Paleobiology of sabretooth cat Smilodon populator in the Pampean Region (Buenos Aires Province, Argentina) around the Last Glacial Maximum: Insights from carbon and nitrogen stable isotopes in bone collagen
Authors:
Bocherens et al
Abstract:
The sabretooth cat Smilodon populator was the largest felid in South America. It appears in the fossil record in the Early Pleistocene, as an immigrant from North America, and becomes extinct around the Pleistocene–Holocene boundary. The carbon and nitrogen stable isotopic values of collagen were measured for skeletal remains from Smilodon specimens ranging in age from 25 to 10 kyr BP, for the first time in the Pampas region of Argentina. By comparison with similar values obtained on co-eval predators such as Protocyon (large canid) and Panthera onca (jaguar) and a range of potential prey, such as giant ground sloths, glyptodontids, Macrauchenia, Toxodon, equids, cervids, and rodents, it could be established that Smilodon consumed essentially large prey from open landscape, such as Macrauchenia and giant ground sloths during the last 15,000 years of the Late Pleistocene in the Pampa region. It was possibly competing with the large canid Protocyon but the jaguar was apparently feeding on smaller size prey. A more humid climate at the beginning of the Holocene might have been unfavorable to this large predator and could have contributed to its extinction. These results also provide an important insight to understand the ecological processes involved in the Great American Biotic Interchange.
Bone taphonomy of the Schöningen “Spear Horizon South” and its implications for site formation and hominin meat provisioning
Authors:
Starkovich et al
Abstract:
This paper presents the faunal remains from the new excavation area at the Lower Paleolithic site of Schöningen. The focus of the study is on the southern extension of the main find horizon (Spear Horizon South), which includes the layer that yielded the famous Schöningen spears (13 II-4). Taxonomic data corroborate previous studies, that hominins primarily hunted Equus mosbachensis, a large Pleistocene horse. Equid body part representation at the site suggests that the animals were hunted and butchered locally. There is no evidence for density-mediated attrition in the assemblage. Weathering damage is uncommon, though there is ample evidence that carnivores had access to the bone. Carnivore bite sizes were measured and compared to experimental data provided by previous authors. Based on relationships between bite size and carnivore behavior and body size, we conclude that the primary modifying agents were large carnivores (i.e., wolves or saber-toothed cats). Previous studies show that carnivores often had secondary access to the remains, after hominins. Cut marks are commonly arranged haphazardly on the bones. This may indicate that multiple hominins participated in the butchery of horse skeletons, or that they were butchered over the course of hours or days. Cut marks on axial elements are more “orderly,” which probably reflects the physical logistics of orienting one's body in relation to a large carcass. These data differ from sites formed by Middle and Upper Paleolithic hominins, which might suggest that in later times, a system of organized meat provisioning was already in place. Taken together, the faunal evidence from the Spear Horizon South indicates that late Lower Paleolithic hominins using the site understood the behaviors of different prey species, hunted socially to take down large game, and successfully competed with large carnivores on the landscape for primary access to ungulate remains.
The European saber-toothed cat (Homotherium latidens) found in the “Spear Horizon” at Schöningen (Germany)
Authors:
Serangeli et al
Abstract:
The 300,000 year old Lower Paleolithic site Schöningen 13 II-4 became world famous with the discovery of the oldest well-preserved and complete wooden spears. Through ongoing excavations, new archaeological discoveries of scientific importance are still being made from the same archaeological layer where the spears were found. In this context, remains of a rare carnivore species, the European saber-toothed cat (Homotherium latidens), were recovered. Here we present five teeth and one humerus fragment that are unambiguously from two individual saber-toothed cats. The humerus is a unique specimen; it shows evidence of hominin impacts and use as a percussor.
The Homotherium remains from Schöningen are the best documented finds of this species in an archaeological setting and they are amongst the youngest specimens of Homotherium in Europe. The presence of this species as a carnivore competitor would certainly have impacted the lives of late Middle Pleistocene hominins. The discovery illustrates the possible day-to-day challenges that the Schöningen hominins would have faced and suggests that the wooden spears were not necessarily only used for hunting, but possibly also as a weapon for self-defense.
Jaw Function in Smilodon fatalis: A Reevaluation of the Canine Shear-Bite and a Proposal for a New Forelimb-Powered Class 1 Lever Model
Author:
Brown
Abstract:
The jaw function of Smilodon fatalis has long been a source of debate. Although modern-day lions subdue large prey through the use of a suffocating throat bite, the dramatically elongated maxillary canines of S. fatalis suggest an alternative bite mechanism. The current literature favors a “canine shear-bite,” in which the depression of the cranium by the ventral neck flexors assists the mandibular adductors in closing the jaws. Although the model makes intuitive sense and appears to be supported by scientific data, the mechanical feasibility of “neck-powered” biting has not been experimentally demonstrated. In the present study, the computer-assisted manipulation of digitized images of a high-quality replica of an S. fatalis neck and skull shows that a rotation of the cranium by the ventral neck flexors will not result in jaw closure. Instead, the cranium and mandible rotate ventrally together (at the atlantooccipital joint), and the jaws remain in an open configuration. The only manner by which rotation of the cranium can simultaneously result in jaw closure is by an anterior rotation at the temporomandibular joint. Based on this finding, the author proposes a new Class 1 lever mechanism for S. fatalis jaw function. In this model, the mandible is immobilized against the neck of the prey and a dorsally directed force from the extension of the forelimbs rotates the cranium anteriorly at the temporomandibular joint. The maxillary canines pierce the prey’s neck and assist in clamping the ventral neck structures. The model is based on a maximum gape angle of approximately 90° and incorporates a secondary virtual point of rotation located slightly anteroventral to the temporomandibular joint. The Class 1 Lever Model is mechanically feasible, consistent with current data on S. fatalis anatomy and ecology, and may provide a basis for similar studies on other fossil taxa.
Promegantereon ogygia (Felidae, Machairodontinae, Smilodontini) from the Vallesian (late Miocene, MN 10) of Spain: morphological and functional differences in two noncontemporary populations
Authors:
Siliceo et al
Abstract:
We compare two populations of the primitive saber-toothed felid Promegantereon ogygia from the late Miocene (Vallesian, MN 10) of Spain. These populations come from two fossil sites, Batallones-1 and Batallones-3, located very close to each other, within the Cerro de los Batallones complex. The sites show differences in age and in their faunal assemblages, with Batallones-1 being older than Batallones-3. We find that the population from this latter site shows slightly derived characters in both dentition and postcranial elements, which clearly indicate evolution within the Promegantereon lineage, but are not strong enough to support a separation at the species level.
Canine Evolution in Sabretoothed Carnivores: Natural Selection or Sexual Selection?
Authors:
1. Marcela Randau (a)
2. Chris Carbone (a)
3. Samuel T. Turvey (a)
Affiliations:
a. Institute of Zoology, Zoological Society of London, Regent’s Park, London, United Kingdom
Abstract
The remarkable elongated upper canines of extinct sabretoothed carnivorous mammals have been the subject of considerable speculation on their adaptive function, but the absence of living analogues prevents any direct inference about their evolution. We analysed scaling relationships of the upper canines of 20 sabretoothed feliform carnivores (Nimravidae, Barbourofelidae, Machairodontinae), representing both dirk-toothed and scimitar-toothed sabretooth ecomorphs, and 33 non-sabretoothed felids in relation to body size in order to characterize and identify the evolutionary processes driving their development, using the scaling relationships of carnassial teeth in both groups as a control. Carnassials display isometric allometry in both sabretooths and non-sabretooths, supporting their close relationship with meat-slicing, whereas the upper canines of both groups display positive allometry with body size. Whereas there is no statistical difference in allometry of upper canine height between dirk-toothed and scimitar-toothed sabretooth ecomorphs, the significantly stronger positive allometry of upper canine height shown by sabretooths as a whole compared to non-sabretooths reveals that different processes drove canine evolution in these groups. Although sabretoothed canines must still have been effective for prey capture and processing by hypercarnivorous predators, canine morphology in these extinct carnivores was likely to have been driven to a greater extent by sexual selection than in non-sabretooths. Scaling relationships therefore indicate the probable importance of sexual selection in the evolution of the hypertrophied sabretooth anterior dentition.
Comparative Biomechanical Modeling of Metatherian and Placental Saber-Tooths: A Different Kind of Bite for an Extreme Pouched Predator
Authors:
1. Stephen Wroe (a,b)
2. Uphar Chamoli (b,c)
3. William C. H. Parr (b)
4. Philip Clausen (a)
5. Ryan Ridgely (d)
6. Lawrence Witmer (d)
Affiliations:
a. School of Biological, Earth and Environmental Sciences, University of New South Wales, Sydney, NSW, Australia
b. School of Engineering, University of Newcastle, Callaghan, NSW, Australia
c. St. George Clinical School, University of New South Wales, Sydney, NSW, Australia
d. Department of Biomedical Sciences, Heritage College of Osteopathic Medicine, Ohio University, Athens, Ohio, United States of America
Abstract:
Questions surrounding the dramatic morphology of saber-tooths, and the presumably deadly purpose to which it was put, have long excited scholarly and popular attention. Among saber-toothed species, the iconic North American placental, Smilodon fatalis, and the bizarre South American sparassodont, Thylacosmilus atrox, represent extreme forms commonly forwarded as examples of convergent evolution. For S. fatalis, some consensus has been reached on the question of killing behaviour, with most researchers accepting the canine-shear bite hypothesis, wherein both head-depressing and jaw closing musculatures played a role in delivery of the fatal bite. However, whether, or to what degree, T. atrox may have applied a similar approach remains an open question. Here we apply a three-dimensional computational approach to examine convergence in mechanical performance between the two species. We find that, in many respects, the placental S. fatalis (a true felid) was more similar to the metatherian T. atrox than to a conical-toothed cat. In modeling of both saber-tooths we found that jaw-adductor-driven bite forces were low, but that simulations invoking neck musculature revealed less cranio-mandibular stress than in a conical-toothed cat. However, our study also revealed differences between the two saber-tooths likely reflected in the modus operandi of the kill. Jaw-adductor-driven bite forces were extremely weak in T. atrox, and its skull was even better-adapted to resist stress induced by head-depressors. Considered together with the fact that the center of the arc described by the canines was closer to the jaw-joint in Smilodon, our results are consistent with both jaw-closing and neck musculature playing a role in prey dispatch for the placental, as has been previously suggested. However, for T. atrox, we conclude that the jaw-adductors probably played no major part in the killing bite. We propose that the metatherian presents a more complete commitment to the already extreme saber-tooth ‘lifestyle’.
In the period just before they went extinct, the American lions and saber-toothed cats that roamed North America in the late Pleistocene were living well off the fat of the land.
That is the conclusion of the latest study of the microscopic wear patterns on the teeth of these great cats recovered from the La Brea tar pits in southern California. Contrary to previous studies, the analysis did not find any indications that the giant carnivores were having increased trouble finding prey in the period before they went extinct 12,000 years ago.
The results, published on Dec. 26 in the scientific journal PLOS ONE, contradicts previous dental studies and presents a problem for the most popular explanations for the Megafaunal (or Quaternary) extinction when the great cats, mammoths and a number of the largest mammals that existed around the world disappeared.
"The popular theory for the Megafaunal extinction is that either the changing climate at the end of the last Ice Age or human activity – or some combination of the two – killed off most of the large mammals," said Larisa DeSantis, assistant professor of earth and environmental sciences at Vanderbilt, who headed the study. "In the case of the great cats, we expect that it would have been increasingly difficult for them to find prey, especially if had to compete with humans. We know that when food becomes scarce, carnivores like the great cats tend to consume more of the carcasses they kill. If they spent more time chomping on bones, it should cause detectable changes in the wear patterns on their teeth."
In 1993, Blaire Van Valkenburgh at UCLA published a paper on tooth breakage in large carnivores in the late Pleistocene. Analyzing teeth of American lions, saber-tooth cats, dire wolves and coyotes from La Brea, she found that they had approximately three times the number of broken teeth of contemporary predators and concluded, "...these findings suggest that these species utilized carcasses more fully and likely competed more intensely for food than present-day large carnivores."
The latest study uses a new technique, called dental microwear texture analysis (DMTA), developed by co-authorPeter Ungar at the University of Arkansas. It uses a confocal microscope to produce a three-dimensional image of the surface of a tooth. The image is then analyzed for microscopic wear patterns. Chowing down on red meat produces small parallel scratches. Chomping on bones adds larger, deeper pits. Previous methods of dental wear analysis relied on researchers to identify and count these different types of features. DMTA relies on automated software and is considered more accurate because it reduces the possibility of observer bias.
DeSantis and Ungar, with the assistance of Blaine Schubert from East Tennessee State University and Jessica Scott from the University of Arkansas, applied DMTA to the fossil teeth of 15 American lions (Panthera atrox) and 15 saber-tooth cats (Smilodon fatalis) recovered from the La Brea tar pits in Los Angeles.
Their analysis revealed that the wear pattern on the teeth of the American lion most closely resembled those of the present-day cheetah, which actively avoids bones when it feeds. Similarly, the saber-tooth cat's wear pattern most closely resembled those of the present-day African lion, which indulges in some bone crushing when it eats. (This differs from a previous microwear study using a different technique that concluded saber-tooth cats avoided bone to a far greater extent.)
The researchers examined how these patterns changed over time by selecting specimens from tar pits of different ages, ranging from about 35,000 to 11,500 years ago. They did not find any evidence that the two carnivores increased their "utilization" of carcasses throughout this period. If anything, their analysis suggests that the proportion of the carcasses that both kinds of cats consumed actually declined toward the end.
The researchers acknowledge the high rate of tooth breakage reported in the previous study, but they argue that it is more likely the result of increased breakage when taking down prey instead of when feeding.
"Teeth can break from the stress of chewing bone but they can also break when the carnivores take down prey," DeSantis pointed out. Species like hyenas that regularly chew and crack bones of their kills are as likely to break the rear teeth they use for chewing as their front canines. Species like the cheetah, however, which avoid bones during feeding are twice as likely to break canines than rear teeth. This suggests that they are more likely to break canines when pulling down prey.
The researchers report that previous examinations of the jaws of the American lions and saber-tooth cats from this period found that they have more than three times as many broken canines and interpret this as additional evidence that supports their conclusion that most of the excess tooth breakage occurred during capture instead of feeding.
In addition, the researchers argue that the large size of the extinct carnivores and their prey can help explain the large number of broken teeth. The saber-toothed cats were about the size of today's African lion and the American lion was about 25 percent larger. The animals that they preyed upon likely included mammoths, four-ton giant ground sloths and 3,500-pound bison.
(Miocene carnivores of Florida, similar, but not the same as below. img credit)
The fossilized fangs of saber-toothed cats hold clues to how the extinct mammals shared space and food with other large predators 9 million years ago.
Led by the University of Michigan and the Museo Nacional de Ciencias Naturales in Madrid, a team of paleontologists has analyzed the tooth enamel of two species of saber-toothed cats and a bear dog unearthed in geological pits near Madrid. Bear dogs, also extinct, had dog-like teeth and a bear-like body and gait.
The researchers found that the cat species—a leopard-sized Promegantereon ogygia and a much larger, lion-sized Machairodus aphanistus—lived together in a woodland area. They likely hunted the same prey—horses and wild boar. In this habitat, the small saber-toothed cats could have used tree cover to avoid encountering the larger ones. The bear dog hunted antelope in a more open area that overlapped the cats' territory, but was slightly separated.
"These three animals were sympatric—they inhabited the same geographic area at the same time. What they did to coexist was to avoid each other and partition the resources," said Soledad Domingo, a postdoctoral fellow at the U-M Museum of Paleontology and the first author of a paper on the findings published in the Nov. 7 edition of Proceedings of the Royal Society B.
Millions of years before the first humans, the predators lived during the late Miocene Period in a forested area that had patches of grassland. Large carnivores such as these are rare in the fossil record, primarily because plant-eating animals lower on the food chain have outnumbered meat-eaters throughout history.
Cerro de los Batallones, where Domingo has been excavating for the past eight years, is special. Of its nine sites, two are ancient pits with an abundance of meat-eating mammal bones. Agile predators, the researchers say, likely leapt into the natural traps in search of trapped prey.
"These sites offer a unique window to understand life in the past," Domingo said.
To arrive at their findings, the researchers conducted what's called a stable carbon isotope analysis on the animals' teeth. Using a dentist's drill with a diamond bit, they sampled teeth from 69 specimens, including 27 saber-toothed cats and bear dogs. The rest were plant-eaters. They isolated the carbon from the tooth enamel. Using a mass spectrometer, which you could think of as a type of scale, they measured the ratio of the more massive carbon 13 molecules to the less-massive carbon 12. An isotope is a version of an element that contains a different number of neutrons in its nucleus.
[..]
Because the researchers can tell what the herbivores ate, they can surmise what their habitat was like. They believe the animals in this study lived in a wooded area that contained patches of grassland.
The cats showed no significant difference in their stable carbon isotope ratios. That means they likely fed on the same prey and lived in the same habitat, but the posits that the species each fed on different-sized prey.
[...]
"The three largest mammalian predators captured prey in different portions of the habitat, as do coexisting large predators today. So even though none of the species in this 9-million year old ecosystem are still alive today (some of their descendants are), we found evidence for similar ecological interactions as in modern ecosystems," said Catherine Badgley, co-author of the new study and assistant professor of ecology and evolutionary biology.
