Dinosaur diet and feeding
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Dinosaur diets and feeding behavior varied widely throughout the clade, including carnivorous, herbivorous, and omnivorous forms.


Ornithopoda


Hadrosauridae

While studying the chewing methods of hadrosaurids in 2009, the paleontologists Vincent Williams, Paul Barrett, and
Mark Purnell Mark Andrew Purnell is a British palaeontologist, Professor of Palaeobiology at the University of Leicester. Purnell is an expert in conodont biostratigraphy (principally Carboniferous) and conodont palaeobiology, focussing especially on atte ...
found that hadrosaurs likely grazed on horsetails and vegetation close to the ground, rather than browsing higher-growing leaves and twigs. This conclusion was based upon the evenness of scratches on hadrosaur teeth, which suggested the hadrosaur used the same series of jaw motions over and over again. As a result, the study determined that the hadrosaur diet probably consisted of leaves, and lacked the bulkier items such as twigs or stems, which might have required a different chewing method and created different wear patterns. However, Purnell said these conclusions were less secure than the more conclusive evidence regarding the motion of teeth while chewing. The hypothesis that hadrosaurs were likely grazers rather than browsers appears to contradict previous findings from preserved stomach contents found in the fossilized guts in previous hadrosaur studies. The most recent such study before the publication of the Purnell study was conducted in 2008, when a team led by
University of Colorado at Boulder The University of Colorado Boulder (CU Boulder, CU, or Colorado) is a public research university in Boulder, Colorado. Founded in 1876, five months before Colorado became a state, it is the flagship university of the University of Colorado sys ...
graduate student Justin S. Tweet found a homogeneous accumulation of millimeter-scale leaf fragments in the gut region of a well-preserved partially grown ''
Brachylophosaurus ''Brachylophosaurus'' ( or ; meaning "short-crested lizard", Greek ''brachys'' = short + ''lophos'' = crest + ''sauros'' = lizard, referring to its small crest) was a mid-sized member of the hadrosaurid family of dinosaurs. It is known from sev ...
''. As a result of that finding, Tweet concluded in September 2008 that the animal was likely a browser, not a grazer. In response to such findings, Purnell said preserved stomach contents are questionable because they do not necessarily represent the usual diet of the animal. The issue remains a subject of debate.This information comes from the aforementioned Alan Boyle source from June 29, 2009. However, this specific information is not included in the body of the article, but rather a response by Boyle to comments in the article. Since the comments were written by Boyle himself, and since they cite information he received specifically from Purnell, they are as legitimate a source of information as the article itself.
Coprolite A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name is de ...
s (fossilized droppings) of some Late
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
hadrosaurs show that the animals sometimes deliberately ate rotting wood. Wood itself is not nutritious, but decomposing wood would have contained fungi, decomposed wood material and
detritus In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts commun ...
-eating
invertebrate Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s, all of which would have been nutritious.


Theropoda

Tyrannosaur Tyrannosauroidea (meaning 'tyrant lizard forms') is a superfamily (or clade) of coelurosaurian theropod dinosaurs that includes the family Tyrannosauridae as well as more basal relatives. Tyrannosauroids lived on the Laurasian supercontinent be ...
tooth marks are the most commonly preserved feeding traces of carnivorous dinosaurs. It is usually not possible to identify tooth marks on bone made by small predatory dinosaurs due to similarities in the denticles on their teeth. However, there are exceptions, like an
ornithomimid Ornithomimidae (meaning "bird-mimics") is a family of theropod dinosaurs which bore a superficial resemblance to modern ostriches. Ornithomimids were fast, omnivorous or herbivorous dinosaurs known mainly from the Late Cretaceous Period of Laura ...
caudal Caudal may refer to: Anatomy * Caudal (anatomical term) (from Latin ''cauda''; tail), used to describe how close something is to the trailing end of an organism * Caudal artery, the portion of the dorsal aorta of a vertebrate that passes into the ...
vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
that has tooth drag marks attributed to ''Saurornitholestes'' and a partial ''
Troodon ''Troodon'' ( ; ''Troödon'' in older sources) is a wastebasket taxon and a dubious genus of relatively small, bird-like dinosaurs known definitively from the Campanian age of the Late Cretaceous period (about 77  mya). It includes at least ...
'' skeleton with preserved puncture marks. Small bones of small theropods that were preyed upon by larger ones may have been swallowed whole and digested frequently enough to affect their abundance in the fossil record. In 2001, Bruce Rothschild and others published a study examining evidence for
stress fractures A stress fracture is a fatigue-induced bone fracture caused by repeated stress over time. Instead of resulting from a single severe impact, stress fractures are the result of accumulated injury from repeated submaximal loading, such as running or ...
in
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaurs and the implications for their behavior. Since stress fractures are caused by repeated trauma they are more likely to be a result of the animal's behavior than fractures obtained during a single injurious event. The distribution of stress fractures also has behavioral significance. Stress fractures to the hand are more likely to result from predatory behavior since injuries to the feet could be obtained while running or migrating. In order to identify stress fractures occurring in the feet of ''
Allosaurus ''Allosaurus'' () is a genus of large carnosaurian theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic epoch (Kimmeridgian to late Tithonian). The name "''Allosaurus''" means "different lizard" alluding to ...
'' specifically due to predatory behavior, the researchers checked to see if the toes which bore the greatest stress while in motion also had the greatest percentage of stress fractures. Since the lower end of the third
metatarsal The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the med ...
would contact the ground first while a theropod was running it would have borne the most stress and should be most predisposed to suffer stress fractures. The lack of such a bias in the examined fossils indicates an origin for the stress fractures from a source other than running, like interaction with prey. They suggested that such injuries could occur as a result of the allosaur trying to hold struggling prey with its feet. Contact with struggling prey is also the likely cause of tendon avulsions found in the forelimbs of ''
Allosaurus ''Allosaurus'' () is a genus of large carnosaurian theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic epoch (Kimmeridgian to late Tithonian). The name "''Allosaurus''" means "different lizard" alluding to ...
'' and ''
Tyrannosaurus ''Tyrannosaurus'' is a genus of large theropoda, theropod dinosaur. The species ''Tyrannosaurus rex'' (''rex'' meaning "king" in Latin), often called ''T. rex'' or colloquially ''T-Rex'', is one of the best represented theropods. ''Tyrannosa ...
''. The authors concluded that the presence of stress fractures provide evidence for "very active" predation-based diets rather than obligate scavenging.


Tyrannosauridae

Tyrannosaur Tyrannosauroidea (meaning 'tyrant lizard forms') is a superfamily (or clade) of coelurosaurian theropod dinosaurs that includes the family Tyrannosauridae as well as more basal relatives. Tyrannosauroids lived on the Laurasian supercontinent be ...
tooth marks are the most commonly preserved feeding traces of carnivorous dinosaurs. They have been reported from
ceratopsians Ceratopsia or Ceratopia ( or ; Greek: "horned faces") is a group of herbivorous, beaked dinosaurs that thrived in what are now North America, Europe, and Asia, during the Cretaceous Period, although ancestral forms lived earlier, in the Jurassic. ...
,
hadrosaurs Hadrosaurids (), or duck-billed dinosaurs, are members of the ornithischian family Hadrosauridae. This group is known as the duck-billed dinosaurs for the flat duck-bill appearance of the bones in their snouts. The ornithopod family, which includ ...
and other
tyrannosaurs Tyrannosauroidea (meaning 'tyrant lizard forms') is a superfamily (or clade) of coelurosaurian theropod dinosaurs that includes the family Tyrannosauridae as well as more basal relatives. Tyrannosauroids lived on the Laurasian supercontinent be ...
.
Tyrannosaurid Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to thirteen genera, including the eponymous ''Tyrannosaurus''. The exact number of genera ...
bones with tooth marks represent about 2% of known fossils with preserved tooth marks.
Tyrannosaurid Tyrannosauridae (or tyrannosaurids, meaning "tyrant lizards") is a family of coelurosaurian theropod dinosaurs that comprises two subfamilies containing up to thirteen genera, including the eponymous ''Tyrannosaurus''. The exact number of genera ...
teeth were used as holdfasts for pulling
meat Meat is animal flesh that is eaten as food. Humans have hunted, farmed, and scavenged animals for meat since prehistoric times. The establishment of settlements in the Neolithic Revolution allowed the domestication of animals such as chic ...
off a body, rather than
knife A knife ( : knives; from Old Norse 'knife, dirk') is a tool or weapon with a cutting edge or blade, usually attached to a handle or hilt. One of the earliest tools used by humanity, knives appeared at least 2.5 million years ago, as evidenced ...
-like cutting functions. Tooth wear patterns hint that complex head shaking behaviors may have been involved in tyrannosaur feeding. In 2001, Bruce Rothschild and others published a study examining evidence for
stress fractures A stress fracture is a fatigue-induced bone fracture caused by repeated stress over time. Instead of resulting from a single severe impact, stress fractures are the result of accumulated injury from repeated submaximal loading, such as running or ...
and
tendon avulsions An avulsion fracture is a bone fracture which occurs when a fragment of bone tears away from the main mass of bone as a result of physical trauma. This can occur at the ligament by the application of forces external to the body (such as a fa ...
in
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaurs and the implications for their behavior. Since stress fractures are caused by repeated trauma they are more likely to be a result of the animal's behavior than fractures obtained during a single injurious event. The distribution of stress fractures also has behavioral significance. Stress fractures to the hand are more likely to result from predatory behavior since injuries to the feet could be obtained while running or migrating. The researchers concluded that contact with struggling prey is the likely cause of a tendon avulsion found in the forelimb of the Tyrannosaurus specimen Sue. The authors concluded that the presence of stress fractures in theropods provide evidence for "very active" predation-based diets rather than obligate scavenging.


''Saurornitholestes'' preyed upon by juvenile tyrannosaur

A. R. Jacobsen published a description of a
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
referred to ''
Saurornitholestes ''Saurornitholestes'' ("lizard-bird thief") is a genus of carnivorous dromaeosaurid theropod dinosaur from the late Cretaceous of Canada (Alberta) and the United States (Montana, New Mexico, Alabama, North Carolina, and South Carolina). Two spec ...
'' with tooth marks. The dentary is about 12 cm long and preserves fifteen tooth positions; ten of these have teeth, with five of those teeth fully erupted and intact, two broken but functional as evidenced by the presence of wearfacets, and three only partially erupted. Three toothmarks were visible on the lingual surface of the dentary. Two of the three marks are series of grooves made by the serrations on the maker's teeth. The first consists of 6-7 parallel grooves within a 4 × 1.3 mm area beneath the
alveolus Alveolus (; pl. alveoli, adj. alveolar) is a general anatomical term for a concave cavity or pit. Uses in anatomy and zoology * Pulmonary alveolus, an air sac in the lungs ** Alveolar cell or pneumocyte ** Alveolar duct ** Alveolar macrophage * Mam ...
of the third tooth and angled at 45° to the dentary's longitudinal axis. The
striation Striations means a series of ridges, furrows or linear marks, and is used in several ways: * Glacial striation * Striation (fatigue), in material * Striation (geology), a ''striation'' as a result of a geological fault * Striation Valley, in Anta ...
s are between .37 mm and .40 mm thick with
cuboidal Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellu ...
cross-sections. The second tooth mark lies between the fifth and sixth
alveoli Alveolus (; pl. alveoli, adj. alveolar) is a general anatomical term for a concave cavity or pit. Uses in anatomy and zoology * Pulmonary alveolus, an air sac in the lungs ** Alveolar cell or pneumocyte ** Alveolar duct ** Alveolar macrophage * ...
and consists of two smaller grooves separated 1.8 and 1.6 mm respectively from a larger central groove, with a V-shaped groove beneath it at an angle of 60° to the longitudinal axis of the jaw. The third mark consists of four parallel grooves in a 2 × 2 mm area on the seventh tooth oriented at 90° to the longitudinal axis of the tooth. The shapes of the preserved serrations are too different from those of ''Saurornitholestes'' for the marks to be the result of injuries incurred during
intraspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organ ...
face biting behaviors. Although the right shape for ''
Dromaeosaurus ''Dromaeosaurus'' (, "running lizard") is a genus of dromaeosaurid theropod dinosaur which lived during the Late Cretaceous period (middle late Campanian and Maastrichtian), sometime between 80 and 69 million years ago, in Alberta, Canada and the ...
'' tooth serrations, the preserved marks are too coarse to have been left by that genus. Although a specific identification cannot be made, the most likely perpetrator would be a juvenile individual of one of the
Dinosaur Park Formation The Dinosaur Park Formation is the uppermost member of the Belly River Group (also known as the Judith River Group), a major geologic unit in southern Alberta. It was deposited during the Campanian stage of the Late Cretaceous, between about 76. ...
's tyrannosaurids, like ''
Gorgosaurus ''Gorgosaurus'' ( ; ) is a genus of tyrannosaurid theropod dinosaur that lived in western North America during the Late Cretaceous Period (Campanian), between about 76.6 and 75.1 million years ago. Fossil remains have been found in the Can ...
'', ''
Daspletosaurus ''Daspletosaurus'' ( ; meaning "frightful lizard") is a genus of tyrannosaurid dinosaur that lived in Laramidia between about 79.5 and 74 million years ago, during the Late Cretaceous Period. The genus ''Daspletosaurus'' contains three species ...
'', or ''
Aublysodon ''Aublysodon'' (“backwards-flowing tooth") is a genus of carnivorous dinosaurs known only from the Judith River Formation in Montana, which has been dated to the late Campanian age of the late Cretaceous period (about 75 million years ago). The ...
''. Jacobsen determined that all of the marks on the jawbone were left by the same animal because the serration marks all share the same morphology.


Dromaeosauridae


''Saurornitholestes''

An ornithomimid caudal vertebra has been discovered that has tooth drag marks attributed to ''Saurornitholestes''.


''Deinonychus''

''
Deinonychus ''Deinonychus'' ( ; ) is a genus of dromaeosaurid theropod dinosaur with one described species, ''Deinonychus antirrhopus''. This species, which could grow up to long, lived during the early Cretaceous Period, about 115–108 million ye ...
'', along with other similar
dromaeosaur Dromaeosauridae () is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek ('), meaning ...
s, were suggested to kill prey in a similar manner to modern
accipitrid The Accipitridae is one of the three families within the order Accipitriformes, and is a family of small to large birds with strongly hooked bills and variable morphology based on diet. They feed on a range of prey items from insects to medium-si ...
birds of prey Birds of prey or predatory birds, also known as raptors, are hypercarnivorous bird species that actively hunt and feed on other vertebrates (mainly mammals, reptiles and other smaller birds). In addition to speed and strength, these predators ...
, due to the similarities of their grasping
talons A claw is a curved, pointed appendage found at the end of a toe or finger in most amniotes (mammals, reptiles, birds). Some invertebrates such as beetles and spiders have somewhat similar fine, hooked structures at the end of the leg or tarsus ...
. The ''Deinonychus'' would pin the prey down with its own weight, restrain it with its hind feet talons, and
dismember Dismemberment is the act of cutting, ripping, tearing, pulling, wrenching or otherwise disconnecting the limbs from a living or dead being. It has been practiced upon human beings as a form of capital punishment, especially in connection with ...
it with its mouth.


Ceratopsia


Ceratopsidae

In 1966
John Ostrom John Harold Ostrom (February 18, 1928 – July 16, 2005) was an American paleontologist who revolutionized modern understanding of dinosaurs in the 1960s. As first proposed by Thomas Henry Huxley in the 1860s, Ostrom showed that dinosaurs were ...
postulated that the diet of late
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
chasmosaurs such as ''
Triceratops ''Triceratops'' ( ; ) is a genus of herbivore, herbivorous Chasmosaurinae, chasmosaurine Ceratopsidae, ceratopsid dinosaur that first appeared during the late Maastrichtian stage of the Late Cretaceous Period (geology), period, about 68 m ...
'' and ''
Torosaurus ''Torosaurus'' ("perforated lizard", in reference to the large openings in its frill) is a genus of herbivorous ceratopsid dinosaur that lived during the late Maastrichtian stage of the Cretaceous period, between 68 and 66 million years ago, tho ...
'' fed on very resistant and fibrous materials like the fronds of
cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk (botany), trunk with a crown (botany), crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants o ...
or palm plants. By extension, all Ceratopsids had a shearing dentition and efficient, powerful jaw mechanics that allowed them to feed on tough vegetation. Mallon and Anderson postulated that Ankylosaurs and Ceratopsids may have partitioned the herb layer in the Dinosaur Park Formation, or that Ceratopsid feeding height was slightly higher. As well as suggesting that the Ornithopods might have made room for the passing Ceratopsid herds by rising up to avoid ecological competition. Notably, they could not distinguish centrosaurine and chasmosaurine ecology apart, though it is likely present as in other megaherbivore clades.Mallon JC, Anderson JS (2013) Skull Ecomorphology of Megaherbivorous Dinosaurs from the Dinosaur Park Formation (Upper Campanian) of Alberta, Canada. PLoS ONE 8(7): e67182. https://doi.org/10.1371/journal.pone.0067182 The results of a NPMANOVA analysis supported the suggestion that Ceratopsids had the strongest bite force of each of the megaherbivore groups, and able to process the toughest plants available.Mallon, J.C. Competition structured a Late Cretaceous megaherbivorous dinosaur assemblage. Sci Rep 9, 15447 (2019). https://doi.org/10.1038/s41598-019-51709-5 Like Ornithopods and unlike all other dinosaurs, Ceratopsians possessed dental batteries that may have been attributable to their success. In 2019, Mallon noted that stratigraphic overlap in the Dinosaur Park Formation between subfamilial taxa was limited, further supporting chasmosaurine and centrosaurine preferencial differences.


See also

*
Bird food Bird food or bird seed is food (often varieties of seeds, nuts, and/or dried fruits) intended for consumption by wild and domestic birds. While most bird food is fed to commercial fowl (such as chicken or turkey), bird food is also used to fee ...
*
Bird food plants Bird food plants are certain trees, shrubs and herbaceous plants bearing fruits which afford food for birds. These have been discovered by observation, and by the scientific examination of the contents of birds' stomachs. By planting those species, ...
*
Bird feeding Bird feeding is the activity of feeding wild birds, often by means of bird feeders. With a recorded history dating to the 6th century, the feeding of wild birds has been encouraged and celebrated in the United States and United Kingdom, with it b ...
* Dietary biology of the golden eagle


Footnotes

{{Reflist, 3, refs= "Introduction," in Abler (2001). Page 84. "Introduction," in Rothschild, et al. (2001); pages 331-332. "Abstract," Rothschild, et al., et al. (2001); page 331. "Introduction," Rothschild, et al. (2001); page 332. "Discussion," Rothschild, et al. (2001); page 334. "Abstract," Jacobsen (2001). Page 58. "Introduction," Jacobsen (2001). Page 59. "Discussion," Jacobsen (2001). Page 61. "Description," Jacobsen (2001). Page 59. "Description," Jacobsen (2001). Page 60. "Discussion," Jacobsen (2001). Page 60.


References

* Abler, W.L. 2001. A kerf-and-drill model of tyrannosaur tooth serrations. p. 84-89. In: ''Mesozoic Vertebrate Life''. Ed.s Tanke, D. H., Carpenter, K., Skrepnick, M. W. Indiana University Press. * Jacobsen, A.R. 2001. Tooth-marked small theropod bone: An extremely rare trace. p. 58-63. In: ''Mesozoic Vertebrate Life''. Ed.s Tanke, D. H., Carpenter, K., Skrepnick, M. W. Indiana University Press. * Rothschild, B., Tanke, D. H., and Ford, T. L., 2001, Theropod stress fractures and tendon avulsions as a clue to activity: In: Mesozoic Vertebrate Life, edited by Tanke, D. H., and Carpenter, K., Indiana University Press, p. 331-336. Dinosaur paleobiology