Appalachia (landmass)
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During most of the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent 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 ''cre ...
(100.5 to 66 million years ago) the eastern half of
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
formed Appalachia (named for the
Appalachian Mountains The Appalachian Mountains, often called the Appalachians, are a mountain range in eastern to northeastern North America. The term "Appalachian" refers to several different regions associated with the mountain range, and its surrounding terrain ...
), an island land mass separated from
Laramidia Laramidia was an island continent that existed during the Late Cretaceous period (99.6–66 Year#SI prefix multipliers, Ma), when the Western Interior Seaway split the continent of North America in two. In the Mesozoic era, Laramidia was an island ...
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, or the Western Interior Sea) was a large inland sea (geology), inland sea that existed roughly over the present-day Great Plains of ...
. 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, and by the end of the Cretaceous, it retreated towards the
Gulf of Mexico The Gulf of Mexico () is an oceanic basin and a marginal sea of the Atlantic Ocean, mostly surrounded by the North American continent. It is bounded on the northeast, north, and northwest by the Gulf Coast of the United States; on the southw ...
and the
Hudson Bay Hudson Bay, sometimes called Hudson's Bay (usually historically), is a large body of Saline water, saltwater in northeastern Canada with a surface area of . It is located north of Ontario, west of Quebec, northeast of Manitoba, and southeast o ...
. 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 great-circle distance, straight-line distance from the northernmost part of Western Can ...
rose. From the
Cenomanian The Cenomanian is, in the International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...
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. Campa ...
ages of the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent 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 ''cre ...
, 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, or the Western Interior Sea) was a large inland sea (geology), inland sea that existed roughly over the present-day Great Plains of ...
retreated in the
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
, Laramidia and Appalachia eventually connected. Until this connection, its
fauna Fauna (: faunae or faunas) is all of the animal life present in a particular region or time. The corresponding terms for plants and fungi are ''flora'' and '' funga'', respectively. Flora, fauna, funga and other forms of life are collectively ...
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, Asia and Europe, during the Cretaceous Period, although ancestral forms lived earlier, in the Late Ju ...
,
hadrosaurid Hadrosaurids (), also hadrosaurs 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 fami ...
, pachycephalosaur and ankylosaurid 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 Year#SI prefix multipliers, Ma), when the Western Interior Seaway split the continent of North America in two. In the Mesozoic era, Laramidia was an island ...
". 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, tran ...
of the Laurentide Ice Sheet during
Quaternary glaciation The Quaternary glaciation, also known as the Pleistocene glaciation, is an alternating series of glacial period, glacial and interglacial, interglacial periods during the Quaternary period that began 2.58 Year#SI prefix multipliers, Ma (million ...
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 The Woodbine Group is a geological formation in east Texas whose strata date back to the Early to Middle Cenomanian age of the Late Cretaceous. It is the producing formation of the giant East Texas Oil Field (also known as the "Black Giant") fr ...
, Navesink Formation, Ellisdale Fossil Site, Mooreville Chalk Formation, Demopolis Chalk Formation, Black Creek Group and the Niobrara Formation, together with ongoing research in the area, have given us a better look into this forgotten world of
paleontology Paleontology, also spelled as palaeontology or palæontology, is the scientific study of the life of the past, mainly but not exclusively through the study of fossils. Paleontologists use fossils as a means to classify organisms, measure ge ...
.


Geography

Appalachia stretched from
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and
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all the way down to the
Eastern United States The Eastern United States, often abbreviated as simply the East, is a macroregion of the United States located to the east of the Mississippi River. It includes 17–26 states and Washington, D.C., the national capital. As of 2011, the Eastern ...
and west to the
Midwestern United States The Midwestern United States (also referred to as the Midwest, the Heartland or the American Midwest) is one of the four census regions defined by the United States Census Bureau. It occupies the northern central part of the United States. It ...
. Fossils found in these regions indicate that the area was covered in
coastal plain A coastal plain (also coastal plains, coastal lowland, coastal lowlands) is an area of flat, low-lying land adjacent to a sea coast. A fall line commonly marks the border between a coastal plain and an upland area. Formation Coastal plains can f ...
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 until the end of the Cretaceous. This archipelago would have allowed dinosaurs to migrate into the Gulf Coast and possibly explain why there are some noticeable distinctions between 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 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, nodosaurs, leptoceratopsians, indeterminate ornithopods, tyrannosauroids,
dromaeosaurs Dromaeosauridae () is a family (biology), family of feathered coelurosaurian Theropoda, theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous period (geology), Period. The name Drom ...
, ornithomimids, and indeterminate maniraptors. There is also plausible fossil evidence of 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, troodontids, and coelurosaurs 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, huge predatory theropods 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'', '' Albertosaurus'' and '' Lythronax'', dryptosaurs had more in common with the basal tyrannosaurs like '' Dilong'' and '' Eotyrannus''. 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'', '' Appalachiosaurus'', and the recently discovered '' Teihivenator'' 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'', 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 were not the only predatory dinosaurs in Appalachia. Indeterminate dromaeosaur fossils, possibly belonging to Velociraptorinae and
Saurornitholestinae Eudromaeosauria (International Phonetic Alphabet, ; "true dromaeosaurs") is a subgroup of terrestrial Dromaeosauridae, dromaeosaurid theropod dinosaurs. They were small to large-sized predators that flourished during the Cretaceous Period (geolo ...
, 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 dromaeosaurid remains, tyrannosauroid 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 maniraptorans 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 as 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 that 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, caenagnathids, dromaeosaurs, and tyrannosaurs. The most of these being the carcharodontosaurid '' Acrocanthosaurus''.


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, Hadrosauroidea, and the
Hadrosauridae Hadrosaurids (), also hadrosaurs 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 fam ...
which is now considered to be their "ancestral homeland"; eventually making their way to Laramidia,
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
, 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 continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...
,
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
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 (also known as the Antarctic Ocean), it contains the geographic South Pole. ...
where they diversified into the lambeosaurine and
saurolophine Saurolophinae is a subfamily (biology), subfamily of hadrosaurid dinosaurs. It has since the mid-20th century generally been called the Hadrosaurinae, a group of largely non-crested hadrosaurs related to the crested sub-family Lambeosaurinae. How ...
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'' representing Hadrosauromorpha, '' Hypsibema crassicauda'' and '' Hypsibema missouriensis'' representing Hadrosauroidea, and '' Hadrosaurus'' and '' Eotrachodon'' representing
Hadrosauridae Hadrosaurids (), also hadrosaurs 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 fam ...
. These hadrosaurs from Appalachia seem to be closely related to the crestless hadrosaurs of Laramidia like '' Gryposaurus'' and '' Edmontosaurus'', despite the fact that they are not considered to be
saurolophine Saurolophinae is a subfamily (biology), subfamily of hadrosaurid dinosaurs. It has since the mid-20th century generally been called the Hadrosaurinae, a group of largely non-crested hadrosaurs related to the crested sub-family Lambeosaurinae. How ...
s. ''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 U.S. state, state in the upper Midwestern United States, Midwestern region of the United States. It borders the Mississippi River to the east and the Missouri River and Big Sioux River to the west; Wisconsin to the northeast, Ill ...
, though in fragmentary remains,
Tennessee Tennessee (, ), officially the State of Tennessee, is a landlocked U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders Kentucky to the north, Virginia to the northeast, North Carolina t ...
, most notably from the Coon Creek Formation. ''Hypsibema crassicauda'', over fifty feet long, was one of the largest eastern hadrosaurs, outgrowing some of more derived western hadrosaurs like ''
Lambeosaurus ''Lambeosaurus'' ( ) is a genus of hadrosaurid dinosaur that lived during the Late Cretaceous period of western North America. The first skull of ''Lambeosaurus'' found was used by palaeontologist Lawrence M. Lambe to justify the creation of ...
'' and '' Saurolophus''. The genus likely took up 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 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 e ...
, 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 remains, mostly similar to ''
Corythosaurus ''Corythosaurus'' (; ) is a genus of hadrosaurid "duck-billed" dinosaur from the Late Cretaceous Period (geology), period, about 77–75.7 million years ago, in what is now Laramidia, western North America. Its name is derived from the Anci ...
'', have been reported from New Jersey's Navesink Formation, Bylot Island and
Nova Scotia, Canada Nova Scotia is a Provinces and territories of Canada, province of Canada, located on its east coast. It is one of the three Maritime Canada, Maritime provinces and Population of Canada by province and territory, most populous province in Atlan ...
. 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. Campa ...
. 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.


Ornithopods

While ornithopod fossils have been unearthed in the eastern United States in the past, including footprints in Virginia, they primarily belonged to scrappy remains and couldn't be described as distinct species, with the exception being '' Tenontosaurus''. However, this all changed with the descriptions of '' Convolosaurus'' and '' Ampelognathus'' from early Cretaceous and late Cretaceous Texas, respectively.


Nodosaurs

The nodosaurids, a group of large, herbivorous armored dinosaurs resembling
armadillo Armadillos () are New World placental mammals in the order (biology), order Cingulata. They form part of the superorder Xenarthra, along with the anteaters and sloths. 21 extant species of armadillo have been described, some of which are dis ...
s, 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'', '' Denversaurus'' and '' Panoplosaurus''; 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–Pal ...
; 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'' "Scutum (shield), 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 Bird anatomy#Scales, birds. The ter ...
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 landlocked U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to the west. Kansas is named a ...
in the Niobrara Formation, Alabama in Ripley Formation, 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'', '' Propanoplosaurus'', '' Niobrarasaurus'', '' Silvisaurus'' and possibly '' Hierosaurus'', 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.


Ceratopsians

While remains of the advanced ceratopsians, most notably the centrosaurines and chasmosaurines which were very common in Laramidia during this time period, they were long thought to be nonpresent in Appalachia, however this notion changed in 2016 when a ceratopsian tooth was unearthed in Mississippi's Owl Creek Formation, which has 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 while the centrosaurines had completely vanished from most of North America, though they were thriving in Asia as in the case of '' Sinoceratops,'' and most likely entered Appalachia as soon as the Western Interior Seaway closed.


Leptoceratopsians

Just like their larger cousins, the leptoceratopsids were thought to be absent from Appalachia until the mid-2010s when a jaw bone of an indeterminate genus was unearthed in North Carolina. 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. Campa ...
-era leptoceratopsid
ceratopsia Ceratopsia or Ceratopia ( or ; Ancient Greek, Greek: "horned faces") is a group of herbivore, herbivorous, beaked dinosaurs that thrived in what are now North America, Asia and Europe, during the Cretaceous Period (geology), Period, although ance ...
n 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. 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 (although it may be a case of preservation bias). It is not completely understood how the leptoceratopsians were able to reach Appalachia; however, the most commonly accepted theory was that they island hopped from Europe 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 ''Ajkaceratops'' (pronounced "oi-ka-sera-tops") is a genus of ornithischian dinosaur described in 2010. It lived during the Late Cretaceous in the western Tethys Ocean, Tethyan archipelago, in what is now Europe. The type species, ''A. kozmai'', ...
'', 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. Campa ...
.


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 Avialae, avialans closely related to the ancestors of modern birds. They inhabited both marine and freshwater habitats in the Northern Hemisphere, and include genus, genera such ...
like '' Hesperornis'', '' Canadaga'', '' Baptornis'', '' Fumicollis'', '' Parahesperornis'', and '' Ichthyornis'',
enantiornithes The Enantiornithes, also known as enantiornithines or enantiornitheans in literature, are a group of extinct Avialae, avialans ("birds" in the broad sense), the most abundant and diverse group known from the Mesozoic era. Almost all retained teet ...
like '' Halimornis'' and ornithurans like '' Apatornis'' and '' Iaceornis'', possibly indicating that Appalachia may have possessed 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 U.S. state, state located in both the Mid-Atlantic States, Mid-Atlantic and Northeastern United States, Northeastern regions of the United States. Located at the geographic hub of the urban area, heavily urbanized Northeas ...
, arguably one of the best records for Cretaceous birds as some specimens were preserved in the greensands 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 lightweight ...
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 ...
, which consisted of '' Graculavus'' and '' Telmatornis'',
anseriformes Anseriformes is an order (biology), order of birds also known as waterfowl that comprises about 180 living species of birds in three families: Anhimidae (three species of screamers), Anseranatidae (the magpie goose), and Anatidae, the largest f ...
as represented by '' Anatalavis'',
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 a representative of the group, and '' Palaeotringa'' 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'' 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 amphibian fauna is present. Amidst
lissamphibia The Lissamphibia (from Greek λισσός (lissós, "smooth") + ἀμφίβια (amphíbia), meaning "smooth amphibians") is a group of tetrapods that includes all modern amphibians. Lissamphibians consist of three living groups: the Salientia ( ...
ns, there is evidence for sirenids (including the large '' Habrosaurus''), the batrachosauroidid salamander '' Parrisia'', hylids, and possible representatives of '' Eopelobates'' and '' Discoglossus''. These show close similarities to European faunas, but aside from '' Habrosaurus'' (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 iguanids, helodermatids, and necrosaurids are also known.


Snakes

Fossilized remains of
snake Snakes are elongated limbless reptiles of the suborder Serpentes (). Cladistically squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales much like other members of the group. Many species of snakes have s ...
s are rare in Appalachia and much more common in Laramidia most likely due to preservation bias. However, in the late 1980s, fossil remains of a Cretaceous era snake, attributed to '' Coniophis'', were unearthed in North Carolina.


Turtles

Amidst
turtle Turtles are reptiles of the order (biology), order Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Crypt ...
s, which are rather common finds in Appalachia, '' Adocus'', '' Bothremys'' and '' Chedighaii'' are well represented, the latter two 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''. Two marine stem group- trionychians, "''Trionyx''" ''priscus'' and "''Trionyx''" ''halophilus'', are known from the eastern coastal margin of Appalachia (New Jersey, Delaware, Maryland and the Carolinas).


Crocodiles

Crocodiles were rather abundant in Appalachia with nine local
crocodilian Crocodilia () is an Order (biology), 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 crocodylomorp ...
genera belonging to several confirmed families, with the possibility of much more undiscovered crocodiles waiting to be unearthed. Goniopholididae is represented by '' Dakotasuchus'' and '' Woodbinesuchus'',
Alligatoridae The family (biology), 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 Am ...
is represented by '' Bottosaurus'', Neosuchia is represented by '' Scolomastax'' and '' Deltasuchus'',
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 m ...
are presented ''
Deinosuchus ''Deinosuchus'' is an extinct genus of eusuchian, either an Alligatoroidea, alligatoroid Crocodilia, crocodilian or a stem-group crocodilian, which lived during the Late Cretaceous around . The first remains were discovered in North Carolina ...
'' and '' Leidyosuchus'', Gavialoidea are represented by '' Thoracosaurus'', '' Eothoracosaurus'', and
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 ...
is presented by '' Borealosuchus'', 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 or superpredator, 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 hig ...
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, loggerh ...
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, most notably '' Dyrosaurus'' and '' Hyposaurus'' fossils, are also known form Appalachia, particularly in New Jersey, Alabama, and South Carolina.


Pholidosauridae

Only one species of
pholidosaurid Pholidosauridae is an extinct Family (biology), family of aquatic neosuchian mesoeucrocodylian crocodylomorphs. Fossils have been found in Europe (Denmark, England, France, Germany, Spain and Sweden), Africa (Algeria, Niger, Mali, Morocco and Tu ...
is known to have lived in Appalachia; ''
Terminonaris ''Terminonaris'' is a genus of extinct pholidosaurid crocodyliforms that lived during the Late Cretaceous (Cenomanian and TuronianWu X-C, Russell AP, & Cumbaa SL. 2001. ''Terminonaris'' (Archosauria: Crocodyliformes): new material from Saskatchew ...
'' whose remains have been unearthed in Texas. Kansas, and Minnesota.


Pterosaurs

Pterosaur Pterosaurs are 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 million to 66 million years ago). Pterosaurs are the earli ...
fossils, mostly similar to ''
Pteranodon ''Pteranodon'' (; from and ) is a genus of pterosaur that included some of the largest known flying reptiles, with ''P. longiceps'' having a wingspan of over . They lived during the late Cretaceous geological period of North America in presen ...
'' and '' Nyctosaurus'', have been unearthed in Georgia, Alabama and Delaware. On a similar note, azhdarchid remains, which belong to '' Arambourgiania'', have been unearthed in Tennessee. Azhdarchid remains have also been unearthed in North Carolina. 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 landlocked U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to the west. Kansas is named a ...
and
Texas Texas ( , ; or ) is the most populous U.S. state, state in the South Central United States, South Central region of the United States. It borders Louisiana to the east, Arkansas to the northeast, Oklahoma to the north, New Mexico to the we ...
.
Pteranodontidae The Pteranodontidae are a Family (biology), family of large pterosaurs from the Late Cretaceous of North America and possibly other continents including Europe and Africa. The family was named in 1876 by Othniel Charles Marsh. Pteranodontids had a ...
seems to be the most common group of pterosaurs in Appalachia and is represented by five species: ''
Pteranodon ''Pteranodon'' (; from and ) is a genus of pterosaur that included some of the largest known flying reptiles, with ''P. longiceps'' having a wingspan of over . They lived during the late Cretaceous geological period of North America in presen ...
'', '' Nyctosaurus'', '' Geosternbergia'', ''
Dawndraco ''Dawndraco'' is a controversial genus of pteranodontidae, pteranodontid pterosaur from the Late Cretaceous of North America. It is likely synonymous with the contemporary pteranodontid ''Geosternbergia.'' ''Dawndraco'' was named by Alexander K ...
'', and '' Alamodactylus''. Members of the Ornithocheiridae and Anhangueridae are represented by '' Aetodactylus'' and '' Cimoliopterus'' respectively.


Choristodera

The remains of indeterminate 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
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s are also present in Ellisdale and in both of the Carolinas. The most common are ptilodontoidean multituberculates, 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 alphadontid, 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 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'; ), also called Pan-Placentalia, is the clade consisting of Placentalia, placental mammals and all therian mammals that are more closely related to placentals than to marsupials. Eutherians ...
'' fossils, most notably molars, have also been unearthed in Mississippi. It is possible that they belong to a creature rather reminiscent to '' Protungulatum''. The genus '' Alphadon'' as well as members of the
Allotheria Allotheria (meaning "other beasts", from the Ancient Greek language, Greek , '–other and , '–wild animal) is an extinct clade of mammals known from the Mesozoic and early Cenozoic. Shared characteristics of the group are the presence of lower ...
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 of jawed fish that contains the cartilaginous fish or chondrichthyans, which all have skeletons primarily composed of cartilage. They can be contrasted with the Osteichthyes or ''bony fish'', which have skeleto ...
,
osteichthyes Osteichthyes ( ; ), also known as osteichthyans or commonly referred to as the bony fish, is a Biodiversity, diverse clade of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondricht ...
, chelonioids,
plesiosaurs The Plesiosauria or plesiosaurs are an Order (biology), order or clade of extinct Mesozoic marine reptiles, belonging to the Sauropterygia. Plesiosaurs first appeared in the latest Triassic Period (geology), Period, possibly in the Rhaetian st ...
, and
mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Ancient Greek, Greek ' meaning 'lizard') are an extinct group of large aquatic reptiles within the family Mosasauridae that lived during the Late Cretaceous. Their first fossil remains wer ...
s, which were the apex predators of their environment at the time. Plesiosaur fossils, belonging to the genus '' Cimoliasaurus'', have been unearthed in New Jersey. Mosasaur remains have also been unearthed in Missouri. Fish fossils are rather common throughout Appalachia, especially in locations abundant in
marl Marl is an earthy material rich in carbonate minerals, Clay minerals, clays, and silt. When Lithification, hardened into rock, this becomes marlstone. It is formed in marine or freshwater environments, often through the activities of algae. M ...
,
shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of Clay mineral, clay minerals (hydrous aluminium phyllosilicates, e.g., Kaolinite, kaolin, aluminium, Al2Silicon, Si2Oxygen, O5(hydroxide, OH)4) and tiny f ...
, and limestone. 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'', ''Liriodendron'', and possibly ''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 Estuary, estuarine, lagoonal, marine, and terrestrial. Plant fossils found in neighboring states such as Delaware and Maryland have revealed that ''ferns'', ''gymnosperms'', and ''Flowering plant, angiosperms'' 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'', ''Lauraceae'', ''Sequoioideae'', ''Moraceae'', ''Pinophyta'', ''Malpighiales'', ''Monocotyledon'', ''Ericaceae'', ''Cinnamomum'', ''Ranunculales'', ''Salicaceae'', ''Torreya'', ''Cupressaceae'', ''Magnoliaceae'' and ''Rhamnaceae''. Plant fossils of Minnesota have revealed that cycads, evergreens, Equisetum, laurels, ferns, willows, redwoods, poplars, tulip trees, and pomegranates were present in the area during the Cretaceous. There is also a huge concentration of ''Normapolles'' 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, Georgia, Crawford County, Georgia show a wide variety of plant diversity such as sporophytes, gametophytes, Magnoliids, Detrusandra, Hamamelidaceae, Actinidiaceae, and a multitude of 63 species of plants have been unearthed in this region. Angiosperm plants have been found in the Woodbine Formation. Pinaceae 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 and deciduous forests. Plants of Pinaceae, Taxodioideae, Araucariaceae, Taxaceae, Cycas and Thallophyte 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