gizzard stone
   HOME

TheInfoList



OR:

A gastrolith, also called a stomach stone or gizzard stone, is a rock held inside a
gastrointestinal tract The gastrointestinal tract (GI tract, digestive tract, alimentary canal) is the tract or passageway of the digestive system that leads from the mouth to the anus. The GI tract contains all the major organs of the digestive system, in humans and ...
. Gastroliths in some species are retained in the muscular
gizzard The gizzard, also referred to as the ventriculus, gastric mill, and gigerium, is an organ found in the digestive tract of some animals, including archosaurs (pterosaurs, crocodiles, alligators, dinosaurs, birds), earthworms, some gastropods, so ...
and used to grind food in animals lacking suitable grinding
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) 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, ...
. In other species the rocks are ingested and pass through the digestive system and are frequently replaced. The
grain size Grain size (or particle size) is the diameter of individual grains of sediment, or the lithified particles in clastic rocks. The term may also be applied to other granular materials. This is different from the crystallite size, which ...
depends upon the size of the animal and the gastrolith's role in digestion. Other species use gastroliths as
ballast Ballast is material that is used to provide stability to a vehicle or structure. Ballast, other than cargo, may be placed in a vehicle, often a ship or the gondola of a balloon or airship, to provide stability. A compartment within a boat, ship ...
.Rondeau, et a
Larval Anurans Adjust Buoyancy in Response to Substrate Ingestion
Copeia: February 2005, Vol. 2005, No. 1, pp. 188-195.
Particles ranging in size from
sand Sand is a granular material composed of finely divided mineral particles. Sand has various compositions but is defined by its grain size. Sand grains are smaller than gravel and coarser than silt. Sand can also refer to a textural class ...
to cobble have been documented.


Etymology

Gastrolith comes from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
γαστήρ (''gastēr''), meaning "stomach", and λίθος (''lithos''), meaning "stone".


Occurrence

Among living
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
s, gastroliths are common among
crocodile Crocodiles (family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to include all extant me ...
s,
alligator An alligator is a large reptile in the Crocodilia order in the genus ''Alligator'' of the family Alligatoridae. The two extant species are the American alligator (''A. mississippiensis'') and the Chinese alligator (''A. sinensis''). Additional ...
s,
herbivorous A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s,
seals Seals may refer to: * Pinniped, a diverse group of semi-aquatic marine mammals, many of which are commonly called seals, particularly: ** Earless seal, or "true seal" ** Fur seal * Seal (emblem), a device to impress an emblem, used as a means of a ...
and
sea lion Sea lions are pinnipeds characterized by external ear flaps, long foreflippers, the ability to walk on all fours, short and thick hair, and a big chest and belly. Together with the fur seals, they make up the family Otariidae, eared seals. ...
s. Domestic
fowl Fowl are birds belonging to one of two biological orders, namely the gamefowl or landfowl (Galliformes) and the waterfowl (Anseriformes). Anatomical and molecular similarities suggest these two groups are close evolutionary relatives; together ...
require access to ''grit''. Stones swallowed by
ostriches Ostriches are large flightless birds of the genus ''Struthio'' in the order Struthioniformes, part of the infra-class Palaeognathae, a diverse group of flightless birds also known as ratites that includes the emus, rheas, and kiwis. There a ...
can exceed a length of . Apparent microgastroliths have also been found in frog tadpoles. Ingestion of silt and gravel by tadpoles of various anuran (frog) species has been observed to improve buoyancy control. Some
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
animals such as
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', ' lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
s appear to have used stones to grind tough plant matter. A rare example of this is the Early Cretaceous theropod ''
Caudipteryx ''Caudipteryx'' (which means "tail feather") is a genus of peacock-sized theropod dinosaurs that lived in the Barremian age of the early Cretaceous (about 124.6 million years ago). They were feathered and extremely birdlike in their overall appea ...
zoui'' from northeastern China, which was discovered with a series of small stones, interpreted as gastroliths, in the area of its skeleton that would have corresponded with its abdominal region. Aquatic animals, such as
plesiosaurs The Plesiosauria (; Greek: πλησίος, ''plesios'', meaning "near to" and ''sauros'', meaning "lizard") or plesiosaurs are an order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared i ...
, may have used them as ballast, to help balance themselves or to decrease their buoyancy, as
crocodile Crocodiles (family Crocodylidae) or true crocodiles are large semiaquatic reptiles that live throughout the tropics in Africa, Asia, the Americas and Australia. The term crocodile is sometimes used even more loosely to include all extant me ...
s do. While some
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
gastroliths are rounded and polished, many stones in living birds are not polished at all. Gastroliths associated with
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
fossils can weigh several kilograms.


Paleontology


History of discovery

In 1906,
George Reber Weiland George may refer to: People * George (given name) * George (surname) * George (singer), American-Canadian singer George Nozuka, known by the mononym George * George Washington, First President of the United States * George W. Bush, 43rd Presiden ...
reported the presence of worn and polished quartz pebbles associated with the remains of
plesiosaurs The Plesiosauria (; Greek: πλησίος, ''plesios'', meaning "near to" and ''sauros'', meaning "lizard") or plesiosaurs are an order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared i ...
and
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', ' lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their ...
dinosaurs Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is the ...
and interpreted these stones as gastroliths. In 1907,
Barnum Brown Barnum Brown (February 12, 1873 – February 5, 1963), commonly referred to as Mr. Bones, was an American paleontologist. Named after the circus showman P. T. Barnum, he discovered the first documented remains of '' Tyrannosaurus'' during a career ...
found gravel in close association with the fossil remains of the duck-billed
hadrosaur 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 ...
''
Claosaurus ''Claosaurus'' ( ; Greek κλάω, ''klao'' meaning 'broken' and , ''sauros'' meaning 'lizard'; "broken lizard", referring to the odd position of the fossils when discovered) is a genus of hadrosauroid dinosaur that lived during the Late Cre ...
'' and interpreted it as gastroliths. Brown was among the first paleontologist to recognize that dinosaurs used gastroliths in their digestive systems to aid in the grinding of food. Other paleontologists over the years were unconvinced. In 1932,
Friedrich von Huene Friedrich von Huene, born Friedrich Richard von Hoinigen, (March 22, 1875 – April 4, 1969) was a German paleontologist who renamed more dinosaurs in the early 20th century than anyone else in Europe. He also made key contributions about v ...
found stones in Late Triassic sediments, in association with the fossil remains of the prosauropod '' Sellosaurus'' and interpreted them as gastroliths. In 1934, the Howe Quarry, a fossil location in northwestern Wyoming also yielded dinosaur bones with their associated gastroliths. In 1942,
William Lee Stokes William Lee Stokes (March 27, 1915, Black Hawk, Carbon County, Utah - December 12, 1994) was a geologist and paleontologist who is best known for his work at Cleveland-Lloyd Dinosaur Quarry in Emery County, Utah. William Stokes lived to be 79 a ...
recognized the presence of gastroliths in the remains of sauropod dinosaurs recovered from Late Jurassic
strata In geology and related fields, a stratum ( : strata) is a layer of rock or sediment characterized by certain lithologic properties or attributes that distinguish it from adjacent layers from which it is separated by visible surfaces known as e ...
.


Identification

Geologist A geologist is a scientist who studies the solid, liquid, and gaseous matter that constitutes Earth and other terrestrial planets, as well as the processes that shape them. Geologists usually study geology, earth science, or geophysics, althou ...
s usually require several pieces of evidence before they will accept that a rock was used by a dinosaur to aid its
digestion Digestion is the breakdown of large insoluble food molecules into small water-soluble food molecules so that they can be absorbed into the watery blood plasma. In certain organisms, these smaller substances are absorbed through the small intest ...
. First, it should be rounded on all edges (and some are polished) because inside a dinosaur's gizzard any genuine gastrolith would have been acted upon by other stones and fibrous materials in a process similar to the action of a rock tumbler. Second, the stone must be unlike the rock found in its geological vicinity, i.e., its geologic context. Many gastroliths have been found in fine grained lake, mud, and swamp deposits. These environs are calm water deposits and could not carry pebbles and cobbles (unlike a river or beach). Oliver Wings also argues that the stone must be found with the
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s of the dinosaur which ingested it. It is this last criterion that causes trouble in identification, as smooth stones found without context can (possibly erroneously in some cases) be dismissed as having been polished by water or wind. Christopher H. Whittle (1988,9) pioneered scanning electron microscope analysis of wear patterns on gastroliths. Wings (2003) found that
ostrich Ostriches are large flightless birds of the genus ''Struthio'' in the order Struthioniformes, part of the infra-class Palaeognathae, a diverse group of flightless birds also known as ratites that includes the emus, rheas, and kiwis. There ...
gastroliths would be deposited outside the skeleton if the carcass was deposited in an aquatic environment for as little as a few days following death. He concludes that this is likely to hold true for all birds (with the possible exception of moa) due to their air-filled bones which would cause a carcass deposited in water to float for the time it needs to rot sufficiently to allow gastroliths to escape. Gastroliths can be distinguished from stream- or beach-rounded rocks by several criteria: gastroliths are highly polished on the higher surfaces, with little or no polish in depressions or crevices, often strongly resembling the surface of worn animal teeth. Stream- or beach-worn rocks, particularly in a high-impact environment, show less polishing on higher surfaces, often with many small pits or cracks on these higher surfaces. Finally, highly polished gastroliths often show long microscopic rilles, presumably caused by contact with stomach acid. Since most gastroliths were scattered when the animal died and many entered a stream or beach environment, some gastroliths show a mixture of these wear features. Others were undoubtedly swallowed by other dinosaurs and highly polished gastroliths may have been swallowed repeatedly. None of the gastroliths examined in a 2001 study of ''
Cedarosaurus ''Cedarosaurus'' (meaning "Cedar lizard" - named after the Cedar Mountain Formation, in which it was discovered) was a nasal-crested macronarian dinosaur genus from the Early Cretaceous Period (Valanginian). It was a sauropod which lived in what ...
'' gastroliths had the "soapy" texture popularly used to distinguish gastroliths from other types of clast. The researchers dismissed using a soapy texture to identify gastroliths as "unreliable". Gastroliths tended to be universally dull, although the colors represented were varied including black, dark brown, purplish red and grey-blue. Reflectance values greater than 50% are very diagnostic for identifying gastroliths. Clasts from beaches and streams tended to have reflectance values of less than 35%. Less than ten percent of beach clasts have reflectance values lying between 50 and 80%. The American Museum of Natural History Photograph # 311488 demonstrates an articulated skeleton of a '' Psittacosaurus mongoliensis'', from the Ondai Sair Formation, Lower
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 ...
Period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
of
Mongolia Mongolia; Mongolian script: , , ; lit. "Mongol Nation" or "State of Mongolia" () is a landlocked country in East Asia, bordered by Russia to the north and China to the south. It covers an area of , with a population of just 3.3 million ...
, showing a collection of about 40 gastroliths inside the
rib cage The rib cage, as an enclosure that comprises the ribs, vertebral column and sternum in the thorax of most vertebrates, protects vital organs such as the heart, lungs and great vessels. The sternum, together known as the thoracic cage, is a sem ...
, about midway between shoulder and pelvis.


Geologic distribution


Jurassic

Gastroliths have sometimes been called Morrison stones because they are often found in the
Morrison Formation The Morrison Formation is a distinctive sequence of Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandstone, siltston ...
(named after the town of Morrison, west of
Denver Denver () is a consolidated city and county, the capital, and most populous city of the U.S. state of Colorado. Its population was 715,522 at the 2020 census, a 19.22% increase since 2010. It is the 19th-most populous city in the Unit ...
,
Colorado Colorado (, other variants) is a state in the Mountain West subregion of the Western United States. It encompasses most of the Southern Rocky Mountains, as well as the northeastern portion of the Colorado Plateau and the western edge of the ...
), a late
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
formation roughly 150 million years old. Some gastroliths are made of
petrified wood Petrified wood, also known as petrified tree (from Ancient Greek meaning 'rock' or 'stone'; literally 'wood turned into stone'), is the name given to a special type of '' fossilized wood'', the fossilized remains of terrestrial vegetation. ' ...
. Most known instances of preserved
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', ' lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their ...
gastroliths are from
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
animals.


Cretaceous

The
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145& ...
Cedar Mountain Formation The Cedar Mountain Formation is the name given to a distinctive sedimentary geologic formation in eastern Utah, spanning most of the early and mid- Cretaceous. The formation was named for Cedar Mountain in northern Emery County, Utah, where Will ...
of Central
Utah Utah ( , ) is a state in the Mountain West subregion of the Western United States. Utah is a landlocked U.S. state bordered to its east by Colorado, to its northeast by Wyoming, to its north by Idaho, to its south by Arizona, and to its ...
is full of highly polished red and black
chert Chert () is a hard, fine-grained sedimentary rock composed of microcrystalline or cryptocrystalline quartz, the mineral form of silicon dioxide (SiO2). Chert is characteristically of biological origin, but may also occur inorganically as a ...
s, and other rounded quartzose clasts, which may partly represent gastroliths. The cherts may themselves contain fossils of ancient animals, such as corals. These stones do not appear to be associated with stream deposits and are rarely more than fist-sized, which is consistent with the idea that they are gastroliths.


Sauropods

Most known instances of preserved
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', ' lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their ...
gastroliths are from
Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...
animals. The largest known gastroliths found in association with sauropod skeletons are approximately ten centimeters in length.Martin, A.J. (2006). Introduction to the Study of Dinosaurs. Second Edition. Oxford, Blackwell Publishing. 560 pp. .


''Cedarosaurus weiskopfae''

In 2001 Frank Sanders, Kim Manley, and
Kenneth Carpenter Kenneth Carpenter (born September 21, 1949, in Tokyo, Japan) is a paleontologist. He is the former director of the USU Eastern Prehistoric Museum and author or co-author of books on dinosaurs and Mesozoic life. His main research interests ...
published a study on 115 gastroliths discovered in association with a ''
Cedarosaurus ''Cedarosaurus'' (meaning "Cedar lizard" - named after the Cedar Mountain Formation, in which it was discovered) was a nasal-crested macronarian dinosaur genus from the Early Cretaceous Period (Valanginian). It was a sauropod which lived in what ...
'' specimen. The stones were identified as gastroliths on the basis of their tight spatial distribution, partial matrix support, and an edge-on orientation indicative of their being deposited while the carcass still had soft tissue. Their high surface
reflectance The reflectance of the surface of a material is its effectiveness in reflecting radiant energy. It is the fraction of incident electromagnetic power that is reflected at the boundary. Reflectance is a component of the response of the electronic ...
values are consistent with other known dinosaur gastroliths. Nearly all of the ''Cedarosaurus'' gastroliths were found within a .06 m volume of space in the gut region of the skeleton. The total mass of the gastroliths themselves was . Most were less than in volume. The least massive clast was and the most was , with most of them being toward the smaller end of that range. The clasts tended to be close to spherical in shape, although the largest specimens were also the most irregular. The largest gastroliths contributed the most to the total surface area of the set. Some gastroliths were so large and irregularly shaped that they may have been difficult to swallow. The gastroliths were mostly composed of
chert Chert () is a hard, fine-grained sedimentary rock composed of microcrystalline or cryptocrystalline quartz, the mineral form of silicon dioxide (SiO2). Chert is characteristically of biological origin, but may also occur inorganically as a ...
, with some
sandstone Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicat ...
,
siltstone Siltstone, also known as aleurolite, is a clastic sedimentary rock that is composed mostly of silt. It is a form of mudrock with a low clay mineral content, which can be distinguished from shale by its lack of fissility.Blatt ''et al.'' 1980, ...
, and
quartzite Quartzite is a hard, non- foliated metamorphic rock which was originally pure quartz sandstone.Essentials of Geology, 3rd Edition, Stephen Marshak, p 182 Sandstone is converted into quartzite through heating and pressure usually related to tec ...
clasts also included. Since some of the most irregular gastroliths are also the largest, it is unlikely that they were ingested by accident. ''Cedarosaurus'' may have found irregular clasts to be attractive potential gastroliths or was not selective about shape. The clasts were generally of dull coloration, suggesting that color was not a major factor for the sauropod's decision making. The high surface area to volume ratio of the largest clasts suggests that the gastroliths may have broken down ingested plant material by grinding or crushing it. The sandstone clasts tended to be fragile and some broke in the process of collection. The sandstone gastroliths may have been rendered fragile after deposition by loss of cement caused by the external chemical environment. If the clasts had been that fragile while the animal was alive, they probably rolled and tumbled in the digestive tract. If they were more robust, they could have served as part of a ball-mill system.


Migration

Paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
s and geologists are researching new methods of identifying gastroliths that have been found disassociated from animal remains, because of the important information they can provide, if indeed they are trace fossils. If the validity of such gastroliths can be verified, it may be possible to trace gastrolithic rocks back to their original source area where the dinosaur first swallowed the rock. This may provide important information on how dinosaurs migrated. Because the number of suspected gastroliths is substantial, they might provide significant new information and insights into the lives and behaviour of dinosaurs.


See also

*
Bezoar A bezoar is a mass often found trapped in the gastrointestinal system, though it can occur in other locations. A pseudobezoar is an indigestible object introduced intentionally into the digestive system. There are several varieties of bezoar, s ...
*
Calculus (medicine) A calculus (plural calculi), often called a stone, is a concretion of material, usually mineral salts, that forms in an organ or duct of the body. Formation of calculi is known as lithiasis (). Stones can cause a number of medical conditions. S ...
*
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 ...
*
Enterolith An enterolith is a mineral concretion or calculus formed anywhere in the gastrointestinal system. Enteroliths are uncommon and usually incidental findings but, once found, they require at a minimum watchful waiting. If there is evidence of compl ...
*
Regurgitalith Regurgitaliths
are the


Footnotes


References

* Darby, D.G. and Ojakangas, J. (1980). Gastroliths from an Upper Cretaceous Plesiosaur. Journal of Paleontology 54:3 * Whittle, C. (1989). On the Origins of Gastroliths: Determining the Weathering Environment of Rounded and Polished Stones by Scanning Electron Microscope Analysis.
Geological Society of America The Geological Society of America (GSA) is a nonprofit organization dedicated to the advancement of the geosciences. History The society was founded in Ithaca, New York, in 1888 by Alexander Winchell, John J. Stevenson, Charles H. Hitch ...
Bulletin 51:5. * Whittle, C. (1988). On the Origins of Gastroliths. Journal of
Vertebrate Paleontology Vertebrate paleontology is the subfield of paleontology that seeks to discover, through the study of fossilized remains, the behavior, reproduction and appearance of extinct animals with vertebrae or a notochord. It also tries to connect, by us ...
, Supplement to 3:28. * Wings, Oliver (2003): Observations on the Release of Gastroliths from Ostrich Chick Carcasses in Terrestrial and Aquatic Environments. ''Journal of Taphonomy'' 1(2): 97-103
PDF fulltext
* Wings, Oliver (2004): Identification, distribution, and function of gastroliths in dinosaurs and extant birds with emphasis on ostriches (''Struthio camelus''). Ph.D. Thesis, The University of Bonn, Bonn, Germany, 187 pp. URN: urn:nbn:de:hbz:5N-0462
PDF fulltext
* Wings, Oliver (2007): A review of gastrolith function with implications for fossil vertebrates and a revised classification. ''Acta Palaeontologica Polonica'' 52(1): 1-16
PDF fulltext
* * Stokes, W. L. 1987. Dinosaur gastroliths revisited. Journal of Paleontology 61: 1242–1246. * {{Authority control Trace fossils Stomach Digestive system