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Egg fossils are 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 preserve ...
ized remains of
egg An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the ...
s laid by
ancient Ancient history is a time period from the beginning of writing and recorded human history through late antiquity. The span of recorded history is roughly 5,000 years, beginning with the development of Sumerian cuneiform script. Ancient h ...
animal Animals are multicellular, eukaryotic organisms in the Biology, biological Kingdom (biology), kingdom Animalia (). With few exceptions, animals heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, ...
s. As evidence of the
physiological Physiology (; ) is the science, scientific study of function (biology), functions and mechanism (biology), mechanisms in a life, living system. As a branches of science, subdiscipline of biology, physiology focuses on how organisms, organ syst ...
processes of an animal, egg fossils are considered a type of
trace fossil A trace fossil, also called an ichnofossil (; ), is a fossil record of biological activity by lifeforms, but not the preserved remains of the organism itself. Trace fossils contrast with body fossils, which are the fossilized remains of part ...
. Under rare circumstances a fossil egg may preserve the remains of the once- developing
embryo An embryo ( ) is the initial stage of development for a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male sp ...
inside, in which case it also contains body fossils. A wide variety of different animal groups laid eggs that are now preserved in the fossil record beginning in the
Paleozoic The Paleozoic ( , , ; or Palaeozoic) Era is the first of three Era (geology), geological eras of the Phanerozoic Eon. Beginning 538.8 million years ago (Ma), it succeeds the Neoproterozoic (the last era of the Proterozoic Eon) and ends 251.9 Ma a ...
. Examples include
invertebrate Invertebrates are animals that neither develop nor retain a vertebral column (commonly known as a ''spine'' or ''backbone''), which evolved from the notochord. It is a paraphyletic grouping including all animals excluding the chordata, chordate s ...
s like
ammonoids Ammonoids are extinct, (typically) coiled-shelled cephalopods comprising the subclass Ammonoidea. They are more closely related to living octopuses, squid, and cuttlefish (which comprise the clade Coleoidea) than they are to nautiluses (family N ...
as well as
vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...
s like
fishes A fish (: fish or fishes) is an aquatic, anamniotic, gill-bearing vertebrate animal with swimming fins and a hard skull, but lacking limbs with digits. Fish can be grouped into the more basal jawless fish and the more common jawed ...
, possible
amphibian Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
s, and
reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...
s. The latter group includes the many
dinosaur egg Dinosaur eggs are the organic vessels in which a dinosaurs, dinosaur embryo develops. When the first scientifically documented remains of non-avian dinosaurs were being described in England during the 1820s, it was presumed that dinosaurs had la ...
s that have been recovered from
Mesozoic The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
strata In geology and related fields, a stratum (: strata) is a layer of Rock (geology), rock or sediment characterized by certain Lithology, lithologic properties or attributes that distinguish it from adjacent layers from which it is separated by v ...
. Since the organism responsible for laying any given egg fossil is frequently unknown, scientists classify eggs using a parallel system of taxonomy separate from but modeled after the Linnaean system. This " parataxonomy" is called ''veterovata''.


History

The first named oospecies was '' Oolithes bathonicae'', a name given provisionally by Professor J. Buckman to a group of eggs which Buckman believed were laid by a teleosaur. However, modern scientists no longer think it is possible to determine what kind of reptile laid these eggs. In 1859, the first scientifically documented dinosaur egg fossils were discovered in southern France by a Catholic
priest A priest is a religious leader authorized to perform the sacred rituals of a religion, especially as a mediatory agent between humans and one or more deity, deities. They also have the authority or power to administer religious rites; in parti ...
and amateur naturalist named Father Jean-Jacques Poech, however he thought they were laid by giant birds. The first scientifically ''recognized'' dinosaur egg fossils were discovered serendipitously in 1923 by an
American Museum of Natural History The American Museum of Natural History (AMNH) is a natural history museum on the Upper West Side of Manhattan in New York City. Located in Theodore Roosevelt Park, across the street from Central Park, the museum complex comprises 21 interconn ...
crew while looking for evidence of early humans in Mongolia. Egg discoveries continued to mount all over the world, leading to the development of multiple competing classification schemes. In 1975 Chinese paleontologist
Zhao Zi-Kui Zhao may refer to: * Zhao (surname) (赵), a Chinese surname ** commonly spelled Chao in Taiwan or up until the early 20th century in other regions ** Chiu, from the Cantonese pronunciation ** Cho (Korean surname), represent the Hanja 趙 (Chine ...
started a revolution in fossil egg classification by developing a system of " parataxonomy" based on the traditional Linnaean system to classify eggs based on their physical qualities rather than their hypothesized mothers. Zhao's new method of egg classification was hindered from adoption by Western scientists due to language barriers. However, in the early 1990s Russian paleontologist Konstantin Mikhailov brought attention to Zhao's work in the English language
scientific literature Scientific literature encompasses a vast body of academic papers that spans various disciplines within the natural and social sciences. It primarily consists of academic papers that present original empirical research and theoretical ...
.


Diversity


Invertebrates

Eggs laid by invertebrate animals are known from the fossil record. Among these are eggs laid by ancient
cephalopods A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...
. Eggs laid by
ammonoids Ammonoids are extinct, (typically) coiled-shelled cephalopods comprising the subclass Ammonoidea. They are more closely related to living octopuses, squid, and cuttlefish (which comprise the clade Coleoidea) than they are to nautiluses (family N ...
are the best known cephalopod egg fossils. The best preserved fossil ammonite eggs were preserved in the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
Kimmeridge Clay The Kimmeridge Clay is a sedimentary rock, sedimentary deposit of fossiliferous marine clay which is of Late Jurassic to lowermost Cretaceous age and occurs in southern and eastern England and in the North Sea. This rock formation (geology), form ...
of
England England is a Countries of the United Kingdom, country that is part of the United Kingdom. It is located on the island of Great Britain, of which it covers about 62%, and List of islands of England, more than 100 smaller adjacent islands. It ...
. Nevertheless, the
fossil record 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 ...
of cephalopod eggs is scant since their soft, gelatinous eggs decompose quickly and have little chance to fossilize. Another major group of Mesozoic cephalopods, the belemnoids, have no documented eggs in the fossil record whatsoever, although this may be because scientists have not properly searched for them rather than an actual absence from the fossil record.


Fishes and amphibians

Fossil fish eggs have an extensive record going at least as far back as the
Devonian The Devonian ( ) is a period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...
and spanning into the
Cenozoic era The Cenozoic Era ( ; ) is Earth's current geological Era (geology), era, representing the last 66million years of Earth's history. It is characterized by the dominance of mammals, Insect, insects, birds and flowering plant, angiosperms (floweri ...
. The eggs of many different fish
taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
have contributed to this record, including
lobe-finned fish Sarcopterygii (; )—sometimes considered synonymous with Crossopterygii ()—is a clade (traditionally a class or subclass) of vertebrate animals which includes a group of bony fish commonly referred to as lobe-finned fish. These vertebrates ar ...
, placoderms, and
sharks Sharks are a group of elasmobranch cartilaginous fish characterized by a ribless endoskeleton, dermal denticles, five to seven gill slits on each side, and pectoral fins that are not fused to the head. Modern sharks are classified within the ...
. Occasionally eggs are preserved still within the mother's body, or associated with fossil
embryos An embryo ( ) is the initial stage of development for a multicellular organism. In organisms that reproduce sexually, embryonic development is the part of the life cycle that begins just after fertilization of the female egg cell by the male spe ...
. Some fossil eggs possibly laid by fish cannot be confidently distinguished from those laid by
amphibians Amphibians are ectothermic, anamniote, anamniotic, tetrapod, four-limbed vertebrate animals that constitute the class (biology), class Amphibia. In its broadest sense, it is a paraphyletic group encompassing all Tetrapod, tetrapods, but excl ...
. Several fossilized fish or amphibian eggs have been classified as ichnogenera, including '' Mazonova'', '' Archaeoovulus'', '' Chimaerotheca'', '' Fayolia'', and '' Vetacapsula''.


Reptiles

The fossil record of reptile eggs goes back at least as far as the
Early Permian 01 or 01 may refer to: * The year 2001, or any year ending with 01 * The month of January * 1 (number) Music * '01 (Richard Müller album), ''01'' (Richard Müller album), 2001 * 01 (Urban Zakapa album), ''01'' (Urban Zakapa album), 2011 * ''01011 ...
. However, since the earliest reptile eggs probably had soft shells with little preservation potential, reptilian eggs may go back significantly farther than their fossil record. Many ancient reptile groups are known from egg fossils including
crocodilia Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
ns,
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
s, and turtles. Some ancient reptiles, like ichthyosaursEllis, Richard, (2003) ''Sea Dragons - Predators of the Prehistoric Oceans''. University Press of Kansas. . and plesiosaurs are known to have given live birth and are therefore not anticipated to have left behind egg fossils. Dinosaur eggs are among the most well known kind of fossil reptile eggs.


Classification

Fossil eggs are classified according to the parataxonomic system called Veterovata. There are three broad categories in the scheme, on the pattern of organismal phylogenetic classification, called oofamilies, oogenera and oospecies (collectively known as ootaxa).Olga Amo, Gloria Cuenca–Bescós & José Ignacio Canudo (1999). José Ignacio Canudo & Gloria Cuenca-Bescós, ed. "Vertebrate eggshell fragments from the Lower Cretaceous (Lower Barremian) of Camino Canales (Galve Bassin, Province of Teruel, NE Spain)" (PDF). IV European Workshop on Vertebrate Palaeontology. Albarracín, Spain: Universidad de Zaragoza. The names of oogenera and oofamilies conventionally contain the root "oolithus" meaning "stone egg", but this rule is not always followed. They are divided up into several basic types: Testudoid, Geckoid, Crocodiloid, Dinosauroid-spherulitic, Dinosauroid-prismatic, and Ornithoid. Veterovata does not always mirror the taxonomy of the animals which laid the eggs.Carpenter, Kenneth (1999). ''Eggs, Nests, and Baby Dinosaurs: A Look at Dinosaur Reproduction (Life of the Past)'', Indiana University Press; .


Parataxonomy

The oogenus level parataxonomy of Veterovata, following Lawver and Jackson (2014) for Testudoid, Hirsch (1996) for Geckonoid eggs, and Mikhailov et al. (1996) for the rest unless otherwise noted: Testudoid * ''Spheruflexibilis'' morphotype ** Oofamily Testudoflexoolithidae *** '' Testudoflexoolithus'' * ''Spherurigidis'' morphotype ** Oofamily Testudoolithidae *** '' Testudoolithus'' *** '' Emydoolithus'' *** '' Haininchelys'' *** '' Chelonoolithus'' Geckoid * Geckonoid morphotype ** Oofamily Gekkoolithidae *** '' Gekkoolithus'' *** '' Gekkonidovum'' Crocodiloid * Oogenus
Mycomorphoolithus ''Mycomorphoolithus'' (meaning "fungus-shaped stone egg") is an oogenus of fossil eggs found in Spain and England. They possibly represent eggshells of non-eusuchian Crocodylomorpha, crocodylomorphs, and are similar to, but not part of, the Krokol ...
* Oofamily
Krokolithidae Krokolithidae is an oofamily of fossil crocodylomorph eggs. The oogenus ''Mycomorphoolithus ''Mycomorphoolithus'' (meaning "fungus-shaped stone egg") is an oogenus of fossil eggs found in Spain and England. They possibly represent eggshells of ...
** ''
Bauruoolithus ''Bauruoolithus'' is an oogenus of fossilized eggs belonging to an extinct crocodyliform from the Late Cretaceous ( late Campanian- early Maastrichtian) of Brazil. ''Bauruoolithus'' eggs were most likely laid by the notosuchian crocodyliform '' ...
'' ** ''
Krokolithes ''Krokolithes'' is an oogenus of Crocodiloid eggs. These eggs were laid by an extinct species of Crocodylian. It contains three oospecies: ''K. dinophilus'', ''K. wilsoni'' and ''K. helleri''. 1 Fossils of the oogenus have been found in the Oligo ...
'' ** '' Suchoolithus'' Mosasauroid * '' Antarcticoolithus'' Dinosauroid-spherulitic * '' Placoolithus'' * '' Sphaerovum'' * '' Stromatoolithus'' * '' Tacuarembovum'' * Oofamily Cairanoolithidae ** '' Cairanoolithus''Sellés, A. G., & Galobart, À. (2015). Reassessing the endemic European Upper Cretaceous dinosaur egg Cairanoolithus. Historical Biology, (ahead-of-print), 1-14. * Oofamily Stalicoolithidae ** '' Coralloidoolithus'' ** '' Shixingoolithus'' ** '' Stalicoolithus'' * Oofamily Spheroolithidae ** '' Guegoolithus'' ** '' Spheroolithus'' ** '' Paraspheroolithus'' * Oofamily Phaceloolithidae ** '' Phaceloolithus'' * Oofamily Ovaloolithidae ** '' Ovaloolithus'' * Oofamily Megaloolithidae ** '' Megaloolithus'' ** '' Pseudomegaloolithus'' * Oofamily Similifaveoloolithidae ** ''
Similifaveoloolithus ''Similifaveoloolithus'' is an oogenus of fossil dinosaur egg from the Tiantai County, Tiantai basin in Zhejiang Province, China. It is the sole known oospecies of the oofamily Similifaveoloolithidae.Wang Qiang, Zhao Zikui, Wang Xiaolin, Zhang Sh ...
'' * Oofamily
Faveoloolithidae Faveoloolithidae is an oofamily of dinosaur eggs. It contains '' Faveoloolithus'', '' Hemifaveoloolithus'', '' Parafaveoloolithus'', and probably '' Sphaerovum''. However, unlike the other Faveoloolthids, ''Sphaerovum'' has compactituberculate ...
** '' Faveoloolithus'' ** '' Hemifaveoloolithus'' ** '' Parafaveoloolithus'' * Oofamily Youngoolithidae ** '' Youngoolithus'' * Oofamily
Dendroolithidae ''Dendroolithus'' is an oogenus of Dendroolithid dinosaur egg found in the late Cenomanian Chichengshan Formation ( Tiantai Group), in the Gong-An-Zhai and Santonian Majiacun Formations of China and the Maastrichtian Nemegt and Campanian B ...
** ''
Dendroolithus ''Dendroolithus'' is an oogenus of Dendroolithid dinosaur egg found in the late Cenomanian Chichengshan Formation ( Tiantai Group), in the Gong-An-Zhai and Santonian Majiacun Formations of China and the Maastrichtian Nemegt and Campanian B ...
'' * Oofamily Dictyoolithidae ** ''
Dictyoolithus ''Dictyoolithus'' is an oogenus of dinosaur egg from the Cretaceous of China. It is notable for having over five superimposed layers of eggshell units. Possibly, it was laid by megalosauroid dinosaurs. Distribution ''Dictyoolithus'' is known fro ...
'' ** '' Paradictyoolithus'' ** '' Protodictyoolithus'' * Oofamily Polyclonoolithidae ** '' Polyclonoolithus''Xie, J.-F., Zhang, S.-K., Jin, X.-S., Li, D.-Q., and Zhou, L.-Q. (2016)
A new type of dinosaur eggs from Early Cretaceous of Gansu Province, China.
" ''Vertebrata PalAsiatica'', 54(1):1-10.
Dinosauroid-prismatic * '' Pseudogeckoolithus'' * Oofamily Arriagadoolithidae ** '' Arriagadoolithus'' ** '' Triprismatoolithus'' * Oofamily Prismatoolithidae ** '' Preprismatoolithus'' ** '' Prismatoolithus'' ** ''
Protoceratopsidovum ''Protoceratopsidovum'' is an oogenus of dinosaur egg from Mongolia. Despite its name (which means "eggs of ''Protoceratops''"Zelenitsky, D., and Currie, P. (2004) "A Cladistic Analysis of Theropod Ootaxa." ''Journal of Vertebrate Paleontology.'' ...
'' ** '' Sankofa'' ** '' Spheruprismatoolithus''E. S. Bray. 1999. Eggs and eggshell from the Upper Cretaceous North Horn Formation, central Utah. In D. D. Gillette (ed.), Vertebrate Paleontology in Utah, Utah Geological Survey Miscellaneous Publication 99-1:361-375 ** '' Trigonoolithus'' Ornithoid * Ornithoid-ratite Morphotype ** '' Ageroolithus'' ** '' Diamantornis'' ** '' Ornitholithus'' ** '' Reticuloolithus''D. K. Zelenitsky and W. J. Sloboda. 2005. Eggshells. In P. J. Currie and E. B. Koppelhus (eds.), Dinosaur Provincial Park: A Spectacular Ancient Ecosystem Revealed. Indiana University Press, Bloomington 398-404 ** '' Struthiolithus'' ** '' Tristraguloolithus'' ** '' Tubercuoolithus'' ** Oofamily
Elongatoolithidae Elongatoolithidae is an oofamily of fossil eggs, representing the eggs of oviraptorosaurs (with the exception of the Bird, avian ''Ornitholithus''). They are known for their highly elongated shape. Elongatoolithids have been found in Europe, Asia ...
*** '' Continuoolithus'' *** '' Ellipsoolithus'' *** '' Elongatoolithus'' *** '' Heishanoolithus'' *** ''
Macroelongatoolithus ''Macroelongatoolithus'' is an oogenus of large theropod dinosaur eggs, representing the eggs of giant caenagnathid oviraptorosaurs. They are known from Asia and from North America. Historically, several oospecies have been assigned to ''Macroelo ...
'' *** ''
Macroolithus ''Macroolithus'' is an oogenus (fossil-egg genus) of dinosaur egg belonging to the oofamily Elongatoolithidae. The Type (biology), type oospecies, ''M. rugustus'', was originally described under the now-defunct oogenus name ''Oolithes''. Three o ...
'' *** '' Nanhsiungoolithus'' *** '' Paraelongatoolithus'' *** '' Porituberoolithus'' *** '' Rodolphoolithus'' *** '' Spongioolithus'' *** '' Trachoolithus'' *** '' Undulatoolithus'' ** Oofamily Laevisoolithidae *** '' Laevisoolithus'' *** '' Subtiliolithus'' *** '' Tipoolithus''Garcia, G., T. Rodolphe, H. Cappetta, B. Marandat, I. Bentaleb, A. Benabdallah and M. Vianey-Liaud. (2003). "First Record of Dinosaur Eggshels and Teeth from The North-West African Maastrichtian (Morocco)." ''Palaeovertebrata'', Montpellier, 32 (2-4): 59-69, ** Oofamily Medioolithidae *** '' Incognitoolithus'' *** '' Microolithus'' *** '' Mediolithus'' ** Oofamily Montanoolithidae *** '' Montanoolithus'' ** Oofamily Oblongoolithidae *** '' Oblongoolithus'' * Ornithoid-prismatic Morphotype ** '' Dispersituberoolithus'' ** Oofamily Gobioolithidae *** ''
Gobioolithus ''Gobioolithus'' is an oogenus of fossil bird egg native to Mongolia. They are small, smooth-shelled, and elongated eggs that were first discovered in the 1960s and early 70s during a series of fossil-hunting expeditions in the Gobi Desert. Two ...
'' ''
Incertae sedis or is a term used for a taxonomy (biology), taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
/Unclassified'' * '' Oolithes'' * '' Metoolithus'' * ''
Mosaicoolithus ''Mosaicoolithus'' is an oogenus of fossil egg from the Cenomanian Chichengshan Formation (Tiantai Group) and Albian to Cenomanian Laijia Formation (Qujiang Group) of Tiantai County, Zhejiang Province in China. Its classification is incertae sed ...
'' * ''
Mycomorphoolithus ''Mycomorphoolithus'' (meaning "fungus-shaped stone egg") is an oogenus of fossil eggs found in Spain and England. They possibly represent eggshells of non-eusuchian Crocodylomorpha, crocodylomorphs, and are similar to, but not part of, the Krokol ...
'' * '' Nipponoolithus'' * '' Parvoblongoolithus'' * '' Parvoolithus''Mikhailov, K.E. (1997). Fossil and recent eggshell in amniotic vertebrates: Fine structure, comparative morphology and classification. Special Papers in Palaeontology 56. The Palaeontological Association. London. (page 58). * '' Plagioolithus'' * '' Styloolithus'' * Oofamily Pachycorioolithidae ** '' Pachycorioolithus''


See also

* Fossil footprint *
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 ...


References


External links


UCMP's online fossil egg exhibit

Paleofile
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