Difficulties in assigning trace makers from theropodan bite marks: an example from a young diplodocoid sauropod

Bite marks on the bones of dinosaurs are relatively rare for non‐tyrannosaur dominated faunas, and few have been described in detail. Here, we describe a femur of a young diplodocoid sauropod in the Carnegie Quarry (Late Jurassic Morrison Formation) at Dinosaur National Monument that shows extensive bite marks to the proximal part of the bone. This is the only record of bite marks from this extensive quarry of over 1500 vertebrate elements, making this a most unusual find. Identification of the tracemaker is difficult as multiple large theropods are known from the quarry. Furthermore, we show that subtly different actions of feeding can potentially result in very different spacing of bite marks making matches to tooth patterns in the jaws of potential bite makers very uncertain. Although identification is uncertain, the tracemaker is clearly not a tyrannosaurid, but the selective scrape feeding pattern seen here is similar to the of tyrannosaurid theropods. This technique may be more widely distributed among large carnivorous theropods than previously realized.     

An Additional Baurusuchid from the Cretaceous of Brazil with Evidence of Interspecific Predation among Crocodyliformes

A new Baurusuchidae (Crocodyliformes, Mesoeucrocodylia), Aplestosuchus sordidus, is described based on a nearly complete skeleton collected in deposits of the Adamantina Formation (Bauru Group, Late Cretaceous) of Brazil. The nesting of the new taxon within Baurusuchidae can be ensured based on several exclusive skull features of this clade, such as the quadrate depression, medial approximation of the prefrontals, rostral extension of palatines (not reaching the level of the rostral margin of suborbital fenestrae), cylindrical dorsal portion of palatine bar, ridge on the ectopterygoid-jugal articulation, and supraoccipital with restricted thin transversal exposure in the caudalmost part of the skull roof. A newly proposed phylogeny of Baurusuchidae encompasses A. sordidus and recently described forms, suggesting its sixter-taxon relationship to Baurusuchus albertoi, within Baurusuchinae. Additionally, the remains of a sphagesaurid crocodyliform were preserved in the abdominal cavity of the new baurusuchid. Direct fossil evidence of behavioral interaction among fossil crocodyliforms is rare and mostly restricted to bite marks resulting from predation, as well as possible conspecific male-to-male aggression. This is the first time that a direct and unmistaken evidence of predation between different taxa of this group is recorded as fossils. This discovery confirms that baurusuchids were top predators of their time, with sphagesaurids occupying a lower trophic position, possibly with a more generalist diet.     

A sauropod rib with an embedded theropod tooth: direct evidence for feeding behaviour in the Jehol group,China

Xing, L, Bell, P.R., Currie, P.J., Shibata, M., Tseng, K. & Dong, Z. 2012: A sauropod rib with an embedded theropod tooth: direct evidence for feeding behaviour in the Jehol group, China. Lethaia, Vol. 45, pp. 500–506. A fragmentary rib from the Lower Cretaceous (Barremian) Yixian Formation in northeastern China preserves rare, direct evidence of feeding behaviour by an unidentified theropod. The rib, which comes from the holotype of Dongbetitan, preserves an embedded, broken theropod tooth. Comparison of the tooth with all known theropods from the Yixian Formation suggests that it belongs to a new taxon of medium‐sized theropod. Given the large size difference between the sauropod and the theropod and the absence of reactive bone growth around the tooth, the bite likely occurred post‐mortem during scavenging. Recognition of a new, medium‐sized theropod increases the known diversity of taxa from the Yixian Formation and helps fill a gap in the theropod palaeoecology of that formation, which previously consisted of only small (<2 m) forms. □China, Cretaceous, feeding behaviour, theropod, titanosauriformes, sauropod.     

Mole’s humerus speaks. A rebuttal to Furió 2016

The discovery of small, very well-defined and perfectly preserved tooth marks on the humerus of a mole, Talpa cf. europaea (TE9, Sima del Elefante, Sierra de Atapuerca, Burgos), is extraordinary. To date, no micromammal fossil is known with puncture prints produced by a bite with a clear or delimited morphology that would permit its detailed study. The exceptional character of the finding may raise questions and suspicions about alteration and taphonomic agents. However, we have evidence that both the marks in the mole humerus are due to the action of biting and that this bite corresponds to the dentition of Beremendia fissidens. After all, not only large predators bite, as this article intends to demonstrate.     

Cretaceous small scavengers: feeding traces in tetrapod bones from Patagonia, Argentina

Ecological relationships among fossil vertebrate groups are interpreted based on evidence of modification features and paleopathologies on fossil bones. Here we describe an ichnological assemblage composed of trace fossils on reptile bones, mainly sphenodontids, crocodyliforms and maniraptoran theropods. They all come from La Buitrera, an early Late Cretaceous locality in the Candeleros Formation of northwestern Patagonia, Argentina. This locality is significant because of the abundance of small to medium-sized vertebrates. The abundant ichnological record includes traces on bones, most of them attributable to tetrapods. These latter traces include tooth marks that provde evidence of feeding activities made during the sub-aerial exposure of tetrapod carcasses. Other traces are attributable to arthropods or roots. The totality of evidence provides an uncommon insight into paleoecological aspects of a Late Cretaceous southern ecosystem.     

A tyrannosaur jaw bitten by a confamilial: scavenging or fatal agonism?

Bell, P.R. & Currie, P.J. 2009: A tyrannosaur jaw bitten by a confamilial: scavenging or fatal agonism?. Lethaia, Vol. 43, pp. 278–281. A partial dentary of an adult tyrannosaur from the Dinosaur Park Formation of Alberta, Canada, preserves the embedded tooth of another tyrannosaur within the bone. The specimen’s incompleteness precludes generic identification of either the jaw or the embedded tooth, although Gorgosaurus and/or Daspletosaurus are most likely given the stratigraphic position. The absence of healing around the lesion indicates the bite took place either post‐mortem or within weeks prior to the death of this animal. A post‐mortem bite can be explained by confamilial or cannibalistic scavenging. Alternatively, the bite would represent a perimortem instance of intrafamilial aggression that may have resulted in the death of that animal. An estimated 6053N of bite force was required to produce the bite mark. This specimen provides the best evidence for aggressive peri‐ or post‐mortem confamilial interaction among tyrannosaurs and corroborates previous studies based on inferred tooth marks. □Alberta, behaviour, Campanian, Cretaceous, Dinosaur Park Formation, Theropoda, Tyrannosauridae.     

Taphonomic study of a pleurosternid turtle shell from the Early Cretaceous of Angeac‐Charente,southwest France

The feeding behaviour of crocodilians leads to the formation of particular assemblages by accumulation of carcasses on the bottom or at the edge of water points, in addition to the characteristic marks it leaves on bones, making crocodilians important taphonomic agents. We report here the case of a Pleurosternidae turtle shell, Pleurosternon bullockii (Owen 1842 ), discovered in 2014 in Angeac‐Charente in south‐western France. The shell shows a range of tooth marks left by a crocodyliform. Several elements designate Goniopholis as the most likely author of the marks and support the use of the ‘nutcracker’ technique. Some marks appear to have healed, while others suggest that this P. bullockii individual suffered from shell diseases. Many turtle remains and some ornithomimosaur dinosaur bones from Angeac‐Charente bear marks of a crocodyliform attack.     

Bone-Breaking Bite Force of Basilosaurus isis (Mammalia,Cetacea) from the Late Eocene of Egypt Estimated by Finite Element Analysis

Bite marks suggest that the late Eocence archaeocete whale Basilosaurus isis (Birket Qarun Formation, Egypt) fed upon juveniles of the contemporary basilosaurid Dorudon atrox. Finite element analysis (FEA) of a nearly complete adult cranium of B. isis enables estimates of its bite force and tests the animal’s capabilities for crushing bone. Two loadcases reflect different biting scenarios: 1) an intitial closing phase, with all adductors active and a full condylar reaction force; and 2) a shearing phase, with the posterior temporalis active and minimized condylar force. The latter is considered probable when the jaws were nearly closed because the preserved jaws do not articulate as the molariform teeth come into occulusion. Reaction forces with all muscles active indicate that B. isis maintained relatively greater bite force anteriorly than seen in large crocodilians, and exerted a maximum bite force of at least 16,400 N at its upper P³. Under the shearing scenario with minimized condylar forces, tooth reaction forces could exceed 20,000 N despite lower magnitudes of muscle force. These bite forces at the teeth are consistent with bone indentations on Dorudon crania, reatract-and-shear hypotheses of Basilosaurus bite function, and seizure of prey by anterior teeth as proposed for other archaeocetes. The whale’s bite forces match those estimated for pliosaurus when skull lengths are equalized, suggesting similar tradeoffs of bite function and hydrodynamics. Reaction forces in B. isis were lower than maxima estimated for large crocodylians and carnivorous dinosaurs. However, comparison of force estimates from FEA and regression data indicate that B. isis exerted the largest bite forces yet estimated for any mammal, and greater force than expected from its skull width. Cephalic feeding biomechanics of Basilosaurus isis are thus consistent with habitual predation.     

Functional morphology of bite mechanics in the great barracuda (Sphyraena barracuda)

The great barracuda, Sphyraena barracuda, is a voracious marine predator that captures fish with a swift ram feeding strike. While aspects of its ram feeding kinematics have been examined, an unexamined aspect of their feeding strategy is the bite mechanism used to process prey. Barracuda can attack fish larger than the gape of their jaws, and in order to swallow large prey, can sever their prey into pieces with powerful jaws replete with sharp cutting teeth. Our study examines the functional morphology and biomechanics of 'ram-biting' behavior in great barracuda where the posterior portions of the oral jaws are used to slice through prey. Using fresh fish and preserved museum specimens, we examined the jaw mechanism of an ontogenetic series of barracuda ranging from 20 g to 8.2 kg. Jaw functional morphology was described from dissections of fresh specimens and bite mechanics were determined from jaw morphometrics using the software MandibLever (v3.2). High-speed video of barracuda biting (1500 framess(-1)) revealed that prey are impacted at the corner of the mouth during capture in an orthogonal position where rapid repeated bites and short lateral headshakes result in cutting the prey in two. Predicted dynamic force output of the lower jaw nearly doubles from the tip to the corner of the mouth reaching as high as 58 N in large individuals. A robust palatine bone embedded with large dagger-like teeth opposes the mandible at the rear of the jaws providing for a scissor-like bite capable of shearing through the flesh and bone of its prey.     

Killing in the Pliocene: shark attack on a dolphin from Italy

Abstract: Shark bite marks, including striae, sulci and abrasions, in a well‐preserved fossil dolphin skeleton referred to Astadelphis gastaldii (Cetacea, Delphinidae) from Pliocene sediments of Piedmont (northern Italy), are described in detail. The exceptional combination of a fossil dolphin having a significant part of the skeleton preserved and a large number of bite marks on the bones represents one of the few detailed documentations of shark attack in the past. Most bite marks have been referred to a shark about 4 m long with unserrated teeth, belonging to Cosmopolitodus hastalis, on the basis of their shape and their general disposition on the dolphin skeleton. According to our hypothesis, the shark attacked the dolphin with an initial mortal bite to the abdomen from the rear and right, in a similar way as observed for the living white shark when attacking pinnipeds. A second, less strong, bite was given on the dorsal area when the dolphin, mortally injured, probably rolled to the left. The shark probably released the prey, dead or dying, and other sharks or fishes probably scavenged the torn body of the dolphin.     

A diseased Panthera leo spelaea (Goldfuss 1810) lioness from a forest elephant graveyard in the Late Pleistocene (Eemian) interglacial lake at Neumark-Nord,central Germany

A solitary articulated skeleton of a middle-aged and diseased Panthera leo spelaea lioness from the Eemian interglacial has been found amongst numerous articulated skeletons of Palaeoloxodon antiquus forest elephants, in sediments from a small, shallow lake at Neumark-Nord in central Germany, which has Neanderthal settlements along its shoreline. Several pathologies such as a fibula fracture, arthritis in one of the front legs and a lost canine tooth with associated maxillary inflammation and dissolution made the lioness vulnerable to other predators such as hyenas, whose presence is indicated by their bones, coprolites and many scavenging marks on the elephant skeletons and on a femur from a male lion. The scavenging of hyenas and lions at this site is commonly documented by canine bite marks on the joints of elephant bones. Bite and scratch marks on the ventral vertebral columns and pelvises of two P. antiquus forest elephant skeletons suggest that the intestines and inner organs may have been consumed by large predators, as is commonly the case with modern African lions feeding on elephants. The weak and diseased lioness may possibly have been killed during antagonistic battles between hyenas and lions over their larger prey.     

Visual morphology and feeding behaviour of the daggertooth

Dissection of freshly eaten prey suggests that the North Pacific daggertooth Anotopterus nikparini sever the spine of small prey with a single slashing blow. The bite marks indicate that the immobilized prey is manipulated and swallowed head first. Frequent evidence suggests daggertooth wound prey which are too large to consume whole by cutting off strips of flesh. In either case the wounds consist of a single prominent slash mark on one side and a series of inconspicuous marks on the opposite side. Retinal morphology provides supporting evidence that daggertooths are visual ambush predators that attack prey from below, and should be considered as epipelagic rather than mesopelagic fish. Visual acuity appears to be high, suggesting that frequent attacks on prey too large to be killed immediately is not error, but a well-defined feeding behaviour. The daggertooth may be a significant source of mortality for economically important fish such as all sizes of salmon.     

Mammalian tooth marks on the bones of dinosaurs and other Late Cretaceous vertebrates

Abstract: We describe bones from the Late Cretaceous of Alberta – including bones of large dinosaurs, a femur from the aquatic reptile Champsosaurus, and a dentary from the marsupial Eodelphis– that bear tooth marks made by animals with opposing pairs of teeth. Of the animals known from the Late Cretaceous of North America, only mammals are capable of making such tooth marks. In particular, multituberculates, which have paired upper and lower incisors, are the most likely candidates for the makers of these traces. The traces described here represent the oldest known mammalian tooth marks. Although it is possible that some of these tooth marks represent feeding traces, the tooth marks often penetrate deep into the dense cortices of the bone. This raises the possibility that, much as extant mammals gnaw bone and antler, some Cretaceous mammals may have consumed the bones of dinosaurs and other vertebrates as a source of minerals. However, none of the tooth marks described here resemble the extensive gnaw traces produced by Cenozoic multituberculates or rodents. This suggests that specialized gnawing forms may have been rare or absent in the Late Cretaceous of North America.     

Biology meets engineering: The structural mechanics of fossil and extant shark teeth

The majority of studies on the evolution and function of feeding in sharks have focused primarily on the movement of cranial components and muscle function, with little integration of tooth properties or function. As teeth are subjected to sometimes extreme loads during feeding, they undergo stress, strain, and potential failure. As attributes related to structural strength such as material properties and overall shape may be subjected to natural selection, both prey processing ability and structural parameters must be considered to understand the evolution of shark teeth. In this study, finite element analysis was used to visualize stress distributions of fossil and extant shark teeth during puncture, unidirectional draw (cutting), and holding. Under the loading and boundary conditions here, which are consistent with bite forces of large sharks, shark teeth are structurally strong. Teeth loaded in puncture have localized stress concentrations at the cusp apex that diminish rapidly away from the apex. When loaded in draw and holding, the majority of the teeth show stress concentrations consistent with well designed cantilever beams. Notches result in stress concentration during draw and may serve as a weak point; however they are functionally important for cutting prey during lateral head shaking behavior. As shark teeth are replaced regularly, it is proposed that the frequency of tooth replacement in sharks is driven by tooth wear, not tooth failure. As the tooth tip and cutting edges are worn, the surface areas of these features increase, decreasing the amount of stress produced by the tooth. While this wear will not affect the general structural strength of the tooth, tooth replacement may also serve to keep ahead of damage caused by fatigue that may lead to eventual tooth failure. J. Morphol., 2011. © 2010 Wiley‐Liss, Inc.     

Puncture marks on early African anthropoids

Field studies of living primates have shown that primate predation is a rare event. This must also have been true for past primate communities. In the Fayum Oligocene of Egypt, specimens of all four species of Upper Fossil Wood Zone primates show evidence of tooth puncture marks. Of the four potential groups of primate predators--the snakes, the raptors, the crocodiles, and the primitive carnivores or creodonts--only the crocodiles and the creodonts could have made these puncture marks. When one compares the feeding habits of living crocodiles and mammalian carnivores with the evidence from the Fayum, it appears that the Fayum primates were preyed upon and/or scavenged by mammalian carnivore-like animals. The dismemberment of the Fayum primates by Oligocene predators indicates, in part, why the Fayum fossil material is rarely articulated. Bone damage by predators may well set limits on what bone associations can be discovered in the Fayum even before the bones are scattered and buried by depositional processes.     

Eating with a saw for a jaw: Functional morphology of the jaws and tooth‐whorl in Helicoprion davisii

The recent reexamination of a tooth‐whorl fossil of Helicoprion containing intact jaws shows that the symphyseal tooth‐whorl occupies the entire length of Meckel's cartilage. Here, we use the morphology of the jaws and tooth‐whorl to reconstruct the jaw musculature and develop a biomechanical model of the feeding mechanism in these early Permian predators. The jaw muscles may have generated large bite‐forces; however, the mechanics of the jaws and whorl suggest that Helicoprion was better equipped for feeding on soft‐bodied prey. Hard shelled prey would tend to slip anteriorly from the closing jaws due to the curvature of the tooth‐whorl, lack of cuspate teeth on the palatoquadrate (PQ), and resistance of the prey. When feeding on soft‐bodied prey, deformation of the prey traps prey tissue between the two halves of the PQ and the whorl. The curvature of the tooth‐whorl and position of the exposed teeth relative to the jaw joint results in multiple tooth functions from anterior to posterior tooth that aid in feeding on soft‐bodied prey. Posterior teeth cut and push prey deeper into the oral cavity, while middle teeth pierce and cut, and anterior teeth hook and drag more of the prey into the mouth. Furthermore, the anterior‐posterior edges of the teeth facilitate prey cutting with jaw closure and jaw depression. The paths traveled by each tooth during jaw depression are reminiscent of curved pathways used with slashing weaponry such as swords and knifes. Thus, the jaws and tooth‐whorl may have formed a multifunctional tool for capturing, processing, and transporting prey by cyclic opening and closing of the lower jaw in a sawing fashion. J. Morphol. 276:47–64, 2015. © 2014 Wiley Periodicals, Inc.     

Tooth root morphology as an indicator for dietary specialization in carnivores (Mammalia: Carnivora)

The Carnivora occupy a wide range of feeding niches in concordance with the enormous diversity in their skull and dental form. It is well established that differences in crown morphology are linked to variations in the material properties of the foods ingested and masticated. However, how tooth root form is related to dietary specialization is less well known. In the present study, we investigate the relationship between tooth root morphology and dietary specialization in terrestrial carnivores (canids, felids, hyaenids, and ursids). We specifically address the question of how variation in tooth root surface area is related to bite force potentials as one of the crucial masticatory performance parameters in feeding ecology. We applied computed tomography imaging to reconstruct and quantify dental root surface area in 17 extant carnivore species. Moreover, we computed maximal bite force at several tooth positions based on a dry skull model and assessed the relationship of root surface area to skull size, maximal bite force, food properties, and prey size. We found that postcanine tooth root surface areas corrected for skull size serve as a proxy for bite force potentials and, by extension, dietary specialization in carnivores. Irrespective of taxonomic affinity, species that feed on hard food objects have larger tooth roots than those that eat soft or tough foods. Moreover, carnivores that prey on large animals have larger tooth root surface areas. Our results show that tooth root morphology is a useful indicator of bite force production and allows inferences to be made about dietary ecology in both extant and extinct mammals. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105, 456–471.     

Performance of shark teeth during puncture and draw: implications for the mechanics of cutting

The performance of an organism's feeding apparatus has obvious implications for its fitness and survival. However, the majority of studies that focus on chondrichthyan feeding have largely ignored the role of teeth. Studying the functional morphology of shark teeth not only elucidates the biological role that teeth play in feeding, but also provides insight specifically into the evolution of shark feeding because teeth are often the only structures available in the fossil record. In the present study, we investigate the puncture and draw performance of three general categories of extant teeth, tearing‐type, cutting‐type, and cutting–clutching type, as well as three fossil morphologies, utilizing a universal testing system. Differences in puncturing performance occurred among different prey items, indicating that not all ‘soft’ prey items are alike. The majority of teeth were able to puncture different prey items, and differences in puncture performance also occurred among tooth types; however, few patterns emerged. In some cases, broader triangular teeth were less effective at puncturing than narrow‐cusped teeth. There were no differences between the maximum draw forces and maximum puncture forces. Many of the shark teeth in the present study were not only able to perform draw and puncture equally well, but also many tooth morphologies were functionally equivalent to each other. The findings obtained in the present study lend little support to the belief that shark tooth morphology is a good predictor of biological role. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 271–286.     

Taphonomic evidence of a Paleogene mammalian predator–prey interaction

A taphonomic study has been undertaken on an assemblage of bones and teeth of Isoptychus sp. and Thalerimys fordi (extinct rodent family Theridomyidae) from a single bed in a coastal plain setting, in the Late Eocene (Priabonian) Osborne Member, Headon Hill Formation (Hampshire Basin, UK). The vertebrate fossils show good preservation and do not bear the marks of obvious long distance transport. The two theridomyid species show similar patterns of mortality, element representation and surface modifications, which indicate similar mechanisms of accumulation. There is high mortality of juvenile and old individuals indicating accumulation of the assemblage by the action of attritional not catastrophic agents. The postcranial elements show fragmentary states and very low relative abundances. The vast majority of elongate bones (limb bones, phalanges and metapodials) are broken and exhibit a spiral irregular type of fracture with rounded fracture edges indicating that the bones were broken when they were fresh and have subsequently undergone additional modification. The enamel of most of the cheek teeth and incisors shows localized etching to various degrees and most of the bones show etching. By elimination of other modifying agents the observed etching is attributed to digestive corrosion. Collectively, these data indicate that the majority of the theridomyid individuals were eaten and digested by an animal that could cause high fragmentation during ingestion and with stomach juices of relatively high acidity. Both these features characterize mammalian carnivores. The presence of puncture marks on bones of both theridomyid species and comparisons with sizes of bite marks caused by extant mammalian carnivores suggest predation by a small mammalian carnivore about the size of an arctic fox. The extinct amphicyonid carnivoran Cynodictis cf. lacustris occurs in the same bed and the sizes of some of its teeth match well with the sizes of the puncture marks on the theridomyid bones. A predator–prey interaction is, therefore, deduced for the amphicyonid and the two theridomyid species, thereby reconstructing a small part of the continental Paleogene food chain.     

Leftover prey remains: a new taphonomic mode from the Late Miocene Cerro Azul Formation of Central Argentina

The Cerro Azul Formation (La Pampa Province, Argentina) comprises a rich vertebrate fauna of small mammals dominated by notoungulates and rodents. The fauna pertains to the Late Miocene specifically to the Huayquerian Stage/Age. Taphonomic analysis of micro‐mammals from Estancia Ré locality evidenced that the faunal assemblage was accumulated by the activity of a predator. This assemblage was compared with others from the Cerro Azul Formation in Telén and Caleufú localities (La Pampa Province), previously interpreted as products of predator activities. These microfossil accumulations differ from assemblages attributed to pellets and faeces produced by modern predators (nocturnal and diurnal bird raptors and carnivore mammals). However, due to their anatomical representation, degree of bone breakage and presence of tooth marks, they are interpreted as accumulations of uneaten prey remains discarded by the predator. The predator involved could not be determined with certainty, although the presence of tooth marks in some skeletal remains and the presence of coprolites in Telén and Caleufú suggest that it could be a carnivore mammal. Similarities in the accumulation mechanism, patterns of preservation and sedimentary contexts in the three assemblages support the recognition of a new taphonomic mode, termed ‘leftover prey remains’.