stvaranje Adobe PDF dokumenata najpogodnijih za visokokvalitetni ispis prije tiskanja koristite ove postavke. 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