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A critical re-appraisal of the obligate scavenging hypothesis for Tyrannosaurus rex and other tyrant dinosaurs

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AI-generated Abstract

The giant Cretaceous dinosaur Tyrannosaurus rex and its relatives have long been debated in terms of their ecological behavior, specifically their role as scavengers or predators. This paper critically re-evaluates the obligate scavenging hypothesis, discussing the ecological context of carnivory among modern analogs and providing insights into the anatomical adaptations of tyrannosaurids that may indicate a predatory lifestyle. The study highlights the duality of scavenging and predation as ecological strategies in large carnivorous animals, challenging traditional views by presenting evidence of mechanical adaptations that would support a predatory role for Tyrannosaurus rex.

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a eS  - Figure 20.4 plots the ratio of the tibia length to 5 femur length (T/F) against femur length (used as a proxy for body size) for various theropod dinosaurs. Among nonavian theropods, there is a decrease in T/F as femur length increases (as noted by Gatesy 1991; Holtz 1995; and Carrano 1999), The hypothesis that tyrannosaurids have tibiae that are only as long ot shorter than their femora is not supported by the data: this is true for larger specimens, but not for smaller individuals of tyrant dinosaurs. Indeed, the smallest tyrannosaurs had ‘T/F values as high as those of ornithomimosaur: (generally regarded as among the swiftest of dinosaurs; Barsbold and Os- mélska 1990) of the same femoral length and higher than those of other nontyrannosaur, nonornithomimosaur theropods of the same body size. In fact, even the largest individual of Tyrannosaurus rex examined (FMNH PR2081) had a'T/F value (0.86) equal to that of a much smaller Herrerasau- rus ischigualastensis (PVL 2566). If T/F values less than 1.0 independent of other features of the anatomy were sufficient to exclude tyrannosaurs from the possibility of predation, then many other theropods (including allosauroids, Ceratosaurus, and even herrerasaurids with femora as small as 243 mm long) would also be excluded from predation.  In examining absolute limb lengths in theropods, it is found that both
a eS - Figure 20.4 plots the ratio of the tibia length to 5 femur length (T/F) against femur length (used as a proxy for body size) for various theropod dinosaurs. Among nonavian theropods, there is a decrease in T/F as femur length increases (as noted by Gatesy 1991; Holtz 1995; and Carrano 1999), The hypothesis that tyrannosaurids have tibiae that are only as long ot shorter than their femora is not supported by the data: this is true for larger specimens, but not for smaller individuals of tyrant dinosaurs. Indeed, the smallest tyrannosaurs had ‘T/F values as high as those of ornithomimosaur: (generally regarded as among the swiftest of dinosaurs; Barsbold and Os- mélska 1990) of the same femoral length and higher than those of other nontyrannosaur, nonornithomimosaur theropods of the same body size. In fact, even the largest individual of Tyrannosaurus rex examined (FMNH PR2081) had a'T/F value (0.86) equal to that of a much smaller Herrerasau- rus ischigualastensis (PVL 2566). If T/F values less than 1.0 independent of other features of the anatomy were sufficient to exclude tyrannosaurs from the possibility of predation, then many other theropods (including allosauroids, Ceratosaurus, and even herrerasaurids with femora as small as 243 mm long) would also be excluded from predation. In examining absolute limb lengths in theropods, it is found that both
From this evidence, it is clear that tyrannosaurids would cover more ground for the same angle of femur motion than hadrosaurids and ceratop- sids of the same body size. In other words, they would travel further per unit of time (i.e., would be faster) than their potential prey. Again, as before, this does not require a suspended phase on the part of the tyrannosaurid. Limb proportions do not dismiss the possibility that tyrannosaurids could overtake contemporary herbivores, and indeed they are consistent with a model in which tyrant dinosaurs were faster than their potential prey. (Note that the above discussion does not take into account the great disparity between forelimb and hind limb length in ceratopsians. This might indicate that cera- topsids were necessarily slower than a fully bipedal dinosaur of similar hind limb proportions, unless the forelimbs moved with faster steps than the hind limbs in order to keep pace.) To put it another way, if (as Horner argues) the low T/F values of Tyrannosaurus and its kin would hinder them from run- ning to catch their prey, the equally low or even lower T/F values of hadro- saurids and ceratopsids would even more greatly hinder the ability of these ornithischians in running from a pursuing tyrannosaurid.  Additionally, tyrannosaurids possessed an arctometatarsus, a modifica-  Rowe asf Ebay Pat Plead! Lotnaacs nee tas] eee ete wie bene nem albanien af
From this evidence, it is clear that tyrannosaurids would cover more ground for the same angle of femur motion than hadrosaurids and ceratop- sids of the same body size. In other words, they would travel further per unit of time (i.e., would be faster) than their potential prey. Again, as before, this does not require a suspended phase on the part of the tyrannosaurid. Limb proportions do not dismiss the possibility that tyrannosaurids could overtake contemporary herbivores, and indeed they are consistent with a model in which tyrant dinosaurs were faster than their potential prey. (Note that the above discussion does not take into account the great disparity between forelimb and hind limb length in ceratopsians. This might indicate that cera- topsids were necessarily slower than a fully bipedal dinosaur of similar hind limb proportions, unless the forelimbs moved with faster steps than the hind limbs in order to keep pace.) To put it another way, if (as Horner argues) the low T/F values of Tyrannosaurus and its kin would hinder them from run- ning to catch their prey, the equally low or even lower T/F values of hadro- saurids and ceratopsids would even more greatly hinder the ability of these ornithischians in running from a pursuing tyrannosaurid. Additionally, tyrannosaurids possessed an arctometatarsus, a modifica- Rowe asf Ebay Pat Plead! Lotnaacs nee tas] eee ete wie bene nem albanien af
the distribution of forces of locomotion (Holtz 1995) (Fig. 20.10). Recent analyses by Snively and Russell (2002, 2003) and Snively et al. (2004) have demonstrated that this adaptation might additionally serve to resist tor- sional forces, allowing them to turn more rapidly than they might other- wise without risking mechanical failure of their narrow metatarsi. Ceratop- sids and hadrosaurids lack this adaptation, or other morphological correlates with more cursorial function (Coombs 1978; Carrano 1999).
the distribution of forces of locomotion (Holtz 1995) (Fig. 20.10). Recent analyses by Snively and Russell (2002, 2003) and Snively et al. (2004) have demonstrated that this adaptation might additionally serve to resist tor- sional forces, allowing them to turn more rapidly than they might other- wise without risking mechanical failure of their narrow metatarsi. Ceratop- sids and hadrosaurids lack this adaptation, or other morphological correlates with more cursorial function (Coombs 1978; Carrano 1999).
Van Valkenburgh and Ruff (1987) and Farlow et al. (1991) converted tooth measurements into bending strength indices using beam theory. The present analysis followed the calculations of Farlow et al. and used a rect- angular cross-sectional model rather than an oval model for ziphodont and tyrannosaurid teeth. These values do not represent actual strength values, but rather indices comparing the relative resistance of teeth to loads of unit values.
Van Valkenburgh and Ruff (1987) and Farlow et al. (1991) converted tooth measurements into bending strength indices using beam theory. The present analysis followed the calculations of Farlow et al. and used a rect- angular cross-sectional model rather than an oval model for ziphodont and tyrannosaurid teeth. These values do not represent actual strength values, but rather indices comparing the relative resistance of teeth to loads of unit values.
et al. 1996; Carpenter 2000; Happ this volume) and by the Frenchman coprolite (Chin et al. 1998). However, the incrassate teeth of tyrannosaurs may have also functioned during prey capture and dispatch. Horner and colleagues (Horner 1994, 1997, Horner and Lessem 1993; Horner and Dobb 1997) argue that because the forelimbs of tyrannosaurs could not be used to capture prey, the only other likely implement to seize a victim would be the jaws—a view I find reasonable.
et al. 1996; Carpenter 2000; Happ this volume) and by the Frenchman coprolite (Chin et al. 1998). However, the incrassate teeth of tyrannosaurs may have also functioned during prey capture and dispatch. Horner and colleagues (Horner 1994, 1997, Horner and Lessem 1993; Horner and Dobb 1997) argue that because the forelimbs of tyrannosaurs could not be used to capture prey, the only other likely implement to seize a victim would be the jaws—a view I find reasonable.

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