Appalachia (landmass)

Source: Wikipedia, the free encyclopedia.
For other uses, see Appalachia (disambiguation).

Map of North America during the Campanian

During most of the

pachycephalosaur and ankylosaurid dominated fauna of the western part of North America, known as "Laramidia".[6]

Due to high sea levels, subsequent erosion,

Woodbine Formation, Navesink Formation,[14] Ellisdale Fossil Site,[15] Mooreville Chalk Formation, Demopolis Chalk Formation, Black Creek Group and the Niobrara Formation,[16] together with ongoing research in the area,[17] have given us a better look into this forgotten world of paleontology
.

Geography

Appalachia stretched from

lambeosaur in the area.[19]

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.[20][21][22]

Fauna

Dinosaurs

Main article: List of Appalachian dinosaurs

From the Cenomanian to the Maastrichtian, Appalachia was inhabited by various groups of dinosaurs including, hadrosauroids, hadrosaurs, nodosaurs, leptoceratopsians, indeterminate ornithopods, tyrannosauroids, dromaeosaurs, ornithomimids, and indeterminate maniraptors lived in the area. There is also fossil evidence of possible chasmosaurs, lambeosaurs, sauropods, carcharodontosaurs, caenagnathids, troodontids, and coelurosaurs that might have inhabited the area.[23][24][6][25][26] In 2022, fossils unearthed from the Woodbine Formation in Texas confirmed that carcharodontosaurs, troodontids, and coelurosaurs did indeed inhabit Appalachia.[27]

Tyrannosaurs

In Late Cretaceous North America, the dominant predators were the

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 9 m (30 ft), indicating that diversity in these basal tyrannosauroids remained high during the Late Cretaceous.[31] These fossilized teeth possibly belong to a species of Appalachiosaurus or an undescribed species of a new tyrannosaur.[32][33] 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.[31][34] Fossils in Cenomanian deposits further indicate tyrannosauroids had been a constant presence in Appalachia since the Middle Cretaceous.[35]

Dromaeosaurs

The dryptosaurs were not the only predatory dinosaurs in Appalachia. Indeterminate

maniraptorans were present in Appalachia as well.[41]

Ornithomimids

Various

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.[42][43][39][44] In 2022, fossils from the Eutaw Formation from Mississippi revealed that large ornithomimids did indeed roam Appalachia.[45]

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.[23][24] The most of these being the carcharodontosaurid Acrocanthosaurus.[46][47]

Hadrosaurs

Another common group, arguably the most widespread species in the area,

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, though in fragmentary remains,[64] Tennessee, most notably from the Coon Creek Formation.[65][66]

Hypsibema crassicauda,

deciduous plants.[72]

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.[73] In 2021, new remains of Hypsibema missouriensis, also known as Parrosaurus missouriensis, were unearthed in Missouri.[74][75][76]

Lambeosaurs

Indeterminate

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.[77] 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.[78]

Ornithopods

While ornithopod fossils have been unearthed in the eastern United States in the past, including footprints in Virginia,[79] they primarily belonged to scrappy remains and couldn't be described as distinct species, with the exception being Tenontosaurus.[80][81][82] However, this all changed with the descriptions of Convolosaurus and Ampelognathus from early Cretaceous and late Cretaceous Texas, respectively.[83][84]

Nodosaurs

The nodosaurids, 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,[85] limited to forms like Edmontonia, Denversaurus and Panoplosaurus; perhaps due to competition from the ankylosauridae; though they did thrive in isolation, most notably in Appalachia, as mentioned earlier and in the case of Struthiosaurus,[86] Europe as well. Nodosaurid scutes 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[87] in the Niobrara Formation, Alabama in Ripley Formation,[88] Mississippi, Delaware, Maryland and New Jersey, possibly belonging to a multitude of different species.[89] Five possible and best-known examples of Appalachian nodosaurids, from both the early and late Cretaceous period, include Priconodon, Propanoplosaurus, Niobrarasaurus,[90][91] Silvisaurus[92] and possibly Hierosaurus,[93] 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[94] 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

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.[101][102] 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.[103]

Birds

Several bird remains are known from Appalachian sites, most of them

Neornithes 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, which consisted of Graculavus and Telmatornis, anseriformes as represented by Anatalavis, procellariiformes with Tytthostonyx being a representative of the group, and Palaeotringa and Laornis belonging to a currently unknown group of birds.[104] Hesperornithid fossils have also been unearthed in Arkansas.[109] Some birds found in Canada such as Tingmiatornis and Canadaga were found in areas that were a part of Appalachia.[110][111] Enantiornithine birds are also known from Appalachia, as is the case of Flexomornis from the Woodbine Formation from Texas.[112]

Non-dinosaur herpetofauna

Amphibians

Through the Ellisdale Fossil Site, a good picture of Appalachia's amphibian fauna is present. Amidst lissamphibians, 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.[113]

Lizards

There is also a high degree of endemism in regards to its reptilian fauna: among squamates, the

necrosaurids
are also known.

No fossilized remains of snakes have been discovered in Appalachia during the Cretaceous period, only being found in Laramidia.[114]

Turtles

Amidst turtles, 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.[115] In Santonian Alabama occurred the giant endemic Appalachemys.[116]

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 is represented by Dakotasuchus [117] and Woodbinesuchus,[118] Alligatoridae is represented by Bottosaurus,[119] Neosuchia is represented by Scolomastax[120] and Deltasuchus,[121] Alligatoroidea are presented Deinosuchus and Leidyosuchus, Gavialoidea are represented by Thoracosaurus,[122] Eothoracosaurus,[123] and Crocodilia is presented by Borealosuchus,[124] are well established in Laramidia as well, probably indicative of their ocean crossing capacities. Deinosuchus,[125] being one of the largest crocodilians of the fossil record,[126] was an apex predator that did prey on the dinosaurs[127][128][129] in the area, the same case applies for Laramidia as well,[130][131] despite the fact that the majority of its diet consisted of turtles[132] and sea turtles.[133][134] 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.[135] 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.[136][137]

Dyrosauridae

Dyrosauridae, most notably Dyrosaurus and Hyposaurus fossils, are also known form Appalachia, particularly in New Jersey, Alabama, and South Carolina.[138]

Pholidosauridae

Only one species of

pholidosaurid is known to have lived in Appalachia; Terminonaris whose remains have been unearthed in Texas and Kansas.[139][140]

Pterosaurs

Geosternbergia, Dawndraco, and Alamodactylus.[150] Members of the Ornithocheiridae and Anhangueridae are represented by Aetodactylus and Cimoliopterus respectively.[151]

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.[152] 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.[152]

Mammals

Several types of

Metatherians are also known, including an alphadontid,[156] a stagodontid,[157] and a herpetotheriid.[158] 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.[159] Eutheria fossils, most notably molars, have also been unearthed in Mississippi. It is possible that they belong to a creature rather reminiscent to Protungulatum.[160] The genus Alphadon as well as other members of the Allotheria family have had their remains unearthed in New Jersey.[161]

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.

plesiosaurs, and mosasaurs, which were the apex predators of their environment at the time.[170][171]

Plesiosaur fossils, belonging to the genus Cimoliasaurus, have been unearthed in New Jersey.[172] Mosasaur remains have also been unearthed in Missouri.[173]

Fish fossils are rather common throughout Appalachia, especially in locations abundant in marl, shale, and limestone.[174][175] 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.[176][177]

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

pomegranates were present in the area during the Cretaceous.[181] 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.[182]

Fossils unearthed near from the

gametophytes, Detrusandra, Hamamelidaceae, Actinidiaceae, and a multitude of 63 species of plants have been unearthed in this region.[184] Angiosperm plants have been found in the Woodbine Formation.[185] Pinaceae and Lauraceae fossils have been unearthed in Mississippi and North Carolina respectively.[186][187]

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

deciduous forests.[188] Plants of Pinaceae, Taxodioideae, Araucariaceae, Taxaceae, Cycas and Thallophyte have been found in Georgia and South Carolina.[189]

See also

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