Durophagy

Last updated
Jaws and teeth of Rhinoptera bonasus (cownose ray) Rhinoptera bonasus (cownose stingray teeth & mouthparts) 1.jpg
Jaws and teeth of Rhinoptera bonasus (cownose ray)

Durophagy is the eating behavior of animals that consume hard-shelled or exoskeleton-bearing organisms, such as corals, shelled mollusks, or crabs. [1] It is mostly used to describe fish, but is also used when describing reptiles, [2] including fossil turtles, [3] placodonts and invertebrates, as well as "bone-crushing" mammalian carnivores such as hyenas. [4] Durophagy requires special adaptions, such as blunt, strong teeth and a heavy jaw. [5] Bite force is necessary to overcome the physical constraints of consuming more durable prey and gain a competitive advantage over other organisms by gaining access to more diverse or exclusive food resources earlier in life. [6] Those with greater bite forces require less time to consume certain prey items as a greater bite force can increase the net rate of energy intake when foraging and enhance fitness in durophagous species.

Contents

In the order Carnivora there are two dietary categories of durophagy; bonecrackers and bamboo eaters. Bonecrackers are exemplified by hyenas and borophagines, while bamboo eaters are primarily the giant panda and the red panda. Both have developed similar cranial morphology. However, the mandible morphology reveals more about their dietary resources. Both have a raised and dome-like anterior cranium, enlarged areas for the attachment of masticatory muscles, enlarged premolars, and reinforced tooth enamel. Bamboo eaters tend to have larger mandibles, while bonecrackers have more sophisticated premolars. [7]

Teleost fish (Teleostei)

This Piaractus brachypomus skull exhibits the durophagous dentition that enables it to crack hard nut shells, instead of animal prey. Piaractus brachypomus - squelette mnhn Paris.JPG
This Piaractus brachypomus skull exhibits the durophagous dentition that enables it to crack hard nut shells, instead of animal prey.

Many Teleosts, for example the Atlantic wolffish, exhibit durophagous behaviour and crush hard prey with their appropriately adapted jaws and teeth. Other fish use of their pharyngeal teeth, with the aid of their protrusible mouth for enabling the grabbing of prey to draw it into their mouth. The pharyngeal jaws found in more derived teleosts are more powerful, with left and right ceratobranchials fusing to become one lower jaw and the pharyngeal branchial fusing to create a large upper jaw that articulates with the neurocranium. They also have developed a muscle, the hypertrophied pharyngeal, to crush prey with help from the molariform pharyngeal teeth. This permits the consumption of hard-shelled prey. [8] [9]

As in the Carnivora however, some largely herbivorous or omnivorous Teleost fishes too, exhibit durophagous behaviour in feeding on plant foods, in that they crack the hard stones of fruit that fall into their water: spectacular examples include relatives of the carnivorous piranhas such species include Piaractus brachypomus and Piaractus mesopotamicus.

Triggerfish (Balistidae)

Triggerfish have jaws that contain a row of four teeth on either side, the upper jaw containing an additional set of six plate-like pharyngeal teeth. Triggerfish do not have jaw protrusion and there are enlarged jaw adductor muscles for extra power to crush the protective shells and spines of their prey. [8]

Cichlids (Cichlidae)

Mollusk shells can be crushed to expose soft parts of the prey to digestive juices or the soft parts can be removed from the shell. Species that crush shells are defined by their large and greatly thickened pharyngeal bones. These bones have flat-crowned teeth and along with their dorsal fellows drawn by powerful muscles, create a crushing mill. The jaws are less derived as they are for just for picking up relatively large objects. [10] [11]

The second method cichlids use is to crush mollusk shells between powerful jaws armed with suitable teeth. Cichlids possess short, broad jaws armed with an outer row of relatively few, strong and conical teeth and several inner rows of finer, also conical teeth. Along with these features are the presence of foreshortening of the skull and development of particularly powerful mandibular adductor muscles. To feed with this type of structure the fish can protrude its mouth ventrally to permit muscles to be seized by the jaws and the mouth then is retracted rapidly so the hard-toothed jaws crush the mollusk shell with the resulting force. A series of biting movements completes the process and the shell fragments are spat out and the soft body is swallowed. [10] [11]

Chondrichthyans

Within the chondrichthyans, horn sharks (Heterodontidae), some rays (Myliobatidae) and chimeras (Holocephali) exhibit durophagous behaviour. They have adaptations to allow for this including stout flattened teeth, hypertrophied jaw adductor muscles and robust jaws to feed on hard prey such as crustaceans and molluscs. Sharks that crush prey have teeth with small, low rounded cusps that are numerous per row, or are molariform. The molariform teeth are smoothly rounded, lack cusps, and there are numerous teeth per row. [8]

Horn sharks (Heterodontiformes)

Horn sharks have molariform teeth. The anterior teeth are pointed and are used for grasping while the posterior teeth are molariform and are used for crushing. Horn sharks feed primarily on limpets, bivalve molluscs and blue crabs. [8]

Bonnethead shark (Sphyrna tiburo)

The bonnethead shark Sphyrna tiburo uses ram feeding to capture crab, shrimp and fish which are placed between the molariform teeth where they are crushed. This species also uses suction to transport prey to the esophagus for swallowing. By combining durophagous characteristics with altered kinematic and motor patterns, bonnethead sharks can prey on hard shelled animals. This characteristic distinguishes prey crushing from simply biting, which is a behaviour exhibited by elasmobranchs. While bonnethead sharks feed almost exclusively on crabs, they have the same tooth structure as the Horn sharks (Heterodontiformes). [8]

Chimeras (Holocephali)

Chimeras (Holocephali) have pavement teeth that are flat, hexagonal in shape and interconnect to form an even dental plate. There is the presence of calcified strengthened cartilaginous jaws, calcified struts within the jaws and a lever 'nutcracker' system that amplifies the force of the jaw adductor muscles. The fusion of the palatoquadrate and mandibular symphysis, a restricted gape and asynchronous activation of the jaw adductors are key elements in the 'nutcracker' model of jaw-crushing ability. Chimeras use their pavement teeth for grinding molluscs, gastropods and crabs. [8]

Myliobatidae

Myliobatidae are free-swimming rays whose pectoral fins make up broad, powerful "wings" which include the eagle and cow-nose rays. They feed on molluscs and have dentitions adapted to crushing. Dentitions of durophagous myliobatids show several specializations in the jaws and teeth related to their diet. The cartilaginous jaws are strengthened by calcified struts (trabeculae), and the palatoquadrate and mandibular symphysis are fused. Strong ligaments connecting the upper and lower jaws restrict the jaw gape. The strong adductor muscles can be asynchronously activated. [12] [8]

Eagle (Aetobatus narinari) and cow-nose (Rhinoptera javanica) rays

In eagle ( Aetobatus narinari ) and cow-nose ( Rhinoptera javanica ) rays, teeth are hexagonal and are arranged in anteroposterior files packed closely together in an alternating array to form an almost gap-free pavement, similar to the organization found in Chimeras. The teeth are covered with a layer of enameloid. The tooth pavement is stabilized by vertical surfaces that bear ridges and grooves which are interconnected with those on neighboring teeth. These rays also use their pavement teeth for grinding molluscs, gastropods and crabs. Cow nose rays are specialized suction feeders, which open and close their jaws to generate water movements that are used to excavate buried prey. Food capture is achieved by suction and the prey is then cleaned by actions similar to those used in excavation. [12]

Myliobatis and Aetobatus

In Myliobatis and Aetobatus , anteroposterior ridges of the basal plate extend from the posterior margin of the tooth and these interdigitate with those of the succeeding tooth and also form a shelf on which the body of the neighboring tooth rests. The dentition of the bat ray ( Myliobatis californica ) is made up of a series of seven files of crushing teeth. The central hexagonal plate is very wide, taking up about half the width of the occlusal surface and it is flanked by three lateral files of smaller teeth on each side, the outermost being pentagonal. The crushing surface formed by the teeth of the upper jaw is more curved than that of the lower jaw. [12]

Birds

Shorebirds commonly consume bivalves and snails which are low in chitin but the calcium carbonate shell makes up a large portion of their weight. Bivalves and snails are largely consumed whole by ducks and wading birds. The molluscivores that swallow snails or bivalves whole have large well-modularized gizzards for crushing the strong shells. The gizzard of red-necked stints and red knots is more than ten times larger than the proventriculus. The size of the gizzard is adaptable in these shore birds, becoming atrophied when soft food items like worms are consumed and increasing in size and muscularity following prolonged consumption of snails, cockles or mussels. The production of chitinase for the hydrolysis of chitin is important for birds that consume mollusks. [13]

Marine mammals

Sea otters (Enhydra lutris)

Sea otters preferentially forage on benthic invertebrates, particularly sea urchins, gastropod, bivalve mollusks, and crustaceans. Once prey is caught, the otters use their powerful jaws and sharp teeth to consume their meal quickly, even protective crustacean shells. They have canines that deliver a lethal bite, and molars that can crush bones and the shells of mollusks.

Sea otter molars are broad, flat, multi cuspid teeth and the carnassial are also modified for crushing. Both the temporalis and masseter muscles are well developed, creating a strong bite force. The teeth are extremely broad and carnassial are highly molarized. Captured prey is manipulated with the forepaws or is held temporarily in loose skin pouches in the armpits. For larger, heavier-shelled prey, otters will sometimes exhibit tool-use behavior, breaking open sea urchins and mussels with a false stone used as an anvil. Sea otters can also bite sea urchins and mussels open using their strong jaws and teeth. Adults can crush most of their food items but youngsters have not yet developed powerful enough jaws. Therefore, young otters require the assistance of a tool or stone. Tools may also be used when the molluscs are too large to be crushed in the jaws. [14] [15]

Mammals

Monkeys

All mangabeys appear to be durophagous and possess relatively thick molar enamel and expanded premolars, dental adaptations for processing hard foods. Their diet consists of Sacoglottis gabonensis seeds. These seeds can remain on the ground for months without rotting. With hard-object feeding, Mangabeys needed selection to favour thick molar enamel and flattened molars for crushing seeds. [16]

Giant panda

The giant panda is mainly a herbivore despite its short, relatively unspecialized digestive tract that is characteristic of carnivores. Giant pandas lack microbial digestion in their rumen or caecum that is typical of most herbivores for breaking down cellulose and lignin in plant cell walls. Therefore, Giant Pandas need to get their nutrients from the cell contents and fraction of hemicellulose they can break down. The panda subsists mainly on bamboo and does so with modifications of their jaws. Pandas show elaboration of the crushing features of the dentition. The molars are broad, flat, multi cuspid teeth and are the main grinding surface. Jaw action is not a simple crushing one but rather a definite sideways grinding. Panda jaws have a large zygomatico-mandibularis muscle, which is responsible for the sideways movement of the jaw. The glenoid is very deep, preventing back and forth movement of the jaw. [15] [17]

Bamboo represents a predictable food source which is seasonally abundant. Pandas are able to subsist on it despite its low nutritive content. Pandas do this by moving large quantities through the digestive tract in a short period of time. They also reduce their energy expenditures by resting and only remaining active to feed, and they don't have a hibernation period, allowing them to have more foraging time. They chose security over uncertainty, indicated by their bamboo eating adaptations. [17]

Hyaenids

Bone-crushing eating habits appear to be associated with stronger teeth, as seen is in hyaenids. This is because bone-crushing requires greater bite strength and increases the risk of canine breakage. In hyaenids, the carnassial are slightly less specialized as cutting blades than those of the Felidae. The bone-crushing adaptations relate mainly to the premolars. The anterior and posterior cusps are reduced and the central cusp enlarged and widened, so that the tooth is converted from a blade-like structure to a heavy conical hammer. Strong muscles are also required for bone crushing, and the temporalis attachment on the skull is enlarged by a strong sagittal crest. Heavy, hammer-like teeth and extremely strong jaws and jaw muscles make it possible for hyaenas to crack larger bone than other carnivores are capable of, and their highly efficient cutting carnassials can deal with tough hides and tendons. [15] [17] [18]

Wolverine (Gulo gulo)

The wolverine has jaws and teeth that are extremely powerful and together with its scavenging habits, have earned the wolverine the name "hyena of the north". The wolverine is an effective scavenger, capable of cracking heavy bones and shows the same adaptations in the jaw as the hyenas do. The sagittal crest projects well above the area of attachment of the neck muscles, and in a large animal it extends back far behind the level of the condyles to provide attachments for the relatively enormous temporalis muscles, creating a powerful bite force. [17]

See also

Related Research Articles

<span class="mw-page-title-main">Jaw</span> Opposable articulated structure at the entrance of the mouth

The jaws are a pair of opposable articulated structures at the entrance of the mouth, typically used for grasping and manipulating food. The term jaws is also broadly applied to the whole of the structures constituting the vault of the mouth and serving to open and close it and is part of the body plan of humans and most animals.

<span class="mw-page-title-main">Moray eel</span> Family of fishes

Moray eels, or Muraenidae, are a family of eels whose members are found worldwide. There are approximately 200 species in 15 genera which are almost exclusively marine, but several species are regularly seen in brackish water, and a few are found in fresh water.

<span class="mw-page-title-main">Machairodontinae</span> Extinct subfamily of carnivores

Machairodontinae is an extinct subfamily of carnivoran mammals of the family Felidae. They were found in Asia, Africa, North America, South America, and Europe from the Miocene to the Pleistocene, living from about 16 million until about 10,000 years ago.

<span class="mw-page-title-main">Carnassial</span> Mammal tooth type

Carnassials are paired upper and lower teeth modified in such a way as to allow enlarged and often self-sharpening edges to pass by each other in a shearing manner. This adaptation is found in carnivorans, where the carnassials are the modified fourth upper premolar and the first lower molar. These teeth are also referred to as sectorial teeth.

<span class="mw-page-title-main">Horn shark</span> Species of shark

The horn shark is a species of bullhead shark, in the family Heterodontidae. It is endemic to the coastal waters off the western coast of North America, from California to the Gulf of California. Young sharks are segregated spatially from the adults, with the former preferring deeper sandy flats and the latter preferring shallower rocky reefs or algal beds. A small species typically measuring 1 m (3.3 ft) in length, the horn shark can be recognized by a short, blunt head with ridges over its eyes, two high dorsal fins with large spines, and a brown or gray coloration with many small dark spots.

<i>Globidens</i> Extinct genus of lizards

Globidens is an extinct genus of mosasaurid oceanic lizard classified as part of the Globidensini tribe in the Mosasaurinae subfamily.

A molluscivore is a carnivorous animal that specialises in feeding on molluscs such as gastropods, bivalves, brachiopods and cephalopods. Known molluscivores include numerous predatory molluscs,, arthropods such as crabs and firefly larvae, and, vertebrates such as fish, birds and mammals. Molluscivory is performed in a variety ways with some animals highly adapted to this method of feeding behaviour. A similar behaviour, durophagy, describes the feeding of animals that consume hard-shelled or exoskeleton bearing organisms, such as corals, shelled molluscs, or crabs.

<span class="mw-page-title-main">Kidako moray</span> Species of fish

The Kidako moray is a species of marine fish in the family Muraenidae. It inhabits coral reefs or lagoons and could be found in tropical and subtropical seas near Taiwan, Japan, and Australia. The species is diurnal, which means it is more active in the daytime than the nighttime. It is also piscivorous: it consumes fish, octopus, and squid. Other than the Kidako moray, there are about 200 species of moray eels in the Muraenidae family. The Kidako moray would not attack humans unless they are provoked. However, due to the menacing looks of the Kidako moray and moray eels in general, they are feared by divers and snorkelers.

<span class="mw-page-title-main">Branchial arch</span> Bony "loops" present in fish, which support the gills

Branchial arches, or gill arches, are a series of paired bony "loops" that support the gills in fish. As gills are the primitive feature of vertebrates, all vertebrate embryos develop pharyngeal arches, though the eventual fate of these arches varies between taxa. In jawed fish, the first arch pair develops into the jaw. The second gill arches develop into the hyomandibular complex, which supports the back of the jaw and the front of the gill series. The remaining posterior arches support the gills. In amphibians and reptiles, many pharyngeal arch elements are lost, including the gill arches, resulting in only the oral jaws and a hyoid apparatus remaining. In mammals and birds, the hyoid is simplified further.

<span class="mw-page-title-main">Pharyngeal jaw</span> Mophological feature in some fish

Pharyngeal jaws are a "second set" of jaws contained within an animal's throat, or pharynx, distinct from the primary or oral jaws. They are believed to have originated as modified gill arches, in much the same way as oral jaws. Originally hypothesized to have evolved only once, current morphological and genetic analyses suggest at least two separate points of origin. Based on connections between musculoskeletal morphology and dentition, diet has been proposed as a main driver of the evolution of the pharyngeal jaw. A study conducted on cichlids showed that the pharyngeal jaws can undergo morphological changes in less than two years in response to their diet. Fish that ate hard-shelled prey had a robust jaw with molar-like teeth fit for crushing their durable prey. Fish that ate softer prey, on the other hand, exhibited a more slender jaw with thin, curved teeth used for tearing apart fleshy prey. These rapid changes are an example of phenotypic plasticity, wherein environmental factors affect genetic expression responsible for pharyngeal jaw development. Studies of the genetic pathways suggest that receptors in the jaw bone respond to the mechanical strain of biting hard-shelled prey, which prompts the formation of a more robust set of pharyngeal jaws.

<i>Zygophyseter</i> Extinct genus of sperm whales

Zygophyseter varolai is an extinct sperm whale that lived during the Tortonian age of the Late Miocene 11.2 to 7.6 million years ago. It is known from a single specimen from the Pietra Leccese Formation in Italy. It was a member of a stem group of fossil macroraptorial sperm whales also including Brygmophyseter, Acrophyseter, and Livyatan. It probably grew to be around 6.5 to 7 meters in length and shared some characteristics with other raptorials, such as large teeth with tooth enamel that were functional in both the upper and lower jaws which the modern sperm whale lacks. It also had a beak, the ability to echolocate prey, and could have probably swum faster than the modern-day sperm whale which can reach 4 kilometers per hour (2.5 mph). These were probably used in the capture of large prey, such as large fish, seals, and whales. In fact, its common name, the killer sperm whale, refers to its feeding habits that would have had a resemblance to the modern-day killer whale.

<span class="mw-page-title-main">Pharyngeal teeth</span> Teeth in the throat a number of other fish species

Pharyngeal teeth are teeth in the pharyngeal arch of the throat of cyprinids, suckers, and a number of other fish species otherwise lacking teeth.

<span class="mw-page-title-main">Tooth</span> Hard structure of the mouth

A tooth is a hard, calcified structure found in the jaws of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tearing food, for defensive purposes, to intimidate other animals often including their own, or to carry prey or their young. The roots of teeth are covered by gums. Teeth are not made of bone, but rather of multiple tissues of varying density and hardness that originate from the outermost embryonic germ layer, the ectoderm.

<i>Simosaurus</i> Extinct genus of reptiles

Simosaurus is an extinct genus of marine reptile within the superorder Sauropterygia from the Middle Triassic of central Europe. Fossils have been found in deposits in France and Germany that are roughly 230 million years old. It is usually classified as a nothosaur, but has also been considered a pachypleurosaur or a more primitive form of sauropterygian.

<span class="mw-page-title-main">Fish jaw</span>

Most bony fishes have two sets of jaws made mainly of bone. The primary oral jaws open and close the mouth, and a second set of pharyngeal jaws are positioned at the back of the throat. The oral jaws are used to capture and manipulate prey by biting and crushing. The pharyngeal jaws, so-called because they are positioned within the pharynx, are used to further process the food and move it from the mouth to the stomach.

<i>Simosthenurus occidentalis</i> Species of marsupial

Simosthenurus occidentalis is a species of sthenurine marsupial that existed in Australia during the Pliocene, becoming extinct in the Pleistocene epoch around 42,000 years ago. It was a large herbivorous biped that resembles large kangaroos, but with a heavier body than modern kangaroos. The structure of the skull and teeth - resembling koalas and panda bears - indicates that it consumed tough vegetation.

<i>Cartorhynchus</i> Extinct genus of reptiles

Cartorhynchus is an extinct genus of early ichthyosauriform marine reptile that lived during the Early Triassic epoch, about 248 million years ago. The genus contains a single species, Cartorhynchus lenticarpus, named in 2014 by Ryosuke Motani and colleagues from a single nearly-complete skeleton found near Chaohu, Anhui Province, China. Along with its close relative Sclerocormus, Cartorhynchus was part of a diversification of marine reptiles that occurred suddenly during the Spathian substage, soon after the devastating Permian-Triassic extinction event, but they were subsequently driven to extinction by volcanism and sea level changes by the Middle Triassic.

<span class="mw-page-title-main">Biting</span> Behaviour of opening and closing the jaw found in many animals

Biting is an action involving a set of teeth closing down on an object. It is a common zoological behavior, being found in toothed animals such as mammals, reptiles, amphibians, fish, and arthropods. Biting is also an action humans participate in, most commonly when chewing food. Myocytic contraction of the muscles of mastication is responsible for generating the force that initiates the preparatory jaw abduction (opening), then rapidly adducts (closes) the jaw and moves the top and bottom teeth towards each other, resulting in the forceful action of a bite. Biting is one of the main functions in the lives of larger organisms, providing them the ability to forage, hunt, eat, build, play, fight, protect, and much more. Biting may be a form of physical aggression due to predatory or territorial intentions. In animals, biting can also be a normal activity, being used for eating, scratching, carrying objects, preparing food for young, removing ectoparasites or irritating foreign objects, and social grooming. Humans can have the tendency to bite each other whether they are children or adults.

Delotrochanter is an extinct genus of large-sized carnivoran mammals, belonging to the Amphicyonidae, that lived in the Great Plains of North America during the Oligocene to Early Miocene. This genus includes some of the largest and last surviving members of the subfamily Temnocyoninae. it furthermore possesses adaptions towards both bone-crushing and cursoriality.

Eoarctos is an extinct genus of arctoid carnivorans, known from the latest Eocene to early Oligocene of North Dakota and Nebraska. It is known from several remains, the most notable of which is the almost perfectly preserved skeleton of a large male. It was comparable in build and size to a fisher or small raccoon, with an estimated body mass of 4.3 kg, and possessed a variety of features that indicate a mix of terrestrial and scansorial locomotion. Its most notable feature is its unique dentition, with its massive premolars and hypocarnivorous molars, as well as its robust mandible, indicating that it consumed hard-shelled prey, possibly making it the oldest molluscivorous carnivoran known.

References

  1. Huber, Daniel R.; Dean, Mason N.; Summers, Adam P. (2008-08-06). "Hard prey, soft jaws and the ontogeny of feeding mechanics in the spotted ratfish Hydrolagus colliei". Journal of the Royal Society Interface. 5 (25): 941–953. doi:10.1098/rsif.2007.1325. ISSN   1742-5689. PMC   2607471 . PMID   18238758.
  2. Pregill, Gregory (1984). "Durophagous Feeding Adaptations in an Amphisbaenid". Journal of Herpetology. 18 (2): 186–191. doi:10.2307/1563747. JSTOR   1563747.
  3. Myers, Timothy S.; Polcyn, Michael J.; Mateus, Octávio; Vineyard, Diana P.; Gonçalves, António O.; Jacobs, Louis L. (2017-11-13). "Phylogeny". Data from: A new durophagous stem cheloniid turtle from the lower Paleocene of Cabinda, Angola (Data Set). Dryad Digital Repository. doi:10.5061/dryad.n618q.
  4. Tseng, Zhijie Jack; Wang, Xiaoming (2010-11-01). "Cranial functional morphology of fossil dogs and adaptation for durophagy in Borophagus and Epicyon (Carnivora, Mammalia)". Journal of Morphology. 271 (11): 1386–1398. doi:10.1002/jmor.10881. ISSN   1097-4687. PMID   20799339. S2CID   7150911.
  5. Huber, Daniel R.; Eason, Thomas G.; Hueter, Robert E.; Motta, Philip J. (2005-09-15). "Analysis of the bite force and mechanical design of the feeding mechanism of the durophagous horn shark Heterodontus francisci". Journal of Experimental Biology. 208 (18): 3553–3571. doi: 10.1242/jeb.01816 . ISSN   0022-0949. PMID   16155227.
  6. Pfaller, Joseph B.; Gignac, Paul M.; Erickson, Gregory M. (2011-05-15). "Ontogenetic changes in jaw-muscle architecture facilitate durophagy in the turtle Sternotherus minor". Journal of Experimental Biology. 214 (10): 1655–1667. doi: 10.1242/jeb.048090 . ISSN   0022-0949. PMID   21525311.
  7. Figueirido, Borja; Tseng, Zhijie Jack; Martín-Serra, Alberto (2013-07-01). "Skull Shape Evolution in Durophagous Carnivorans". Evolution. 67 (7): 1975–1993. doi: 10.1111/evo.12059 . ISSN   1558-5646. PMID   23815654. S2CID   23918004.
  8. 1 2 3 4 5 6 7 Wilga, C. D.; Motta, P. J. (2000-09-15). "Durophagy in sharks: feeding mechanics of the hammerhead Sphyrna tiburo". Journal of Experimental Biology. 203 (18): 2781–2796. ISSN   0022-0949. PMID   10952878.
  9. Pierre, Vandewalle; Eric, Parmentier; Michel, Chardon (2000). "The branchial basket in Teleost feeding". Cybium. 24 (4). ISSN   0399-0974.
  10. 1 2 Fryer, Geoffrey; Iles, T. D. (1972). The cichlid fishes of the great lakes of Africa: their biology and evolution. Edinburgh: Oliver and Boyd. ISBN   978-0050023471. OCLC   415879.
  11. 1 2 Delos Gerking, Shelby (1994). Feeding ecology of fish. San Diego: Academic Press. ISBN   978-0122807800. OCLC   29390160.
  12. 1 2 3 Berkovitz, B. K.; Shellis, R. P. (2017). The teeth of non-mammalian vertebrates. London: Academic Press. ISBN   9780128028506. OCLC   960895126.
  13. Klasing, Kirk C. (1998). Comparative avian nutrition. Wallingford, Oxon, UK: Cab International. ISBN   9780851992198. OCLC   37361786.
  14. Berta, Annalisa; Sumich, James L.; Kovacs, Kit M. (2015). Marine Mammals. pp. 397–463. doi:10.1016/b978-0-12-397002-2.00012-0. ISBN   9780123970022.
  15. 1 2 3 F., Ewer, R. (1973). The carnivores. Ithaca, N.Y.: Cornell University Press. ISBN   9780801407451. OCLC   621882.{{cite book}}: CS1 maint: multiple names: authors list (link)
  16. McGraw, W. Scott; Pampush, James D.; Daegling, David J. (2012-02-01). "Brief communication: Enamel thickness and durophagy in mangabeys revisited". American Journal of Physical Anthropology. 147 (2): 326–333. doi:10.1002/ajpa.21634. ISSN   1096-8644. PMID   22101774.
  17. 1 2 3 4 Gittleman, John L. (1989–1996). Carnivore behavior, ecology, and evolution. Ithaca: Comstock Pub. Associates. ISBN   9780801430275. OCLC   18135778.
  18. Hartstone-Rose, Adam; Steynder, Deano D. (2013-05-01). "Hypercarnivory, durophagy or generalised carnivory in the Mio-Pliocene hyaenids of South Africa? : research article". South African Journal of Science. 109 (5–6). doi: 10.1590/sajs.2013/20120040 . hdl: 10520/EJC136400 . ISSN   0038-2353.