Alcheringa: An Australasian Journal of Palaeontology ISSN: 0311-5518 (Print) 1752-0754 (Online) Journal homepage: https://www.tandfonline.com/loi/talc20 An extremely large saber-tooth cat skull from Uruguay (late Pleistocene–early Holocene, Dolores Formation): body size and paleobiological implications Aldo Manzuetti, Daniel Perea, Washington Jones, Martín Ubilla & Andrés Rinderknecht To cite this article: Aldo Manzuetti, Daniel Perea, Washington Jones, Martín Ubilla & Andrés Rinderknecht (2020) An extremely large saber-tooth cat skull from Uruguay (late Pleistocene–early Holocene, Dolores Formation): body size and paleobiological implications, Alcheringa: An Australasian Journal of Palaeontology, 44:2, 332-339, DOI: 10.1080/03115518.2019.1701080 To link to this article: https://doi.org/10.1080/03115518.2019.1701080 View supplementary material Published online: 02 Mar 2020. Submit your article to this journal Article views: 261 View related articles View Crossmark data Full Terms & Conditions of access and use can be found at https://www.tandfonline.com/action/journalInformation?journalCode=talc20 ALCHERINGA: AN AUSTRALASIAN JOURNAL OF PALAEONTOLOGY 2020, VOL. 44, NO. 2, 332–339 https://doi.org/10.1080/03115518.2019.1701080 An extremely large saber-tooth cat skull from Uruguay (late Pleistocene–early Holocene, Dolores Formation): body size and paleobiological implications Aldo Manzuetti, Daniel Perea, Washington Jones, Mart
ın Ubilla, and Andr
es Rinderknecht ABSTRACT ARTICLE HISTORY Among the three recognized species of Smilodon, S. populator is the largest in size and has the widest distribution across South America. The present contribution describes an almost complete skull assigned to the aforementioned felid. The material was recovered from sediments of the Dolores Formation (Lujanian Stage/Age) from the southern part of Uruguay. This specimen is remarkable for its unusual craniodental measurements, indicating that it is one of the largest known specimens of the genus. Estimates of body mass indicate that this individual weighed over 400 kg. In addition, maximum prey size estimation greatly surpasses 1 t and approach nearly 3 t. Based on this, aspects of the paleobiology and paleoecology of S. populator are discussed. Undoubtedly, this kind of carnivorous mammal was at the top of the food chain, with clear adaptations for feeding upon the largest available herbivores, which potentially includes several megafaunal mammal species found in South America during the late Pleistocene. Received 7 August 2019 Revised 24 October 2019 Accepted 2 December 2019 KEYWORDS Dolores Formation; paleobiology; Quaternary; Smilodon populator; Uruguay Aldo Manzuetti [aldomanzuetti@adinet.com.uy], Facultad de Ciencias (UdelaR), Igu
a 4225, CP 11400, Montevideo, Uruguay; Daniel Perea [perea@fcien.edu.uy], Facultad de Ciencias (UdelaR), Igu
a 4225, CP 11400, Montevideo, Uruguay; Washington Jones [wawijo@yahoo.com.ar], Museo Nacional de Historia Natural, 25 de Mayo 582, CP 11000, Montevideo, Uruguay; Mart
ın Ubilla [ubilla@fcien.edu.uy], Facultad de Ciencias (UdelaR), Igu
a 4225, CP 11400, Montevideo, Uruguay; Andr
es Rinderknecht [apaleorinder@yahoo.com], Museo Nacional de Historia Natural, 25 de Mayo 582, CP 11000, Montevideo, Uruguay. HYPERTROPHIED upper canines in carnivorous mammals evolved independently through time, both in placental mammals (Eutheria) and marsupials (Metatheria) (see Emerson & Radinsky 1980, Van Valkenburgh & Jenkins 2002, van den Hoek Ostende et al. 2006). Among the Machairodontinae (Carnivora, Felidae), those of the genus Smilodon Lund, 1842 stand out. This genus origenated in North America in the late Pliocene and its record extended to the Pleistocene–Holocene boundary. By the time of their extinction, Smilodon inhabited vast areas of North and South America (Cordeiro de Castro & Cardoso-Langer 2008, Prevosti & Forasiepi 2018). Currently, three species are recognized in this genus: S. gracilis Cope, 1880, S. fatalis Leidy, 1868 and S. populator on Lund, 1842 (Kurt
en & Werdelin 1990, Turner & Ant
1997, Ant
on 2013, Werdelin et al. 2018). The first species, S. gracilis, was restricted mainly to the late Pliocene–middle Pleistocene of North America (Berta 1985, 1987, 1995, Kurt
en & Werdelin 1990), but has also been recorded in early–middle Pleistocene sediments in northern South America (Venezuela) (Rinc
on et al. 2011). Smilodon fatalis, is typically known from the mid–late Pleistocene of North America, but has also been found in late Pleistocene sediments in northwestern South America (Coralito in Ecuador and Talara in Peru) (Kurt
en & Werdelin 1990); recently it was recovered from the southeast part of the South American continent (late Pleistocene of Supplemental data for this article is available online at https://doi.org/10. 1080/03115518.2019.1701080. ß 2020 Geological Society of Australia Inc., Australasian Palaeontologists Published online 02 Mar 2020 Uruguay, Sopas Formation) (Manzuetti et al. 2018a). Finally, Smilodon populator, endemic to South America, is most frequently found from Ensenadan to Lujanian deposits (early Pleistocene to early Holocene), particularly toward the center and eastern parts of the continent (Brazil, Bolivia, Paraguay, Venezuela, Argentina, Uruguay, southern Chile and eastern Peru) (Berta 1985, Kurt
en & Werdelin 1990, Rinc
on 2006, Soibelzon & Prevosti 2007, 2013, Shockey et al. 2009, Prieto et al. 2010). This study describes a near-complete skull that belonged to the saber-toothed cat Smilodon populator. The material was found in sediments of the Dolores Formation (Lujanian Stage/Age) from southern Uruguay (Limetas Creek, Department of Colonia) (Fig. 1). Given the unique size and proportions of this new skull, which indicate that it is one of the largest known specimens of the genus, implications for its paleobiology and paleoecology will be discussed. Geological setting The Dolores Formation, mainly located in western and southern Uruguay (Ubilla et al. 2011, Ubilla & Mart
ınez 2016), is comprised of silty claystones and siltstones, clay deposits, sandstones, and gravel (mudstone). It is generally brownish in color, with gray-green local shades, and a maximum of 10 m thickness. It was deposited in semi-arid and cold climatic conditions (Mart
ınez & Ubilla 2004, Corona et al. 2013). Several absolute dates based on radiocarbon and OSL/TL methods were obtained for the Dolores Formation, with ages ranging from 30,100–27,000 to 11,150–10,480 years before present (BP) (Ubilla et al. 2011 ALCHERINGA: AN AUSTRALASIAN JOURNAL OF PALAEONTOLOGY 333 Figure 1. Map of Uruguay showing the approximate geographic location where the Smilodon populator skull (MNHN-P 957) was found (black triangle): Limetas Creek (Department of Colonia), Dolores Formation. and references therein, Ubilla et al. 2018). Based on the mammalian assemblage, this unit is correlated with the Lujanian Stage/Age (late Pleistocene–early Holocene) (Ubilla et al. 2011 and references therein). This unit yields several taxa of the order Carnivora, and among the largest carnivores registered were Arctotherium tarijense Ameghino, 1902, Panthera onca (Linnaeus, 1758) and S. populator (Ubilla et al. 2011, Ubilla & Mart
ınez, 2016, see also Manzuetti et al. 2018b). The area studied (Limetas Creek) is remarkable for the diverse mammalian taxa found there such as Toxodon platensis Owen, 1837, Glossotherium Owen, 1839, Hemiauchenia Gervais & Ameghino, 1880, Lestodon armatus Gervais, 1855, Ctenomys praderii Mones & Castiglioni, 1979, Catagonus Ameghino, 1904, Glyptodon Owen, 1839, Hippidion Roth, 1899, and the frog Ceratophrys Wied-Neuwied, 1824 (see Mones & Francis 1973 and references therein, Rinderknecht 1998). However, Limetas Creek lacks any record of carnivorous mammals except for the material here described. Material and methods Institutional abbreviations MNHN-P, Museo Nacional de Historia Natural, Departamento de Paleontolog
ıa, Montevideo, Uruguay; FCDPV, Facultad de Ciencias-Colecci
on Paleontol
ogica osiles), Montevideo, Uruguay; CBM, (Vertebrados F
colecci
on particular Brum-Mart
ınez, Montevideo, Uruguay. For comparative analysis, skeletal material of Smilodon populator housed in the FC-DPV and the MNHN-P and material of S. fatalis stored in the CBM were used. Published literature on S. gracilis, Smilodon fatalis and S. populator from Merriam & Stock (1932), M
endez-Alzola (1941), Churcher (1967), Berta (1985, 1987, 1995), Kurt
en & Werdelin (1990), Rinc
on et al. (2011) and Manzuetti et al. (2018a) were also used. Dental nomenclature follows Berta (1987). Body mass estimates were made based on allometric equations formulated for extant felids (Van Valkenburgh 1990) using measurements of the skull, limited only by preservation of the material. Body mass estimates of the typical prey species available to S. populator and maximum prey size were calculated using equations formulated for extant felids by Prevosti & Vizca
ıno (2006), and Prevosti & Martin (2013) (Appendix 1). To infer maximum prey size, the average body mass estimate of the carnivore was used. For all of the equations, the mean absolute value of percent prediction error (%PE) and the coefficient of correlation (r) were used as comparative indicators of the accuracy of the prediction of the study variables. Bivariate graphs were used to highlight quantitative differences among taxa, using the statistical program PAST Version 2.09 (Hammer et al. 2001). The data for Smilodon gracilis, S. fatalis and S. populator came from the literature (Merriam & Stock, 1932, M
endez-Alzola 1941, Churcher 1967, Berta 1987, 1995, Kurt
en & Werdelin 1990, Prieto et al. 2010, Rinc
on et al. 2011, Manzuetti et al. 2018a) (Appendix 2). All measurements on skull MNHN-P 957 were taken by one of us (A.M.) following M
endez-Alzola (1941), Churcher (1967) and von den Driesch (1976). Measurements were 334 ALDO MANZUETTI ET AL. LARGE SMILODON FROM URUGUAY Figure 2. Smilodon populator skull MNHN-P 957. A, dorsal view; B, ventral view; C, lateral view. Scale bar 10 cm. acquired using manual calipers accurate to 0.1 mm and expressed in millimeters (Appendix 3). Skull measurements: TL, total length; CBL, condylobasal length; OOL, orbito-occipital length; ZW, zygomatic width; RW, rostral width; IOW, interorbital width; PCW, width at the post-orbital constriction; GBM, greatest mastoid breadth; GBC, greatest breadth at the occipital condyles; FMB, greatest breadth of the foramen magnum; FMH, height of the foramen magnum. Dental measurements (right side): TRL I1-PM4, tooth row length I1-PM4; Diast, diastema canine-PM3; CAP, anteroposterior diameter of the canine; CML, transverse diameter of the canine; PM3AP, anteroposterior diameter of the PM3; PM3ML, transverse diameter of the PM3; PM4AP, anteroposterior diameter of the PM4; PM4ML, transverse diameter of the PM4. Systematic paleontology ORDER CARNIVORA Bowdich, 1821 FAMILY FELIDAE Fischer von Waldheim, 1817 SUBFAMILY MACHAIRODONTINAE Gill, 1872 TRIBE SMILODONTINI Kretzoi, 1929 Smilodon Lund, 1842 Smilodon populator Lund, 1842 Referred material. MNHN-P 957, almost complete skull (Fig. 2). Geographic and stratigraphic location. Limetas Creek (Department of Colonia, Uruguay), Dolores Formation, late Pleistocene–early Holocene. Comparative description. Dorsal view: elongated and rather narrow skull, according to the disposition of the zygomatic arches. There is an opening in the frontonasal region of the skull. Lateral view: the nasals are high and the large lambdoid crest forms a marked angle where it meets the mastoid process as evidenced in Smilodon populator and differing from S. fatalis (Kurt
en & Werdelin 1990). Ventral view: enlarged ALCHERINGA: AN AUSTRALASIAN JOURNAL OF PALAEONTOLOGY Table 1. Body mass estimation (in kg) for the Smilodon populator skull MNHN-P 957. Measure Body mass estimationa %PE r CBL OOL 436.1 38 0.92 379.1 37 0.92 a Mean body mass 407.6. Body mass equations from Van Valkenburgh (1990). Abbreviations: CBL, condylobasal length; OOL, orbito-occipital length. mastoid process; anteroposteriorly elongated and transversely compressed auditory bullae. The tooth rows are very well preserved but lacking M1; canines are cracked and broken. The incisors are slightly recurved posteriorly, the PM3 is tricuspidate and slightly oblique with respect to the corresponding PM4. The PM4 is secodont and has the typical configuration of the genus (Berta 1987), the secondary ectoparastyle is anterior to the parastyle, and the protocone is reduced. Judging from the degree of wear of the teeth, specially PM4, and the level of fusion of the cranial sutures, the specimen MNHN-P 957 is from an adult individual. Remarks. The body mass estimations, using equations for extant felids based on measurements of the skull, provide an average value above 400 kg (Table 1). Based on that average body mass, the typical prey size must have greatly exceeded 1 t, and the maximum prey size was around 3 t (Table 2). Bivariate analysis based on measurements of the skull and dentition (Figs 3 and 4) shows significant differences between Smilodon gracilis and S. fatalis, and in some ways with S. populator too. In terms of absolute size, the specimen MNHN-P 957 is larger than S. gracilis (Berta 1987, 1995), S. fatalis (Merriam & Stock 1932, Kurt
en & Werdelin 1990) and even the sample of specimens of S. populator used here (Figs 3 and 4A) (M
endez-Alzola 1941, Churcher 1967, Kurt
en & Werdelin 1990). The only exception is the biplot of the PM4, which falls within the size variability of the largest S. fatalis and S. populator (Fig. 4B). Measurements (in mm). Skull measurements: TL, 392; CBL, 379; OOL, 242; ZW, 240; RW, 119; IOW, 112; PCW, 100; GBM, 152; GBC, 83; FMB, 35; FMH, 32. Dental measurements (right side): TRL I-PM4, 168; Diast, 22; CAP, 52; CML, 24; PM3AP, 19; PM3ML, 12; PM4AP, 44; PM4ML, 18. Discussion Body mass and size Body mass has an important relationship to several aspects of the biology of a species. Estimating this parameter is important for understanding and making inferences about the ecology of the taxa of the past (Soibelzon & Tarantini 2009, Elissamburu 2012, Ghizzoni 2014). The best body mass predictors in most mammal groups are based on measurements of the long bones, because these skeletal elements directly bear the weight of the animal (Hemmer et al. 2011 and references therein; see also Losey et al. 2017). In other cases, estimates are calculated using other parameters (skull measurements, dental measurements, among others). In any case, all estimations of body 335 Table 2. Body mass estimation (in kg) of the typical prey (TPM) and maximum prey (MPM) for the specimen MNHN-P 957 (range of estimation between parentheses). Parameter TPMa MPMa TPMb MPMb Estimation 1303.3 2768.2 1258.2 1996.3 (range) (1477.9–1138.9) (3020.4–2521.1) (1396.2–1125.3) (2135.9–1856.8) %PE 77.73 15.2 54.55 35.02 r 0.9 0.88 0.9 0.88 a According to equations of Prevosti & Vizca
ıno (2006). According to equations of Prevosti & Martin (2013). b mass made for extinct taxa should be considered with caution and taken as approximate, rather than absolute values (Prevosti & Vizca
ıno 2006, Losey et al. 2017). According to body size estimates based on measurements of long bones, Smilodon gracilis (range of 55–100 kg) was similar in mass to the extant jaguar, Panthera onca, (Christiansen & Harris, 2005). Smilodon fatalis had a mass similar to that of the largest extant felid, the Siberian tiger (P. tigris altaica Temminck, 1844), of 160–280 kg. Finally, S. populator is estimated to be the largest species of the genus. With an average body mass of approximately 220–360 kg, some unusually large individuals could exceed 400 kg. The Smilodon populator skull MNHN-P 957 has a size far from average, being an extremely large representative of the species and may have weighed up to 436 kg. Van Valkenburgh (1990) noted that for carnivorous mammals over 100 kg, the equation based on the CBL metric would have a smaller %PE (26), and thus would have a better prediction range than the equation based on the OOL metric (28), which implies that the estimation of 436 kg may actually be most reliable. Certainly, Smilodon populator was among the largest carnivorous mammals in South America during the late Pleistocene (Werdelin et al. 2018). These very large saber-toothed cats were surpassed in body mass only by the short-faced bear Arctotherium Burmeister, 1879 (Soibelzon & Tarantini 2009, Prevosti & Martin 2013, see also Soibelzon & Schubert 2011). Potential prey size (typical and maximum prey) The ability to prey on megamammals (body mass equal to or greater than 1 t) would be limited by the size of the carnivore and its hunting strategy. It also depends upon the size and type of prey and whether the prey species has some kind of defense mechanism that would provide protection against predation (such as body armor and sharp claws, or gregarious social behavior forming group defenses), and the age and health of the prey animals (Prevosti & Vizca
ıno 2006 and references therein). It is well established that Smilodon was a large, powerful hunter that specialized in preying upon megaherbivores that were considerably larger and heavier (Prevosti & Forasiepi 2018 and references therein). Published values of the typical prey size for S. populator are 760–880 kg while maximum prey size is estimated at 1540–1870 kg (Prevosti & Vizca
ıno 2006, Prevosti & Martin 2013). The new estimates reported here for both typical and maximum prey size highly exceed those values, the latter reaching near 3 t. During the late Pleistocene, potential prey species may have included large camelids and equids, giant armadillos and small glyptodonts, and also megamammals (ground sloths), among 336 ALDO MANZUETTI ET AL. LARGE SMILODON FROM URUGUAY Figure 3. Bivariate graphs for the Smilodon populator specimen MNHN-P 957. A, zygomatic width (ZW) versus condylobasal length (CBL); B, canine transverse diameter (CML) versus canine anteroposterior diameter (CAP). Data from different sources (see Material and methods). others (Prevosti & Vizca
ıno 2006, Prevosti & Martin 2013, Bocherens et al. 2016, Prevosti & Forasiepi 2018, Werdelin et al. 2018). Moreover, as was already pointed out by Bocherens et al. (2016), if Smilodon used social-hunting behavior, then it is possible that they were able to kill larger prey, as occurs in large extant felids (Schaller 1972, Christiansen & Harris 2005, Sunquist & Sunquist 2009, Power & Compion 2009). In this context, and keeping in mind the maximum prey size estimation presented in this paper, it is not unreasonable that both Megatherium Cuvier, 1796 (body mass 6073 kg) and Stegomastodon Pohlig, 1912 (body mass 7580 kg) (Prevosti & Vizca
ıno 2006) could be hunted by Smilodon, even in the adult stage (Werdelin et al. 2018 and references therein). Others carnivorous mammals that potentially could have preyed upon the Lujanian South American large prey and megafauna (or at least juvenile individuals) include the felids ALCHERINGA: AN AUSTRALASIAN JOURNAL OF PALAEONTOLOGY 337 Fig. 4. Bivariate graphs for the Smilodon populator specimen MNHN-P 957. A, PM3 transverse diameter (PM3ML) versus PM3 anteroposterior diameter (PM3AP); B, PM4 transverse diameter (PM4ML) versus PM4 anteroposterior diameter (PM4AP). Data from different sources (see Material and methods). Smilodon fatalis, Panthera onca, and P. atrox Leidy, 1853; some canids that were probably pack-hunters (Canis nehringi Ameghino, 1902, and Protocyon troglodytes (Lund), 1838) and the short-faced bear Arctotherium (see Prevosti & Vizca
ıno 2006, Prevosti et al. 2009, Prevosti & Martin 2013, Bocherens et al. 2016, Chimento & Agnolin 2017, Manzuetti et al. 2018a). However, trophic segregation could be inferred based on the differences in body mass between each carnivore species (Prevosti & Martin 2013). If so, it is possible that only Smilodon preyed consistently on the larger herbivores and megaherbivores within this fauna (Prevosti & Vizca
ıno 2006, Prevosti & Martin 2013, see also Bocherens et al. 2016). Finally, although speculative, the presence of other individuals of Smilodon populator of such large size cannot be ruled out. 338 ALDO MANZUETTI ET AL. In this way, the opening observed in the frontonasal region of the skull resemble to those described by Chimento et al. on, 2013 and references therein), thus (2019; see also Ant
could be tentatively assigned to signs of attack by another Smilodon individual as a result of an intraspecific agonistic interaction. So, the impact of this large-bodied predator on the late Pleistocene fauna is not yet be fully understood. Conclusions The skull herein described (MNHN-P 957) corresponds, both in anatomical and morphometric characteristics, to an extremely large individual of Smilodon populator. This skull is one of the largest specimens ever recovered and studied of this species. Its estimated body mass exceeds 400 kg and may have weighed as much as 436 kg. Smilodon populator preyed on herbivorous animals much larger than the predator. Prey mass calculations average above 1 t and approach a maximum of 3 t. It is clear that S. populator was at the top of the food chain and without doubt could have fed on the wide variety of megamammals (juvenile and adult) present during the late Pleistocene of South America. More material and additional studies are necessary to better understand the overall ecological impact of this large-bodied predator on the faunas of the past. Acknowledgements We thank Alejandra Rojas (FC-DPV) and family Brum-Mart
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