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During most of the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...
(100.5 to 66 million years ago) the eastern half of North America formed Appalachia (named for the
Appalachian Mountains The Appalachian Mountains, often called the Appalachians, (french: Appalaches), are a system of mountains in eastern to northeastern North America. The Appalachians first formed roughly 480 million years ago during the Ordovician Period. They ...
), an island land mass separated from
Laramidia Laramidia was an island continent that existed during the Late Cretaceous period (99.6–66 Ma), when the Western Interior Seaway split the continent of North America in two. In the Mesozoic era, Laramidia was an island land mass separated from A ...
to the west by the
Western Interior Seaway The Western Interior Seaway (also called the Cretaceous Seaway, the Niobraran Sea, the North American Inland Sea, and the Western Interior Sea) was a large inland sea that split the continent of North America into two landmasses. The ancient sea ...
. This seaway had split North America into two massive landmasses due to a multitude of factors such as tectonism and sea-level fluctuations for nearly 40 million years. The seaway eventually expanded, divided across
the Dakotas The Dakotas is a collective term for the U.S. states of North Dakota and South Dakota. It has been used historically to describe the Dakota Territory, and is still used for the collective heritage, culture, geography, fauna, sociology, econo ...
, and by the end of the Cretaceous, it retreated towards the
Gulf of Mexico The Gulf of Mexico ( es, Golfo de México) is an ocean basin and a marginal sea of the Atlantic Ocean, largely surrounded by the North American continent. It is bounded on the northeast, north and northwest by the Gulf Coast of the United ...
and the Hudson Bay. This left the island masses joined in the continent of North America as the
Rocky Mountains The Rocky Mountains, also known as the Rockies, are a major mountain range and the largest mountain system in North America. The Rocky Mountains stretch in straight-line distance from the northernmost part of western Canada, to New Mexico ...
rose. From the Cenomanian to the end of the
Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campani ...
ages of the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...
, Appalachia was separated from the rest of North America. As the
Western Interior Seaway The Western Interior Seaway (also called the Cretaceous Seaway, the Niobraran Sea, the North American Inland Sea, and the Western Interior Sea) was a large inland sea that split the continent of North America into two landmasses. The ancient sea ...
retreated in the
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval ...
, Laramidia and Appalachia eventually connected. Because of this, its
fauna Fauna is all of the animal life present in a particular region or time. The corresponding term for plants is ''flora'', and for fungi, it is ''funga''. Flora, fauna, funga and other forms of life are collectively referred to as ''Biota (ecology ...
was isolated, and developed very differently from the tyrannosaur,
ceratopsian 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 ...
, hadrosaurid, pachycephalosaur and
ankylosaurid Ankylosauridae () is a family of armored dinosaurs within Ankylosauria, and is the sister group to Nodosauridae. The oldest known Ankylosaurids date to around 122 million years ago and went extinct 66 million years ago during the Cretaceous–Pal ...
dominated fauna of the western part of North America, known as "
Laramidia Laramidia was an island continent that existed during the Late Cretaceous period (99.6–66 Ma), when the Western Interior Seaway split the continent of North America in two. In the Mesozoic era, Laramidia was an island land mass separated from A ...
". Due to high sea levels, subsequent erosion, and the lack of orogenic input of sediment into the Western Interior Seaway unlike the east coast of Laramidia, no terrestrially formed deposits have survived, with most dinosaur remains originating from seaborne carcasses that were transported into marine environments. This same lack of terrestrial sediments is also present on the western coast of Laramidia. Some sediments in the northern part of the range have been removed by
glacial erosion Erosion is the action of surface processes (such as Surface runoff, water flow or wind) that removes soil, Rock (geology), rock, or dissolved material from one location on the Earth's crust#Crust, Earth's crust, and then sediment transport, tra ...
of the
Laurentide Ice Sheet The Laurentide Ice Sheet was a massive sheet of ice that covered millions of square miles, including most of Canada and a large portion of the Northern United States, multiple times during the Quaternary glacial epochs, from 2.58 million year ...
during
Quaternary glaciation The Quaternary glaciation, also known as the Pleistocene glaciation, is an alternating series of glacial and interglacial periods during the Quaternary period that began 2.58 Ma (million years ago) and is ongoing. Although geologists describ ...
s, but it is difficult to ascertain how much sediment has been removed, or whether these sediments would have been any more productive than those that remain. Thus relatively little is known about Appalachia in comparison to Laramidia, with the exception of plant life, marine life and the insects trapped in amber from New Jersey. In addition, due to a lack of interest in Appalachia, many fossils that have been found in Appalachia lie unstudied and remain in the inaccurate genera to which they were assigned in the days of E. D. Cope and O. C. Marsh. Only a few fossils of the terrestrial creatures that were found in this region have given us a brief glimpse into what life was like here during the Cretaceous period. However, the area has seen a bit of a resurgence of interest due to several discoveries made in the past few years. As mentioned earlier, not much is known about Appalachia, but some fossil sites, such as the Woodbine Formation, Navesink Formation, Ellisdale Fossil Site, Mooreville Chalk Formation, Demopolis Chalk Formation, Black Creek Group and the
Niobrara Formation The Niobrara Formation , also called the Niobrara Chalk, is a geologic formation in North America that was deposited between 87 and 82 million years ago during the Coniacian, Santonian, and Campanian stages of the Late Cretaceous. It is com ...
, together with ongoing research in the area, have given us a better look into this forgotten world of
paleontology 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 ...
.


Geography

Appalachia stretched from
Quebec Quebec ( ; )According to the Canadian government, ''Québec'' (with the acute accent) is the official name in Canadian French and ''Quebec'' (without the accent) is the province's official name in Canadian English is one of the thirtee ...
and
Newfoundland and Labrador Newfoundland and Labrador (; french: Terre-Neuve-et-Labrador; frequently abbreviated as NL) is the easternmost provinces and territories of Canada, province of Canada, in the country's Atlantic Canada, Atlantic region. The province comprises t ...
all the way down to the
Eastern United States The Eastern United States, commonly referred to as the American East, Eastern America, or simply the East, is the region of the United States to the east of the Mississippi River. In some cases the term may refer to a smaller area or the East C ...
and west to the
Midwestern United States The Midwestern United States, also referred to as the Midwest or the American Midwest, is one of four census regions of the United States Census Bureau (also known as "Region 2"). It occupies the northern central part of the United States. I ...
. Fossils found in these regions indicate that the area was covered in
coastal plain A coastal plain is flat, low-lying land adjacent to a sea coast. A fall line commonly marks the border between a coastal plain and a piedmont area. Some of the largest coastal plains are in Alaska and the southeastern United States. The Gulf Co ...
s and coastal lowlands during the Cretaceous period. Some scientists have proposed the idea that an archipelago of islands had formed during the time that the Western Interior Seaway had divided Laramidia and Appalachia apart until the near end of the Cretaceous. This would allow dinosaurs to migrate into the Gulf Coast and why it would possibly explain why there are some noticeable distinctions with the fauna in the two assemblages of Appalachia. For instance, the southeastern assemblage (which consists of the Carolinas and the Gulf Coast sites) has some tyrannosauroids such as ''Appalachiosaurus'', some hadrosauroids such as ''Eotrachodon'' and ''Lophorhothon'', nodosaurs, dromaeosaurs, and new leptoceratopsian while the northern assemblage (which consists of New Jersey, Delaware, and Maryland) has some tyrannosauroids such as ''Dryptosaurus'', hadrosauroids such as ''Hadrosaurus'', smaller theropods, and a possible
lambeosaur Lambeosaurinae is a group of crested hadrosaurid dinosaurs. Classification Lambeosaurines have been traditionally split into the tribes or clades Parasaurolophini (''Parasaurolophus'', ''Charonosaurus'', others (?).) and Lambeosaurini (''Cor ...
in the area.


Range

Vertebrate fossils have been found along the Atlantic Seaboard as well as other states like Alabama, Georgia, North Carolina, South Carolina, Mississippi, Missouri, Kentucky, Tennessee, Kansas, Nebraska, Iowa, and Minnesota. Parts of Canada that were a part of Appalachia during the Cretaceous include, Manitoba, Ontario, Quebec, Nunavut, New Brunswick, Newfoundland and Labrador, and Nova Scotia.


Fauna


Dinosaurs

From the Cenomanian to the Maastrichtian, Appalachia was inhabited by various groups of dinosaurs including, hadrosauroids,
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 incl ...
, nodosaurs, leptoceratopsians, indeterminate ornithopods, tyrannosauroids,
dromaeosaurs 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 ...
, ornithomimids, and indeterminate maniraptors lived in the area. There is also fossil evidence of possible chasmosaurs, lambeosaurs,
sauropods 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 b ...
, carcharodontosaurs,
caenagnathids Caenagnathidae is a family of bird-like maniraptoran theropod dinosaurs from the Cretaceous of North America and Asia. They are a member of the Oviraptorosauria, and close relatives of the Oviraptoridae. Like other oviraptorosaurs, caenagnathi ...
,
troodontids Troodontidae is a clade of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil discov ...
, and
coelurosaurs Coelurosauria (; from Greek, meaning "hollow tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, tyran ...
that might have inhabited the area. In 2022, fossils unearthed from the Woodbine Formation in Texas confirmed that carcharodontosaurs, troodontids, and coelurosaurs did indeed inhabit Appalachia.


Tyrannosaurs

In Late Cretaceous North America, the dominant predators were the
Tyrannosauroidea 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 supercontinen ...
, huge predatory
theropods 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 ca ...
built for ripping flesh from their prey. In Appalachia, dryptosaurs were the top predators in this clade. Rather than developing the huge heads and massive bodies of their western relatives such as ''
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 Ca ...
'', ''
Albertosaurus ''Albertosaurus'' (; meaning "Alberta lizard") is a genus of tyrannosaurid theropod dinosaurs that lived in western North America during the Late Cretaceous Period, about 71 million years ago. The type species, ''A. sarcophagus'', wa ...
'' and '' Lythronax'', dryptosaurs had more in common with the basal tyrannosaurs like ''
Dilong ''Dilong'' (; lit. "earth dragon") is a Chinese dragon name that is also used to mean "earthworm" in traditional Chinese medicine and '' Geosaurus'' in zoological nomenclature. Dragon In Chinese mythology, ''dilong'' "earth dragon" is one of ...
'' and ''
Eotyrannus ''Eotyrannus'' (meaning "dawn tyrant") is a genus of tyrannosauroid theropod dinosaur hailing from the Early Cretaceous Wessex Formation beds, included in Wealden Group, located in the southwest coast of the Isle of Wight, United Kingdom. The r ...
''. Dryptosaurs were characterized by long arms with three fingers; while they were not as large as the largest tyrannosaurids, fossils from the Potomac Formation in New Jersey show that some of them did evolve some of the large-bodied features that can be found on other tyrannosaurs. Three genera of valid Appalachian tyrannosaurs are known, ''
Dryptosaurus ''Dryptosaurus'' ( ) is a genus of tyrannosauroid that lived approximately 67 million years ago (mya) during the latter part of the Cretaceous period, New Jersey. ''Dryptosaurus'' was a large, bipedal, ground-dwelling carnivore, that grow up to ...
'', '' Appalachiosaurus'', and the recently discovered ''
Teihivenator This list of informally named dinosaurs is a listing of dinosaurs (excluding Aves; birds and their extinct relatives) that have never been given formally published scientific names. This list only includes names that were not properly published ...
'' while other indeterminate fossils lie scattered throughout most of the southern United States like Georgia, North Carolina, and South Carolina. Fossil foot bones from Appalachian deposits indicate another, unnamed tyrannosauroid measuring , indicating that diversity in these basal tyrannosauroids remained high during the Late Cretaceous. These fossilized teeth possibly belong to a species of ''Appalachiosaurus'' or an undescribed species of a new tyrannosaur. There is also the possibility of a fourth tyrannosaur known from Applachia known as ''
Diplotomodon ''Diplotomodon'' (meaning "double cutting tooth") is a dubious genus of theropod dinosaur, from New Jersey. It was possibly a member of the Tyrannosauroidea, the clade that also contains ''Tyrannosaurus''. ''Diplotomodon'' is only known from a s ...
'', but the genus is considered dubious. Fossils from New Jersey and Delaware, most notably in the Mt. Laurel Formation and Merchantville Formations, have revealed that the primitive tyrannosauroids were much more diverse than expected, and some of them grew to lengths of 8 to 9 meters long, making them around the same size as some of the more advanced tyrannosaurs found in Laramidia. Fossils in Cenomanian deposits further indicate tyrannosauroids had been a constant presence in Appalachia since the Middle Cretaceous.


Dromaeosaurs

The dryptosaurs weren't the only predatory dinosaurs in Appalachia. Indeterminate
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 ...
fossils, possibly belonging to
Velociraptorinae Velociraptorinae is a subfamily of the theropod group Dromaeosauridae. The earliest velociraptorines are probably ''Nuthetes'' from the United Kingdom, and possibly '' Deinonychus'' from North America. However, several indeterminate velociraptor ...
and
Saurornitholestinae Saurornitholestinae is a subfamily of the theropod group Dromaeosauridae. The saurornitholestines currently include three monotypic genera: ''Atrociraptor marshalli'', ''Bambiraptor feinbergi'', and ''Saurornitholestes langstoni''. All are medium ...
, and teeth, most closely matching those of '' Saurornitholestes'', have also been unearthed in Appalachia as well; mostly in the southern states like Missouri, North Carolina, South Carolina, Alabama, Mississippi and Georgia. Finds from the Campanian Tar Heel Formation in North Carolina indicate that there may have been dromaeosaurids of considerable size; intermediate between genera such as ''Saurornitholestes'' and '' Dakotaraptor''. Though known only from teeth, the discovery indicates large dromaeosaurids were part of Appalachia's fauna. Along with the dromaeosauridae remains, tyrannosauroidea and possible ornithomimid remains have been unearthed in Missouri as well. It should also be noted that dromaeosaur remains are more commonly found in the southern region of Appalachia when compared to the northern region, with their sizes ranging from smaller forms to gigantic forms. Recent fossils unearthed in New Jersey show that maniraptors were present in Appalachia as well.


Ornithomimids

Various ornithomimid bones, such as '' Coelosaurus'', have also been reported from Appalachia from Missouri, Mississippi, Alabama, Georgia and as far north in states like Maryland, New Jersey, and Delaware, but it is now believed that some of these are the bones of juvenile dryptosaurs while others belong to various undescribed species of ornithomimids. As of 2019, no distinct species of ornithomimosaurs have been identified yet, mostly due to the fact that no complete skeleton has been unearthed yet. However, it can be assumed that most of them were around the same size of their Laramidian relatives, though there is one specimen that could have reached a large size similar to '' Gallimimus'' or '' Beishanlong''. In 2022, fossils from the Eutaw Formation from Mississippi revealed that large ornithomimids did indeed roam Appalachia.


Other Theropods

Fossils from the Woodbine Formation in Texas, one of the few fossil sites which is one of Appalachia's more well preserved fossils, reveal that other theropods might have roamed Appalachia around the time when the Western Interior Seaway first formed, they include possible specimens of allosauroids,
troodontids Troodontidae is a clade of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil discov ...
,
caenagnathids Caenagnathidae is a family of bird-like maniraptoran theropod dinosaurs from the Cretaceous of North America and Asia. They are a member of the Oviraptorosauria, and close relatives of the Oviraptoridae. Like other oviraptorosaurs, caenagnathi ...
, dromaeosaurs, and tyrannosaurs.


Hadrosaurs

Another common group, arguably the most widespread species in the area, of Appalachian dinosaurs were the hadrosaurs which were represented by three groups including
Hadrosauromorpha Hadrosauromorpha is a clade of iguanodontian ornithopods, defined in 2014 by David B. Norman to divide Hadrosauroidea into the basal taxa with compressed manual bones and a pollex, and the derived taxa that lack them. The clade is defined as ...
,
Hadrosauroidea Hadrosauroidea is a clade or superfamily of ornithischian dinosaurs that includes the "duck-billed" dinosaurs, or hadrosaurids, and all dinosaurs more closely related to them than to ''Iguanodon''. Their remains have been recovered in Asia, Europ ...
, and the Hadrosauridae which is now considered to be their "ancestral homeland"; eventually making their way to Laramidia,
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...
, Bolotsky, Y.L. & Kurzanov, S.K. 1991. he hadrosaurs of the Amur Region.In: 'Geology of the Pacific Ocean Border'' Blagoveschensk: Amur KNII. 94-103. n Russian/ref>
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a subcontinent of Eurasia and it is located entirel ...
,
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the sout ...
and
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest cont ...
where they diversified into the lambeosaurine and saurolophine dinosaurs, though some of the primitive hadrosaurs were still present until the end of the Mesozoic. While the fossil record shows a staggering variety of hadrosaur forms in Laramidia, hadrosaur remains for Appalachia show less diversity due to the relative uncommon number of fossil beds. However, a decent number of hadrosaurs are known from Appalachia with '' Protohadros'', '' Claosaurus'', and ''
Lophorhothon ''Lophorhothon'' is a genus of hadrosauroid dinosaur from the Late Cretaceous, the first genus of dinosaur discovered in Alabama, in the United States. Discovery and naming Remains of this small, poorly known perhaps saurolophine dinosaur ...
'' representing
Hadrosauromorpha Hadrosauromorpha is a clade of iguanodontian ornithopods, defined in 2014 by David B. Norman to divide Hadrosauroidea into the basal taxa with compressed manual bones and a pollex, and the derived taxa that lack them. The clade is defined as ...
, ''
Hypsibema ''Hypsibema'' is a little-known genus of dinosaur from the Late Cretaceous (Campanian stage, around 75 million years ago). Its giant fossils were found in the U.S. states of North Carolina and possibly Missouri. It is believed to be a hadro ...
crassicauda'' and '' Hypsibema missouriensis'' representing
Hadrosauroidea Hadrosauroidea is a clade or superfamily of ornithischian dinosaurs that includes the "duck-billed" dinosaurs, or hadrosaurids, and all dinosaurs more closely related to them than to ''Iguanodon''. Their remains have been recovered in Asia, Europ ...
, and ''
Hadrosaurus ''Hadrosaurus'' (; ) is a genus of hadrosaurid ornithopod dinosaurs that lived in North America during the Late Cretaceous Period in what is now the Woodbury Formation about 80 million to 78 million years ago. The holotype specimen was found in f ...
'' and ''
Eotrachodon ''Eotrachodon orientalis'' (meaning "dawn ''Trachodon'' from the east") is a species of hadrosaurid that was described in 2016. The holotype was found in the Mooreville Chalk Formation (Upper Santonian) in Alabama in 2007 and includes a well ...
'' representing Hadrosauridae. These hadrosaurs from Appalachia seem to be closely related to the crestless hadrosaurs of Laramidia like ''
Gryposaurus ''Gryposaurus'' (meaning "hooked-nosed (Greek ''grypos'') lizard"; sometimes incorrectly translated as "griffin (Latin ''gryphus'') lizard") was a genus of duckbilled dinosaur that lived about 80 to 75 million years ago, in the Late Cretaceo ...
'' and '' Edmontosaurus'', despite the fact that they are not considered to be saurolophines. ''Claosaurus'' is known from a specimen which floated into the Interior Seaway and was found in Kansas, might also be from Appalachia, since it was found closer to the Appalachia side of the seaway and is unknown from Western North America. Hadrosaur remains have even been found in
Iowa Iowa () is a state in the Midwestern region of the United States, bordered by the Mississippi River to the east and the Missouri River and Big Sioux River to the west. It is bordered by six states: Wisconsin to the northeast, Illinois to th ...
, though in fragmentary remains,
Tennessee Tennessee ( , ), officially the State of Tennessee, is a landlocked state in the Southeastern region of the United States. Tennessee is the 36th-largest by area and the 15th-most populous of the 50 states. It is bordered by Kentucky to th ...
, most notably from the
Coon Creek Formation The Coon Creek Formation is a geologic formation located in western Tennessee and extreme northeast Mississippi. It is a sedimentary sandy marl deposit, Late Cretaceous (Maastrichtian) in age, about 70 million years old. The formation is known for ...
. ''Hypsibema crassicauda'', over fifty feet long, was one of the largest eastern hadrosaurs, outgrowing some of more derived western hadrosaurs like '' Lambeosaurus'' and '' Saurolophus''. The genus likely took the environmental niche occupied by large
sauropods 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 b ...
in other areas, possibly grown to colossal sizes to that of '' Magnapaulia'' and '' Shantungosaurus''. ''Hypsibema missouriensis'', was another large species of hadrosaur, but it grew up to 45 to 49 feet, which wasn't as large as ''Hypsibema crassicauda''. When it was first discovered in 1945, it was mistaken for a species of sauropod. Hypsibema missouriensis, possibly even all of the other hadrosaurs living on Appalachia, had serrated teeth for chewing the vegetation in the area. Hadrosaur fossils from the Kanguk Formation in Axel Heiberg Island in Nunavut, Canada show that hadrosaurs were rather widespread throughout Appalachia. The fossils found in the Kanguk Formation also revealed the dietary preferences of hadrosaurs, which revealed that they had a diet of conifers, stems, twigs, and various deciduous plants. In 2020, the remains of one small-bodied hadrosaur and two small-bodied hadrosauromorphs were unearthed in the New Egypt Formation in New Jersey. The fossils were dated to the Maastrichtian, which was the last stage of the Cretaceous period that ended with the extinction the dinosaurs. This information would imply that Appalachia probably had a rich diversity of life, but research will be need in order to get a better picture of this lost world. In 2021, new remains of ''Hypsibema missouriensis'', also known as ''Parrosaurus missouriensis'', were unearthed in Missouri.


Lambeosaurs

Indeterminate
lambeosaurinae Lambeosaurinae is a group of crested hadrosaurid dinosaurs. Classification Lambeosaurines have been traditionally split into the tribes or clades Parasaurolophini ('' Parasaurolophus'', '' Charonosaurus'', others (?).) and Lambeosaurini ('' C ...
remains, mostly similar to '' Corythosaurus'', have been reported from New Jersey's Navesink Formation, Bylot Island and Nova Scotia, Canada. It cannot yet be explained how lambeosaurines might have reached Appalachia though some have theorized that a land bridge must have formed sometime during the
Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campani ...
. In 2020, a forelimb belonging to a lambeosaur was unearthed in the New Egypt Formation from New Jersey with evidence of sharks scavenging on its remains.


Nodosaurs

The
nodosaurids Nodosauridae is a family of ankylosaurian dinosaurs, from the Late Jurassic to the Late Cretaceous period in what is now North America, South America, Europe, and Asia. Description Nodosaurids, like their close relatives the ankylosaurids, were ...
, a group of large, herbivorous armored dinosaurs resembling armadillos, are another testament to Appalachia's difference from Laramidia. During the early Cretaceous, the nodosaurids prospered and were one of the most widespread dinosaurs throughout North America. However, by the latest Cretaceous, nodosaurids were scarce in western North America, limited to forms like ''
Edmontonia ''Edmontonia'' is a genus of panoplosaurin nodosaurid dinosaur from the Late Cretaceous Period. It is part of the Nodosauridae, a family within Ankylosauria. It is named after the Edmonton Formation (now the Horseshoe Canyon Formation in Ca ...
'', ''
Denversaurus ''Denversaurus'' (meaning "Denver lizard") is a genus of panoplosaurin nodosaurid dinosaur from the Late Cretaceous (late Maastrichtian) of western North America. Although at one point treated as a junior synonym of ''Edmontonia'' by some t ...
'' and ''
Panoplosaurus ''Panoplosaurus'' is a genus of armoured dinosaur from the Late Cretaceous of Alberta, Canada. Few specimens of the genus are known, all from the middle Campanian of the Dinosaur Park Formation, roughly 76 to 75 million years ago. It was first d ...
''; perhaps due to competition from the
ankylosauridae Ankylosauridae () is a family of armored dinosaurs within Ankylosauria, and is the sister group to Nodosauridae. The oldest known Ankylosaurids date to around 122 million years ago and went extinct 66 million years ago during the Cretaceous–Pa ...
; though they did thrive in isolation, most notably in Appalachia, as mentioned earlier and in the case of '' Struthiosaurus'', Europe as well. Nodosaurid
scute A scute or scutum (Latin: ''scutum''; plural: ''scuta'' "shield") is a bony external plate or scale overlaid with horn, as on the shell of a turtle, the skin of crocodilians, and the feet of birds. The term is also used to describe the anterior po ...
s have been commonly found in eastern North America, while fossil specimens are very rare. Often the findings are not diagnostic enough to identify the species, but the remains attest to a greater number of these armored dinosaurs in Appalachia. Multiple specimens have been unearthed in
Kansas Kansas () is a state in the Midwestern United States. Its capital is Topeka, and its largest city is Wichita. Kansas is a landlocked state bordered by Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to th ...
in the Niobrara Formation, Alabama in
Ripley Formation The Ripley Formation is a geological formation in North America found in the U.S. states of Alabama, Georgia, Mississippi, Missouri, and Tennessee. The lithology is consistent throughout the layer. It consists mainly of glauconitic sandstone. I ...
, Mississippi, Delaware, Maryland and New Jersey, possibly belonging to a multitude of different species. Five possible and best-known examples of Appalachian nodosaurids, from both the early and late Cretaceous period, include ''
Priconodon ''Priconodon'' (meaning "saw cone tooth") is an extinct genus of dinosaur (perhaps nodosaurid), known from its large teeth. Its remains have been found in the Aptian-Albian age Lower Cretaceous Arundel Formation of Muirkirk, Prince George's Count ...
'', '' Propanoplosaurus'', ''
Niobrarasaurus ''Niobrarasaurus'' (meaning "Niobrara lizard") is an extinct genus of nodosaurid ankylosaur which lived during the Cretaceous 87 to 82 million years ago. Its fossils were found in the Smoky Hill Chalk Member of the Niobrara Formation, in western ...
'', '' Silvisaurus'' and possibly ''
Hierosaurus ''Hierosaurus'' (meaning "sacred lizard") is an extinct genus of nodosaurid ankylosaur which lived during the Late Cretaceous 87 to 82 million years ago. Its fossils were found in the Smoky Hill Chalk Member of the Niobrara Formation, in western ...
'', though its validity is disputed. Just like the ''Claosaurus'' specimen, it is possible that the specimens of ''Niobrarasaurus'', ''Silvisaurus'' and ''Hierosaurus'' floated into the Interior Seaway from the east, since these two species of nodosaurids were discovered in the famous chalk formations of Kansas and are not known from any location from Western North America. Kansas was also a part of Appalachia when the other parts were covered by oceans, which were a part of the Western Interior Seaway.


Leptoceratopsians

While remains of the advanced ceratopsians, most notably the centrosaurines and
chasmosaurines Chasmosaurinae is a subfamily of ceratopsid dinosaurs. They were one of the most successful groups of herbivores of their time. Chasmosaurines appeared in the early Campanian, and became extinct, along with all other non- avian dinosaurs, durin ...
which were very common in Laramidia during this time period, where not found in Appalachia, the leptoceratopsids somehow managed to inhabit that location. A
Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campani ...
-era
leptoceratopsid Leptoceratopsidae is an extinct family of neoceratopsian dinosaurs from Asia, North America and Europe. Leptoceratopsids resembled, and were closely related to, other neoceratopsians, such as the families Protoceratopsidae and Ceratopsidae, bu ...
ceratopsian has been found in the Tar Heel Formation, marking the first discovery of a ceratopsian dinosaur in the Appalachian zone. This specimen bears a uniquely long, slender and downcurved upper jaw, suggesting that it was an animal with a specialized feeding strategy, yet another example of speciation on an island environment. Recently, a ceratopsian teeth were unearthed in Mississippi's Owl Creek Formation, which have been dated to be 67 million years old. The owner of this one particular tooth was probably a chasmosaurine since by the end of the Cretaceous, the centrosaurines had completely vanished from North America, though they were thriving in Asia as in the case of ''
Sinoceratops ''Sinoceratops'' is an extinct genus of ceratopsian dinosaur that lived approximately 73 million years ago during the latter part of the Cretaceous Period in what is now Shandong province in China. It was named in 2010 by Xu Xing ''et al.'' f ...
''. While leptoceratopsid remains, the few that have been discovered in recent years, have been unearthed in the southern part of Appalachia, they appear to be completely absent from the northern part of Appalachia, states like New Jersey, Delaware, and Maryland. Suggesting the idea, proposed by paleontologist David R. Schwimmer, that there was a possible providence during the Late Cretaceous. It is not completely understood how the leptoceratopsians were able to reach Appalachia, however, the most commonly accepted theory was that they island hopped during the time that the Western Interior Seaway split the North American continent into two different land masses in a way that some species of leptoceratopsids, most notably '' Ajkaceratops'', were able to reach Europe. It should also be noted that there is a distinct difference with how the leptoceratopsians evolved in Appalachia and Laramida. The Appalachian leptoceratopsian that was unearthed in the Tar Heel Formation, which grew to the size of a large dog, had a more slender jaw that teeth that curved downward and outward in its beak. This would imply a specialized feeding strategy for feeding on the foliage that was native to Appalachia during the
Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campani ...
.


Birds

Several bird remains are known from Appalachian sites, most of them sea birds like
Hesperornithes Hesperornithes is an extinct and highly specialized group of aquatic avialans closely related to the ancestors of modern birds. They inhabited both marine and freshwater habitats in the Northern Hemisphere, and include genera such as '' Hesperorn ...
like ''
Hesperornis ''Hesperornis'' (meaning "western bird") is a genus of cormorant-like bird that spanned the first half of the Campanian age of the Late Cretaceous period (83.5–78 mya). One of the lesser-known discoveries of the paleontologist O. C. Marsh i ...
'', ''
Canadaga ''Canadaga'' (meaning "Canadian bird") is a flightless bird genus from the Late Cretaceous. The single known species is ''Canadaga arctica''. It lived in the shallow seas around what today is Bylot Island in Nunavut, Canada. Its fossils were fou ...
'', ''
Baptornis ''Baptornis'' ("diving bird") is a genus of flightless, aquatic birds from the Late Cretaceous, some 87-80 million years ago (roughly mid-Coniacian to mid-Campanian faunal stages). The fossils of ''Baptornis advenus'', the type species, were dis ...
'', ''
Fumicollis ''Fumicollis'' is a genus of prehistoric flightless birds from the Late Cretaceous (Coniacian-Santonian) Niobrara Chalk of Kansas. Description Diagnostic traits of ''Fumicollis'' include presacral vertebrae with expanded ventral processes, an e ...
'', ''
Parahesperornis ''Parahesperornis'' is a genus of prehistoric flightless birds from the Late Cretaceous. Its range in space and time may have been extensive, but its remains are rather few and far between, at least compared with its contemporary relatives in ''H ...
'', and ''
Ichthyornis ''Ichthyornis'' (meaning "fish bird", after its fish-like vertebrae) is an extinct genus of toothy seabird-like ornithuran from the late Cretaceous period of North America. Its fossil remains are known from the chalks of Alberta, Alabama, ...
'', Enantiornithes like '' Halimornis'' and '' Ornithuraes'' like '' Apatornis'' and '' Iaceornis'', possibly indicating that Appalachia might have had a diverse variety of birds that were endemic to the region. Of particular interest are possible lithornithid remains in
New Jersey New Jersey is a state in the Mid-Atlantic and Northeastern regions of the United States. It is bordered on the north and east by the state of New York; on the east, southeast, and south by the Atlantic Ocean; on the west by the Delaware ...
, arguably one of the best records for Cretaceous birds as some specimens were preserved in the
greensand Greensand or green sand is a sand or sandstone which has a greenish color. This term is specifically applied to shallow marine sediment that contains noticeable quantities of rounded greenish grains. These grains are called ''glauconies'' and co ...
s in the area, which would represent a clear example of palaeognath
Neornithes 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 ...
in the Late Cretaceous. However, this issue is still under debate. Examples of birds that were endemic to Appalachia include a few groups such as the
Charadriiformes Charadriiformes (, from ''Charadrius'', the type genus of family Charadriidae) is a diverse order of small to medium-large birds. It includes about 390 species and has members in all parts of the world. Most charadriiform birds live near water an ...
, which consisted of '' Graculavus'' and '' Telmatornis'', Anseriformes as represented by ''
Anatalavis ''Anatalavis'' is genus of prehistoric birds related to ducks and geese, perhaps in particular the magpie-goose. The species ''Anatalavis rex'' – formerly placed in ''Telmatornis'' – is known from the Hornerstown Formation (Late Cretaceous o ...
'',
Procellariiformes Procellariiformes is an order (biology), order of seabirds that comprises four family (biology), families: the albatrosses, the Procellariidae, petrels and shearwaters, and two families of storm petrels. Formerly called Tubinares and still call ...
, with '' Tytthostonyx'' being one such example, and ''
Palaeotringa ''Palaeotringa'' is a prehistoric bird genus that was discovered by O. C. Marsh during the late 19th century American bone wars. Its remains were found in the controversial Hornerstown Formation of New Jersey (United States) which straddles the ...
'' and '' Laornis'' belonging to a currently unknown group of birds. Hesperornithid fossils have also been unearthed in Arkansas. Some birds found in Canada such as '' Tingmiatornis'' and ''
Canadaga ''Canadaga'' (meaning "Canadian bird") is a flightless bird genus from the Late Cretaceous. The single known species is ''Canadaga arctica''. It lived in the shallow seas around what today is Bylot Island in Nunavut, Canada. Its fossils were fou ...
'' were found in areas that were a part of Appalachia. '' Enantiornithine birds'' are also known from Appalachia, as is the case of '' Flexomornis'' from the Woodbine Formation from Texas.


Non-dinosaur herpetofauna


Amphibians

Through the Ellisdale Fossil Site, a good picture of Appalachia's non-dinosaurian fauna is present. Amidst lissamphibians, there is evidence for sirenids (including the large ''
Habrosaurus ''Habrosaurus'' (, meaning "graceful lizard") is an extinct genus of prehistoric salamanders, and the oldest known member of the family Sirenidae. Two species are known, ''H. prodilatus'' from the middle Campanian Dinosaur Park Formation of Albe ...
''), the batrachosauroidid salamander '' Parrisia'', hylids, possible representatives of '' Eopelobates'' and ''
Discoglossus ''Discoglossus'' (common name: painted frogs) is a genus of frogs in the family Alytidae (formerly Discoglossidae) found in southern Europe and northwestern Africa Africa is the world's second-largest and second-most populous continent, ...
'', showing close similarities to European faunas, but aside from ''
Habrosaurus ''Habrosaurus'' (, meaning "graceful lizard") is an extinct genus of prehistoric salamanders, and the oldest known member of the family Sirenidae. Two species are known, ''H. prodilatus'' from the middle Campanian Dinosaur Park Formation of Albe ...
'' (which is also found on Laramidia) there is a high degree of endemism, suggesting no interchanges with other landmasses throughout the Late Cretaceous.


Lizards

There is also a high degree of endemism in regards to its reptilian fauna: among squamates, the teiid Prototeius is exclusive to the landmass, and native representatives of
Iguanidae The Iguanidae is a family of lizards composed of the iguanas, chuckwallas, and their prehistoric relatives, including the widespread green iguana. Taxonomy Iguanidae is thought to be the sister group to the collared lizards (family Crotaph ...
,
Helodermatidae The Helodermatidae or beaded lizards are a small family of lizards endemic to North America today, but formerly more widespread in the ancient past. Traditionally, the Gila monster and the Mexican beaded lizard were the only species recognized, ...
, and Necrosauridae. No fossilized remains of
snake Snakes are elongated, limbless, carnivorous reptiles of the suborder Serpentes . Like all other squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales. Many species of snakes have skulls with several more j ...
s have been discovered in Appalachia during the Cretaceous period, only being found in Laramidia.


Turtles

Amidst
turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked t ...
s, which are rather common finds in Appalachia, '' Adocus'', '' Apalone'', and '' Bothremys'' are well represented, the latter in particular more common in Appalachian sites than Laramidian ones. '' Pleurochayah'', a bothremyid, is known from Texas. In Santonian Alabama occurred the giant endemic '' Appalachemys''.


Crocodiles

Crocodiles were rather abundant in Appalachia with nine local crocodilian genera belonging to several confirmed families, with the possibility of much more undiscovered crocodiles waiting to be unearthed. ''
Goniopholididae Goniopholididae is an extinct family of moderate-sized semi-aquatic neosuchian crocodyliformes. Their bodyplan and morphology are convergent on living crocodilians. They lived across Laurasia (Asia, Europe and North America) between the Middle J ...
'' is represented by '' Dakotasuchus'' and ''
Woodbinesuchus ''Woodbinesuchus'' (meaning "Woodbine Formation crocodile") is an extinct genus of goniopholidid mesoeucrocodylian. Its fossils have been recovered from the Cenomanian-age Upper Cretaceous Woodbine Formation of Texas. Crocodyliform fossils are wi ...
'',
Alligatoridae The family Alligatoridae of crocodylians includes alligators, caimans and their extinct relatives. Phylogeny The superfamily Alligatoroidea includes all crocodilians (fossil and extant) that are more closely related to the American alligator ...
is represented by '' Bottosaurus'', ''
Neosuchia Neosuchia is a clade within Mesoeucrocodylia that includes all modern extant crocodilians and their closest fossil relatives. It is defined as the most inclusive clade containing all crocodylomorphs more closely related to ''Crocodylus niloticu ...
'' is represented by '' Scolomastax'' and ''
Deltasuchus ''Deltasuchus'' ("delta crocodile") is a genus of neosuchian crocodyliform from the Late Cretaceous of Texas, specifically in the Woodbine Formation, which was a part of the Appalachian continent during the Cretaceous. It is known from one speci ...
'', ''
Alligatoroidea Alligatoroidea is one of three superfamilies of crocodylians, the other two being Crocodyloidea and Gavialoidea. Alligatoroidea evolved in the Late Cretaceous period, and consists of the alligators and caimans, as well as extinct members mor ...
'' are presented ''
Deinosuchus ''Deinosuchus'' () is an extinct genus of alligatoroid crocodilian, related to modern alligators and caimans, that lived 82 to 73 million years ago (Ma), during the late Cretaceous period. The name translates as "terrible crocodile" and i ...
'' and ''
Leidyosuchus ''Leidyosuchus'' (meaning " Leidy's crocodile") is an extinct genus of alligatoroid from the Late Cretaceous of Alberta. It was named in 1907 by Lawrence Lambe, and the type species is ''L. canadensis''. It is known from a number of specimens fro ...
'', ''
Gavialoidea Gavialoidea is one of three superfamilies of crocodylians, the other two being Alligatoroidea and Crocodyloidea. Although many extinct species are known, only the gharial ''Gavialis gangeticus'' and the false gharial ''Tomistoma schlegelii'' ...
'' are represented by '' Thoracosaurus'', '' Eothoracosaurus'', and
Crocodilia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest livi ...
is presented by ''
Borealosuchus ''Borealosuchus'' (meaning "boreal crocodile") is an extinct genus of crocodyliforms that lived from the Late Cretaceous to the Eocene in North America. It was named by Chris Brochu in 1997 for several species that had been assigned to '' Leidyo ...
'', are well established in Laramidia as well, probably indicative of their ocean crossing capacities. ''Deinosuchus'', being one of the largest crocodilians of the fossil record, was an
apex predator An apex predator, also known as a top predator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the highest trophic lev ...
that did prey on the dinosaurs in the area, the same case applies for Laramidia as well, despite the fact that the majority of its diet consisted of turtles and
sea turtle Sea turtles (superfamily Chelonioidea), sometimes called marine turtles, are reptiles of the order Testudines and of the suborder Cryptodira. The seven existing species of sea turtles are the flatback, green, hawksbill, leatherback, loggerhe ...
s. However, crocodiles still preyed on the endemic dinosaurs whenever they got the chance to do so; there is evidence of crocodile bite marks on the femur of large ornithomimosaur that indicates the predatory behavior of native crocodiles. Fossils unearthed in South Carolina and New Jersey shows that some of the crocodilians endemic to Appalachia survived the extinction of the dinosaurs and even persisted into the Cenozoic.


Dyrosauridae

''
Dyrosauridae Dyrosauridae is a family of extinct neosuchian crocodyliforms that lived from the Late Cretaceous (Maastrichtian) to the Eocene. Dyrosaurid fossils are globally distributed, having been found in Africa, Asia, Europe, North America and South Ame ...
'', most notably '' Dyrosaurus'' and ''
Hyposaurus ''Hyposaurus'' is a genus of extinct marine dyrosaurid crocodyliform. Fossils have been found in Paleocene aged rocks of the Iullemmeden Basin in West Africa, Campanian–Maastrichtian (Late Cretaceous) Shendi Formation of Sudan and Maastrichtia ...
'' fossils, are also known form Appalachia, particularly in New Jersey, Alabama, and South Carolina.


Pholidosauridae

Only one species of '' Pholidosauridae'' is known to have lived in Appalachia; '' Terminonaris'' whose remains have been unearthed in Texas and Kansas.


Pterosaurs

Pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 ...
fossils, mostly similar to '' Pteranodon'' and ''
Nyctosaurus ''Nyctosaurus'' (meaning "night lizard" or "bat lizard") is a genus of nyctosaurid pterosaur from the Late Cretaceous period of what is now the Niobrara Formation of the mid-western United States, which, during the time ''Nyctosaurus'' was al ...
'', have been unearthed in Georgia, Alabama and Delaware. On a similar note, azhdrachid remains, which belong to ''
Arambourgiania ''Arambourgiania'' is an extinct genus of azhdarchid pterosaur from the Late Cretaceous period (Maastrichtian stage) of Jordan, and possibly the United States.Harrell, T. Lynn Jr.; Gibson, Michael A.; Langston, Wann Jr. (2016). "A cervical ver ...
'', have been unearthed in Tennessee. There have been a number of specimens of pterosaurs unearthed in areas that were a part of Appalachia during the time that the Western Interior Seaway had divided North America into two landmasses. Fossils of Appalachian pterosaurs have also been unearthed in
Kansas Kansas () is a state in the Midwestern United States. Its capital is Topeka, and its largest city is Wichita. Kansas is a landlocked state bordered by Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to th ...
and
Texas Texas (, ; Spanish: ''Texas'', ''Tejas'') is a state in the South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2020, it is the second-largest U.S. state by ...
. ''
Pteranodontidae The Pteranodontidae are a family of large pterosaurs of the Cretaceous Period of North America and Africa. The family was named in 1876 by Othniel Charles Marsh. Pteranodontids had a distinctive, elongated crest jutting from the rear of the head ( ...
'' seem the most common groups of pterosaurs in Appalachia and represented by five species: '' Pteranodon'', ''
Nyctosaurus ''Nyctosaurus'' (meaning "night lizard" or "bat lizard") is a genus of nyctosaurid pterosaur from the Late Cretaceous period of what is now the Niobrara Formation of the mid-western United States, which, during the time ''Nyctosaurus'' was al ...
'', '' Geosternbergia'', '' Dawndraco'', and '' Alamodactylus''. Members of the ''
Ornithocheiridae Ornithocheiridae (or ornithocheirids, meaning "bird hands") is a group of pterosaurs within the suborder Pterodactyloidea. These pterosaurs were among the last to possess teeth. Members that belong to this group lived from the Early to Late Cr ...
'' and ''
Anhangueridae Anhangueridae is a group of pterosaurs within the suborder Pterodactyloidea. They were among the last pterosaurs to possess teeth. A recent study discussing the group considered the Anhangueridae to be typified by a premaxillary crest and a later ...
'' are represented by ''
Aetodactylus ''Aetodactylus'' (meaning "eagle finger") is a genus of targaryendraconian pterosaur. It is known from a lower jaw discovered in Upper Cretaceous rocks of northeastern Texas, United States. Description ''Aetodactylus'' is based on SMU 763 ...
'' and '' Cimoliopterus'' respectively.


Choristodera

The remains of indetermine choristoderans have turned up in the Navesink Formation; the only known genus of choristodere during the Late Cretaceous was '' Champsosaurus''. As a whole Appalachian choristodere fossils are very rare, speculated to the result of the lack of a suitable cold freshwater environment as seen in Laramidia at similar latitudes; the animals are speculated to have been more common at higher latitudes and altitudes.


Mammals

Several types of mammals are also present in Ellisdale and in both of
the Carolinas The Carolinas are the U.S. states of North Carolina and South Carolina, considered collectively. They are bordered by Virginia to the north, Tennessee to the west, and Georgia to the southwest. The Atlantic Ocean is to the east. Combining Nor ...
. The most common are ptilodontoidean
multituberculates Multituberculata (commonly known as multituberculates, named for the multiple tubercles of their teeth) is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, a ...
, such as '' Mesodma'', '' Cimolodon'' and a massively-sized species. The sheer diversity of species on the landmass, as well as the earlier appearance compared to other Late Cretaceous locales, suggests that ptilodontoideans evolved in Appalachia.
Metatherian Metatheria is a mammalian clade that includes all mammals more closely related to marsupials than to placentals. First proposed by Thomas Henry Huxley in 1880, it is a more inclusive group than the marsupials; it contains all marsupials as well ...
s are also known, including an
alphadon ''Alphadon'' (meaning "first tooth") is an extinct genus of small, primitive mammal that was a member of the metatherians, a group of mammals that includes modern-day marsupials. Its fossils were first discovered and named by George Gaylord Simp ...
tid, a stagodontid, and a herpetotheriid. Unlike ptilodontoideans, metatherians show a lesser degree of endemism, implying a degree of interchange with Laramidia and Europe. Research in this area has revealed that the''
Taeniolabidoidea Taeniolabidoidea is a group of extinct mammals known from North America and Asia. They were the largest members of the extinct order Multituberculata, as well as the largest non-therian mammals. ''Lambdopsalis'' even provides direct fossil eviden ...
'' mammals can trace their origins here and that there were several species of multituberculates endemic to Appalachia. ''
Eutheria Eutheria (; from Greek , 'good, right' and , 'beast'; ) is the clade consisting of all therian mammals that are more closely related to placentals than to marsupials. Eutherians are distinguished from noneutherians by various phenotypic tra ...
'' fossils, most notably molars, have also been unearthed in Mississippi. It is possible that they belong to a creature rather reminiscent to ''
Protungulatum ''Protungulatum'' ('first ungulate') is a extinct genus of pan-euungulate mammals within extinct family Protungulatidae, and also one of the earliest known placental mammals in the fossil record, that lived in North America from the Late Creta ...
''. The genus
Alphadon ''Alphadon'' (meaning "first tooth") is an extinct genus of small, primitive mammal that was a member of the metatherians, a group of mammals that includes modern-day marsupials. Its fossils were first discovered and named by George Gaylord Simp ...
as well as other members of the ''
Allotheria Allotheria (meaning "other beasts", from the Greek , '–other and , '–wild animal) is an extinct branch of successful Mesozoic mammals. The most important characteristic was the presence of lower molariform teeth equipped with two longitudin ...
'' family have had their remains unearthed in New Jersey.


Marine life

While not much was known about Appalachia's land-based fauna until recently, the marine life that in the area, as well as the life that lived in the nearby Western Interior Seaway, has been well studied by paleontologists for years. Such examples of fossil sites that have preserved the remains of marine life from that period include the Niobrara Formation, the Demopolis Chalk Formation, and the Mooreville Chalk Formation are just a few examples of the fossil sites that have successfully preserved the skeletal remains of various marine creatures from the Cretaceous. Examples of the marine fauna that lived near Appalachia include ''
Chondrichthyes Chondrichthyes (; ) is a class that contains the cartilaginous fishes that have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fishes'', which have skeletons primarily composed of bone tissue. ...
'', '' Osteichthyes'', '' Chelonioidea'', ''
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 ''
Mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ' meaning 'lizard') comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht on ...
s'', which were the apex predators of their environment at the time. Mosasaur remains have even been unearthed in Missouri. Plesiosaur fossils, belonging to the genus ''
Cimoliasaurus ''Cimoliasaurus'' was a plesiosaur that lived during the Late Cretaceous (Maastrichtian) of New Jersey. It grew up to long and weighed up to . Etymology The name is derived from the Greek , meaning "white chalk", and , meaning "lizard", in ...
'', have been unearthed in New Jersey. Fish fossils are rather common throughout Appalachia, especially in locations abundant in marl, shale, and
limestone Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms whe ...
. Fish fossils, as well as a lot of Cretaceous era marine fauna, are rather abundant in regions like the Niobrara Formation in Kansas, which is made up of shale, sandstone and limestone, as well as the Woodbury Formation in New Jersey.


Arthropods

Many species of arthropods are known from the Turonian aged New Jersey amber, situated on the Atlantic coast of Appalachia. Arthropods are also known from the Cenomanian aged Redmond Formation of Labrador, Canada.


Flora

While the fossil sites from the southern part in Appalachia, places like Alabama and the Carolinas, have a very scant amount of Cretaceous plant fossils with the exception of Georgia, the northern parts of Appalachia, such as New Jersey, Maryland, and Delaware have a much better record in terms of plant species being unearthed there, especially with fossils sites like the Ellisdale Fossil Site, and have given us a better glimpse into a forgotten world. In the Ellisdae Fossil Site, excavations have revealed that plants like '' Picea'', ''
Metasequoia ''Metasequoia'', or dawn redwoods, is a genus of fast-growing deciduous trees, one of three species of conifers known as redwoods. The living species '' Metasequoia glyptostroboides'' is native to Lichuan county in Hubei province, China. Althou ...
'', ''
Liriodendron ''Liriodendron'' () is a genus of two species of characteristically large trees, deciduous over most of their populations, in the magnolia family (Magnoliaceae). These trees are widely known by the common name tulip tree or tuliptree for their ...
'', and possibly ''
Rhizophora ''Rhizophora'' is a genus of tropical mangrove trees, sometimes collectively called true mangroves. The most notable species is the red mangrove (''Rhizophora mangle'') but some other species and a few natural hybrids are known. ''Rhizophora'' ...
'' inhabited the region during the late Cretaceous period, implying that the environment during that time period was a coastal forest with a few types of marine environments as well including estuarine,
lagoon A lagoon is a shallow body of water separated from a larger body of water by a narrow landform, such as reefs, barrier islands, barrier peninsulas, or isthmuses. Lagoons are commonly divided into ''coastal lagoons'' (or ''barrier lagoons'') ...
al, marine, and terrestrial. Plant fossils found in neighboring states such as Delaware and Maryland have revealed that ''
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...
s'', ''
gymnosperm The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, '' Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμν ...
s'', and ''
angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants ...
'' did indeed inhabit the area. Angiosperm fossils were unearthed in the Dakota Formation in Nebraska. As mentioned earlier, Georgia has a rich fossil record of plant life dating back to the Cretaceous. Some of the examples of flora that was present in that area include ''
Salicaceae The Salicaceae is the willow family of flowering plants. The traditional family (Salicaceae ''sensu stricto'') included the willows, poplar, aspen, and cottonwoods. Genetic studies summarized by the Angiosperm Phylogeny Group (APG) have greatly ...
'', '' Lauraceae'', ''
Sequoioideae Sequoioideae, popularly known as redwoods, is a subfamily of coniferous trees within the family Cupressaceae. It includes the largest and tallest trees in the world. Description The three redwood subfamily genera are '' Sequoia'' from coasta ...
'', ''
Moraceae The Moraceae — often called the mulberry family or fig family — are a family of flowering plants comprising about 38 genera and over 1100 species. Most are widespread in tropical and subtropical regions, less so in temperate climates; however ...
'', ''
Pinophyta Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All ext ...
'', ''
Malpighiales The Malpighiales comprise one of the largest orders of flowering plants, containing about 36 families and more than species, about 7.8% of the eudicots. The order is very diverse, containing plants as different as the willow, violet, poinsett ...
'', ''
Monocotyledon Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of ...
'', ''
Ericaceae The Ericaceae are a family of flowering plants, commonly known as the heath or heather family, found most commonly in acidic and infertile growing conditions. The family is large, with c.4250 known species spread across 124 genera, making it th ...
'', ''
Cinnamomum ''Cinnamomum'' is a genus of evergreen aromatic trees and shrubs belonging to the laurel family, Lauraceae. The species of ''Cinnamomum'' have aromatic oils in their leaves and bark. The genus contains approximately 250 species, distributed in ...
'', ''
Ranunculales Ranunculales is an order of flowering plants. Of necessity it contains the family Ranunculaceae, the buttercup family, because the name of the order is based on the name of a genus in that family. Ranunculales belongs to a paraphyletic group kno ...
'', ''
Salicaceae The Salicaceae is the willow family of flowering plants. The traditional family (Salicaceae ''sensu stricto'') included the willows, poplar, aspen, and cottonwoods. Genetic studies summarized by the Angiosperm Phylogeny Group (APG) have greatly ...
'', ''
Torreya ''Torreya'' is a genus of conifers comprising six or seven species placed in the family Taxaceae, though sometimes formerly placed in Cephalotaxaceae. Four species are native to eastern Asia; the other two are native to North America. They are ...
'', '' Cupressaceae'', ''
Magnoliaceae The Magnoliaceae () are a flowering plant family, the magnolia family, in the order Magnoliales. It consists of two genera: ''Magnolia'' and '' Liriodendron'' (tulip trees). Unlike most angiosperms, whose flower parts are in whorls (rings), t ...
'' and '' Rhamnaceae''. Plant fossils of Minnesota have revealed that
cycads Cycads are seed plants that typically have a stout and woody ( ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male o ...
, evergreens,
Equisetum ''Equisetum'' (; horsetail, snake grass, puzzlegrass) is the only living genus in Equisetaceae, a family of ferns, which reproduce by spores rather than seeds. ''Equisetum'' is a "living fossil", the only living genus of the entire subclass ...
,
laurels ''Laurus nobilis'' is an aromatic evergreen tree or large shrub with green, glabrous (smooth) leaves. It is in the flowering plant family Lauraceae. It is native to the Mediterranean region and is used as bay leaf for seasoning in cookin ...
,
ferns A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except th ...
,
willows Willows, also called sallows and osiers, from the genus ''Salix'', comprise around 400 speciesMabberley, D.J. 1997. The Plant Book, Cambridge University Press #2: Cambridge. of typically deciduous trees and shrubs, found primarily on moist s ...
, redwoods, poplars, tulip trees, and pomegranates were present in the area during the Cretaceous. There is also a huge concentration of ''
Normapolles ''Normapolles'' is an extinct angiosperm pollen group that arose in the Cenomanian (Late Cretaceous). ''Normapolles'' persisted until the early Oligiocene Epoch (Tertiary Period) but were reduced in diversity by the early Eocene Epoch. ''Normapo ...
'' unearthed in the southeastern United States, suggesting that there was a distinct phytogeography through the area during the Cretaceous. Fossils unearthed near from the Redmond Formation in Labrador have revealed that many of the modern-day angiosperms first appeared in Appalachia. Plant fossils found in
Crawford County Crawford County is the name of eleven counties in the United States: * Crawford County, Arkansas * Crawford County, Georgia * Crawford County, Illinois * Crawford County, Indiana * Crawford County, Iowa * Crawford County, Kansas * Crawford Count ...
, Georgia show a wide variety of plant diversity such as
sporophytes A sporophyte () is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase. Life cycle The sporophyte develops from the zygot ...
,
gametophytes A gametophyte () is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the sex ...
, Detrusandra,
Hamamelidaceae Hamamelidaceae, commonly referred to as the witch-hazel family, is a family of flowering plants in the order Saxifragales. The clade consists of shrubs and small trees positioned within the woody clade of the core Saxifragales. An earlier syste ...
,
Actinidiaceae The Actinidiaceae are a small family of flowering plants. The family has three genera and about 360 species and is a member of the order Ericales. Distribution They are temperate and subtropical woody vines, shrubs, and trees, native to As ...
, and a multitude of 63 species of plants have been unearthed in this region. Angiosperm plants have been found in the Woodbine Formation.
Pinaceae The Pinaceae, or pine family, are conifer trees or shrubs, including many of the well-known conifers of commercial importance such as cedars, firs, hemlocks, larches, pines and spruces. The family is included in the order Pinales, formerly kn ...
and Lauraceae fossils have been unearthed in Mississippi and North Carolina respectively. Plant fossils found in Massachusetts and Rhode Island indicate that the climate here was sub-humid and paratropical too, indicating that some of Appalachia's habitats largely consisted of
coastal plains A coastal plain is flat, low-lying land adjacent to a sea coast. A fall line commonly marks the border between a coastal plain and a piedmont area. Some of the largest coastal plains are in Alaska and the southeastern United States. The Gulf Coa ...
and deciduous forests. Plants of
Pinaceae The Pinaceae, or pine family, are conifer trees or shrubs, including many of the well-known conifers of commercial importance such as cedars, firs, hemlocks, larches, pines and spruces. The family is included in the order Pinales, formerly kn ...
,
Taxodioideae Taxodioideae is a subfamily in Cupressaceae Cupressaceae is a conifer family, the cypress family, with worldwide distribution. The family includes 27–30 genera (17 monotypic), which include the junipers and redwoods, with about 130–140 sp ...
,
Araucariaceae Araucariaceae – also known as araucarians – is an extremely ancient family of coniferous trees. The family achieved its maximum diversity during the Jurassic and Cretaceous periods and the early Cenozoic, when it was distributed almost worldw ...
,
Taxaceae Taxaceae (), commonly called the yew family, is a coniferous family which includes six extant and two extinct genera, and about 30 species of plants, or in older interpretations three genera and 7 to 12 species. Description They are many-bran ...
,
Cycas ''Cycas'' is a genus of plants belonging to a very ancient lineage, the Cycadophyta, which are not closely related to palms, ferns, trees or any other modern group of plants. They are evergreen perennials which achieved their maximum diversity in ...
and
Thallophyte Thallophytes (Thallophyta, Thallophyto or Thallobionta) are a polyphyletic group of non-motile organisms traditionally described as "thalloid plants", "relatively simple plants" or "lower plants". They form a division of kingdom Plantae that inc ...
have been found in Georgia and South Carolina.


See also

*


References

{{Reflist Historical continents Geology of North America Regional geology of the United States Eastern United States