Antediluvian Beasts

Antediluvian Realms of the East: the Tar Heel and Coachman Formations

Morning, and the orange and beige of a celestial titan rising through slivers of cloud coat the swampy landscape with a buzzing, ochre light. Gently sloping mountains in the far distance, each decorated in a single shade of green trees, are the first to grasp at the brightest rays of light from the sun. Giant azhdarchid pterosaurs, drowsy and chilled by the morning air, launch off of their craggy towers in the highlands to search for food in the swamps and marshes. Within a few minutes, they reach the sky above the lowlands and sink down towards waterways beckoning with a biological buffet. The loud morning calls of ancient insects, amphibians, reptiles, and birds illuminate a scene exclusive to the Campanian. The azhdarchids land by the side of a waterhole, already taking notice of some of their company in these marshes. Nearby, a small, rough-skinned leptoceratopsid rips off leaves from some low foliage as it grunts at a small lizard picking up droppings from the dinosaur’s morning feast. This sturdy dinosaur is related to the massive ceratopsids of the west, though here it lies in the shadows of other dinosaur species. The grey and aquamarine leptoceratopsid, taking notice of the azhdarchids, sniffs and scurries into the underbrush. 

Nearby, a flock of ornithomimosaurs stops to drink in the early morning sun. Brilliantly adorned in gleaming black, blue, and grey feathers, the ostrich-mimic dinosaurs huddle close together as they hydrate for the day ahead. Two males, both decked out in feathers designed to impress the females of the crowd, start to kick and peck at each other while displaying the brilliant patterns that spur from their forelimbs. Both males are soon driven off when a mixed heard of Lophorhothon and Hadrosaurus arrive by the side of the waterhole. At 8-9 meters in length, these are all medium-sized hadrosaurs, the Lophorhothon representing a more ancient lineage of duckbills than their Hadrosaurus companions. As the large herbivores lower their heads to drink, Saurornitholestes dromaeosaurs scamper out from their feet. These small predators would do well to mind the size and temper of the hadrosaurs. 

On the opposite side of the waterhole, a herd of true giants triumphantly marches out from the surrounding swamps and bayous. These are Hypsibema crassicauda, 15 meters long and 15 tons in weight, the largest animals of these lands. The herd of hadrosaurs, thirteen strong, lumbers toward the edge of the waterhole to drink. The massive skulls of the titans slowly gulp up the water as smaller dinosaurs and crocodylomorphs flee the scene. A large female Dryptosaurus, 9 meters from head to tail, is also deterred from a quick stop for water by the Hypsibema herd. Several fresh water turtles push themselves back into the murky water. 

Back on the other side of the waterhole, a disturbance spooks the hadrosaurs. Suddenly, an ornithomimosaur is pulled underneath the water. Just off of shore, the head a giant Deinosuchus crocodyliform emerges as the massive predator tries to gulp down the dead ornithomimosaur whole. The azhdarchids take off. They’ve had enough of the lowlands. 

For the majority of the 19th and 20th centuries, the Cretaceous animals of the Carolinas have remained obscure, with decades lacking in finds punctuated by tantalizing glimpses of Cretaceous Carolinian ecosystems by means of fragmentary specimens. Ebenezer Emmons, namer of the Adirondack mountains, described isolated teeth that would later be identified as those of the massive crocodylian Deinosuchus rugosus (e.g., Emmons, 1858; Schwimmer, 2002), an noted paleontologist Edward Drinker Cope designated hadrosaur remains from a marl pit in North Carolina as the type of Hypsibema crassicauda 11 years later (Cope, 1869).

However, the past forty years have seen revelations in our understanding of the Cretaceous species of these states, including the publication of several Cretaceous-age faunas from sites within them. New discoveries of terrestrial and aquatic vertebrates from the Carolinas have illuminated several biogeographic trends from the Cretaceous of North America, supporting the notion that Appalachia was a refugium for certain vertebrate groups like as non-tyrannosaurid tyrannosauroid and basal hadrosauroid dinosaurs.

In North Carolina, perhaps the most important (in terms of terrestrial fossils) site is Phoebus Landing, a tidal-estuarine deposit located near Elizabethtown (e.g., Miller, 1967; Baird and Horner, 1979; Weishampel and Young, 1996). This site, bearing sediments of the middle-late Campanian Tar Heel Formation (e.g., Baird and Horner, 1979; Weishampel and Young, 1996; Weishampel et al., 2004; Self-Trail et al., 2004; Schwimmer et al., 2015), has not only produced skeletal elements from at least four different dinosaur species, but has also preserved teeth and bones from the massive crocodylian Deinosuchus. Among the dinosaur remains discovered here are the morphologically odd (NB: distinct from adult H. foulkii) massively constructed vertebrae of the giant (12+ meter long) hadrosauroid Hypsibema crassicauda (e.g., Baird and Horner, 1979; Weishampel and Young, 1996). Additional evidence of supermassive hadrosauroids in the Tar Heel Formation ecosystem comes in the form of a massive partial humerus only outsized by that of Magnapaulia from Phoebus Landing (Baird and Horner, 1979). Unfortunately, the fragmentary nature of these specimens means that, for now, only tantalizing glimpses of the Cretaceous giants that must have once roamed the Carolinas have been afforded by the marls. Specimens comparable to the corresponding elements in Hadrosaurus foulkii have also been recovered from the sediments at Phoebus Landing (e.g., Baird and Horner, 1979; Weishampel and Young, 1996), and Miller (1967) suggested the presence of “Hadrosaurus minor” (a dubious species of New Jersey hadrosaur possibly representative of juveniles of indeterminate genera) in the fauna present at the site. Finally, specimens comparable to the hadrosauroid Lophorhothon atopus have been recovered (e.g., Baird and Horner, 1979; Weishampel and Young, 1996). From all of this, it seems safe to conclude that two to three species of hadrosaurid were present in the Tar Heel fauna at Phoebus Landing. Importantly, a partial jawbone from a site nearby Phoebus Landing was recently described as Appalachia’s first neoceratopsian (Longrich, 2016).


Fossils from Phoebus Landing were probably deposited in a tidal or estuarine setting. Photo by the author, 2017. 


Theropods also left an important record at Phoebus Landing. Teeth and hindlimb elements possibly referable to Dryptosaurus aquilunguis have been recovered and described from the site, as have ornithomimosaur bones and dromaeosaurid teeth (e.g., Miller, 1967; Baird and Horner, 1979; Weishampel and Young, 1996; Kiernan and Schwimmer, 2004). Along with the data from ornithischian remains, the Tar Heel supported at least seven different dinosaur genera.


Lophorhothon and other hadrosauroids from the Tar Heel Formation were probably similar in form to Claosaurus (above). Photo by the author, 2017. 

Of overlapping time and geographically adjacent to the Tar Heel Formation is Coachman Formation, which in South Carolina has recently been found to contain several species of dinosaur. Schwimmer et al. (2015), in their extensive review of several sites with Coachman Formation outcrops, described remains assignable to the tyrannosauroid Appalachiosaurus montgomeriensis, the dromaeosaurid Saurornitholestes langstoni, at least one additional dromaeosaurid represented by teeth, indeterminate ornithomimosaurs, indeterminate maniraptorans, specimens comparable to “hadrosaurines”, and indeterminate hadrosauroid and ornithischian material. In addition, many species of turtles, fish, and crocodylians (including Deinosuchus) were described by Schwimmer et al. (2015) from Coachman Formation sediments, further bettering our understanding of that ecosystem.


The dromaeosaurid Saurornitholestes would have been similar to Velociraptor (skull imaged above). Photo by the author, 2015. 


The Tar Heel and Coachman Formations represent important glimpses of Cretaceous life in the Carolinas that are becoming clearer with new research into their faunas. The formations’ geographic location, sandwiched between the historically notable fossiliferous formations of New Jersey and Delaware and the more recently recognized ones of the states of Alabama, Mississippi, Tennessee, Missouri, and Georgia, mean the faunas of the Tar Heel and Coachman are essential to our understanding of the biogeography of vertebrates on Appalachia. What remains to be discovered in these deposits will certainly reveal more and more about the life forms of the obscure landmass that sat east of the Western Interior Seaway over 68 million years ago.

For more on some of the dinosaurs of the Tar Heel and Coachman Formations, see these previous posts:

Antediluvian Beasts of the East: Hypsibema crassicauda

Antediluvian Beasts of the East: Dryptosaurus aquilunguis 

PaleoNews #18


Schwimmer DR. 2002. King of the Crocodylians: the Paleobiology of Deinosuchus. Bloomington: Indiana University Press.

Cope ED. 1869. Remarks on Eschrichtius polyporusHypsibema crassicaudaHadrosaurus tripos, and Polydectes biturgidusProceedings of the Academy of Natural Sciences of Philadelphia 21:191-192.

Miller HW. 1967. Cretaceous Vertebrates from Phoebus Landing, North Carolina. Proceedings of the Academy of Natural Sciences of Philadelphia 119:219-239.

Baird D and Horner JR. 1979. Cretaceous dinosaurs of North Carolina. Brimleyana 2:1-28.

Weishampel DB and Young L. 1996. Dinosaurs of the East Coast. Baltimore: Johns Hopkins University Press. 

Weishampel DB, Barrett PM, Coria RA, Loeuff JL, Xing X, Xijin Z, Sahni A, Gomani EMP, and Noto CR. 2004. Dinosaur Distribution. In: Weishampel DB, Dodson P, and Osmólska H, eds: The Dinosauria, 2nd Edition. Berkeley: University of California Press. p. 517-617.

Self-Trail JM, Christopher RA, Prowell D, and Weems RE. 2004. The Age of Dinosaur-Bearing Strata at Phoebus Landing, Cape Fear River, North Carolina. Geological Society of America Abstracts with Programs 36(2):117.

Schwimmer DR, Sanders AE, Erickson BR, and Weems RE. 2015. A Late Cretaceous Dinosaur and Reptile Assemblage from South Carolina, USA. Transactions of the American Philosophical Society 105(2):1-157.

Longrich NR. 2016. A ceratopsian dinosaur from the Late Cretaceous of eastern North America, and implications for dinosaur biogeography. Cretaceous Research 57:199-207.

Kiernan K and Schwimmer DR. 2004. First record of a Velociraptorine theropod (Tetanurae, Dromaeosauridae) from the eastern Gulf Coastal United States. The Mosasaur 7:89-93.


Antediluvian Beasts of the East: Carnufex carolinensis

A slight breeze blows the morning dew drops from the leaves of the plants surrounding the creek as a herd of herbivorous Coahomasuchus browse horsetails and ferns. These small armored reptiles are joined by a herd of large Gorgetosuchus, which bare magnificent spikes and ridges from their armored backs. These duck-billed armored herbivores or aetosaurs as they’re known are among the larger creatures to roam these forests, although they often fall prey to the many terrestrial and aquatic predators with which they coexist. 

Among such predators are phytosaurs, large, scaly hunters which bask along the banks of rivers waiting to snap at any passers-by. These cranky reptiles are hardly the devils of these forests , and often are frightened away by little but a scratch from one of the Gorgetosuchus’s shoulder spikes. Mobs of proterosuchids also line the water’s edge, but prefer fish and amphibians to the well-defended terrestrial herbivores. Often the aetosaurs must be more wary of the smaller but poisonous Uatchitodon, which scurry around in search of insects and small vertebrates to eat. When threatened, the small carnivores cower in a defensive posture. The small size of the Uatchitodon make them hard to spot, and often an aetosaur gains a scar from accidental confrontation with such small poisonous reptiles. 

Herds of large, cranky Placerias also pose a threat to the armored aetosaurs. The sparsely haired skulls of these cow-like herbivores bare large tusks that could easily cripple an ignorant aetosaur, and individuals from both Placerias and Gorgetosuchus herds bare evidence of confrontation with their large, herbivorous contemporaries. 

The aetosaurs’ greatest foe is an aged Carnufex bull. The old predator is a giant among the inhabitants of these woods, stretching 14 feet from nose to tail. He was once longer, but a chance encounter with a 3 meter subadult Carnufex caused the old bruiser to lose a small portion of the end of his tail. The old bull has a huge territory spanning almost six square miles, and has held the area for many years. The aetosaurs are easy prey for the aged reptile, who’s instinct gives away around the spiked herbivores’ armor. 

The aetosaurs, now finding themselves on a hill, start to chew through the vegetation covering the ground. Nearby is their silent enemy, the Carnufex bull, who hides behind a large boulder. The aetosaurs sense his presence, and as he appears from his bunker lower their bodies in defense. The gnarly scarred skin of the bull Carnufex gleams in the sun as he slowly creeps across the rocky soil in an austere manner, undaunted by the glimmering spikes of his targets. The bull suddenly erupts into a fury of predatory energy, latching onto the relatively unprotected head of one of the aetosaurs. His teeth sink into the aetosaurs face, blinding the poor herbivore. Disoriented, the herbivore stumbles while the other aetosaurs flee. The Carnufex bull now has the upper hand. Using his bodyweight, he pushes over the bloodied aetosaur, exposing its unprotected stomach. All the fleeing aetosaurs hear are the moans and growls of the combatants. Soon all falls silent, and another Triassic night sets in. 

Eastern North America has preserved an excellent record of the Late Triassic, with many formations bearing the remains of strange creatures of lineages long gone. Among those which contain the most complete record of this time is the Pekin Formation, which outcrops in the states of North Carolina. The Pekin Formation dates to approximately 231 million years ago, preserving some of the oldest known Triassic archosaur faunas in North America (Zanno et. al., 2015). Among the larger predators of this formation was Carnufex carolinensis, a relative of the ancestor of the group crocodylomorpha, which includes all modern day crocodylians and many other clades of extinct genera. Carnufex carolinensis would have been a large predator in life. The type specimen, a juvenile, would have already measured 3 meters long in life (Zanno et. al., 2015).

The most interesting thing about Carnufex carolinensis is not its size, but where as a predator it was in time. The Triassic provided an open door for evolution, as the Permian extinction had created unstable biological communities and altered ecosystems (Roopnarine et. al., 2007; Bambach, Bush & Erwin, 2007). New types of large predatory animals had room to grow.


The skeleton of Prestosuchus, a large Triassic pseudosuchian. Pseudosuchians were among the groups which gave rise to large predators during the Triassic. Photo by the author, 2015. Note the phytosaur skull in the background.

What Carnufex carolinensis shows is that crocodylomorphs were more diverse during the Triassic than previously thought, and alongside early dinosaurs were diversifying to become both apex and non-apex predators (Zanno et. al., 2015). Carnufex carolinensis itself is the largest known terrestrial predator known from the Pekin Formation, and was much larger than the earliest known North American theropods (Zanno et. al., 2015).

Within the Pekin Formation, Carnufex carolinensis would have coexisted with both a variety of predatory and a variety of herbivorous animals. Bulky dicynodonts would have lumbered around next to the armored aetosaurs Gorgetosuchus, Coahomasuchus, and Lucasuchus (Green et. al., 2005; Heckert et. al., 2015; Heckert, 2012). Predatory animals like cynodonts and phytosaurs also inhabited the formation (Liu & Sues, 2010; Baird, 1986).

Carnufex represents a group of archosaurs responding to new opportunities in a recovering ecosystem. However, the large, terrestrial crocodylomorphs of the Triassic were not to last. The End Triassic Extinction would see the replacement of such large bodied forms by theropod dinosaurs (Zanno et. al., 2015). Carnufex and its contemporaries in the Pekin Formation ecosystem represent a bouncing back from extinction, a rise from oblivion.

IMG_0005 (1)

Carnufex carolinensis by the author. Pencils on paper, 2016. 




  1. Zanno LE, Drymala S, Nesbitt SJ, Scheider VP. 2015.Early crocodylomorph increases top tier predator diversity during rise of dinosaurs. Scientific Reports 5: 9276.
  2. Roopnarine PD, Angielczyk KD, Wang SC, Hertog R. 2007. Trophic Network Models Explain Instability of Early Triassic Terrestrial Communities. Proceeding of the Royal Society of London B: Biological Sciences 274: 2077–2086.
  3. Bambach RK, Bush AM & Erwin DH. 2007. Autecology and the Filling of Ecospace: Key Metazoan Radiations. Palaeontology 50: 1–22.
  4. Green JL, Schneider VP, Schweitzer M, Clarke J. 2005. New evidence for non-Placerias Dicynodonts in the Late Triassic (Carnian-Norian) of North America. Journal of Vertebrate Paleontology Programs Abstracts 25: 65-66.
  5. Heckert AB, Schneider VP, Fraser NC, Webb RA. 2015. A New Aetosaur (Archosauria, Suchia) from the Upper Triassic Pekin Formation, Deep River Basin, North Carolina, U.S.A., and its Implications for Early Aetosaur Evolution. Journal of Vertebrate Paleontology 35: e881831.
  6. Heckert AB. 2012. Two New Aetosaurs (Reptilia: Archosauria) from the Upper Triassic Pekin Formation (Deep River Basin: Newark Supergroup) of North Carolina and the Phylogeny and Distribution of Aetosaurs. Geological Society of America Abstracts with Programs 44(7): 233.
  7. Liu J & Sues HD. 2010. Dentition and Tooth Replacement of Boreogomphodon (Cynodontia: Traversodontidae) from the Upper Triassic of North Carolina, U.S.A. Vertebrate Paleontology Asiatica 48: 169–184.
  8. Baird D. 1986. Some Upper Triassic Reptiles, Footprints and an Amphibian from New Jersey. The Mosasaur 3: 125-153.

Antediluvian Beasts of the East: Sauripterus taylori

Dust swirls through the water as a school of Bothriolepis fish hustle their way through the busy stream. Bothriolepis are the bullies of these waters, shoving off other fish using their bulky bodies and stabbing those which do not give them right of way with their sharp front fins. Bothriolepis are like tanks, the armored carapace which adorns the front of their body making the armored fish sink to the bottom of stream bed. 

These Bothriolepis belong to an ancient clade of fish called placoderms. Now, the group is extremely successful, but an extinction which soon shall exterminate a variety of creatures both flourishing in the seas and just colonizing land. Bothriolepis, though tyrannical in the way they treat the other fish of the stream, are little in this regard compared to their relatives in the open sea, wihch use giant body mouth plates to crush almost any prey animal they can find. Bothriolepis are also much more vulnerable than their cousins, as predators large enough to crush their protective armor still squeeze through the cramped waterways in which the placoderms live. 

The school of Bothriolepis swim through these busy waters for a purpose. Its breeding season for these fish, and both males and females are anxious to breed and escape back into the gloom of deep lagoons that jut into the land from the seemingly infinite greenish blue sea. The Bothriolepis are taking a risk, as their only predators in these waters prefer the large lakes where the armored fish go to breed. 

The school of armored fish arrives at their choice breeding spot. The Bothriolepis mate at the bottom of the lake, away from the watch of the wretched early tetrapods which line the banks to strike at sudden movement. These sluggish beats do well to wait on the banks, as they know bigger hunters occupy the water. 

On the side of the lake, ripples stirred by a the shadow in the water set the tetrapods galloping into the woods. The shadow, almost six feet long, is long and slender in shape. The figure does not yet spook the armored placoderms, who go on mating. Suddenly, a pair of Bothriolepis are sent flying out of the water, and the head of a massive fish bites off the unarmored abdomen and tail of the male fish, sending blood flying in all directions. The female, in shock, escapes the teeth of this monstrous predator. The hunter is a rhizodont, a type of large lobe-finned fish. The skin of this beastly fish is littered with scars, showing the age and stoicism of the predator. The rest of the Bothriolepis flee, but the giant rhizodont is still able to obtain two more meals. Once the armored fish are all gone, the rhizodont paddles back into the gloom of the lake, waiting for the memory of the attack to be forgotten and for new victims to enter its domain. 

Rhizodonts were giants among the Devonian fishes which pioneered the rivers and lakes of land. These fish are famous for their large size, with largest member of the group laying claim to the record of the largest known freshwater fish ever. However, some species included in the rhizodontida have evaded the interest of the public, including a species named Sauripterus taylori which swam through the lakes and rivers of the Northeastern United States over 300 million years ago.

Sauripterus taylori is known from a couple different specimens, all retrieved from the Catskill Formation of Pennsylvania. The Catskill Formation is a mostly terrestrial deposit, and was set down in the Late Devonian during a time known as the Famennian (Davis & Shubin, 2004). In the ecosystem the Catskill represents, Sauripterus would have swam alongside fish like the armored placoderm Bothriolepis (Stein, 2002). Along the shores of the lakes and rivers the rhizodonts inhabited, plants like Euphyllophytina would soak up sunlight (Stein, 2002). Remains of possible juvenile Sauripterus have also been unearthed from the famous Red Hill locality in Pennsylvania, where the large fishes would have coexisted with both tetrapods like Hynerpeton bassetti (Shubin, 2009) and the famously giant fish Hyneria lindae (Daeschler et. al., 2007).  

The fossils known of Sauripterus taylori provide key information on the structure of rhizodontid fins. The original specimen consisted of fragments of the animal’s head, scales, vertebrae, and a well-preserved right pectoral fin (Broom, 1913). More recently, another well-preserved right pectoral fin has been described (Davis & Shubin, 2004), suggesting that the pectoral fins of Sauripterus were well-adapted for propelling the large fish through the water and pushing the animal from the bottom of the lakes and rivers in which it inhabited. The limbs of Sauripterus would have allowed it to be both gigantic and powerful, enabling it to hunt both swift and slow-moving armored prey within the waterways of Devonian Pennsylvania.

Broom, 1913 suggested the original right pectoral fin specimen of Sauripterus taylori to be nearly identical to the ancestral tetrapod limb morphology. However, it has recently been found that the similarities of the limbs of early tetrapods and rhizodonts are a case of independently evolved traits (Davis & Shubin, 2004).

Sauripterus would have been an incredible predator in life. Its large size combined with its agility would have made the fish quite an imposing predator. Like in other rhizodonts, fangs and large tusks would have peeked from the front of the jaws of Sauripterus, sinking into the flesh of fish like fishing hooks. A powerful bite would have aided the large lobe-finned fish in attacking prey. Though we known little about this ancient predator, further excavations at sites like Red Hill may result in the discovery of better specimens, revealing a hunter long forgotten among the Devonian rocks of the Eastern United States.


Sauripterus taylori by the author. Pencils on paper, 2016.


Happy Earth Day! To read the last Antediluvian Beasts of the East installment, click the link below:

Antediluvian Beasts of the East: Dryptosaurus aquilunguis




  1. Davis MC & Shubin NI. 2004. A new specimen of Sauripterus taylori (Sarcopterygii; Osteichthyes) from the Famennian Catskill Formation of North America. Journal of Vertebrate Paleontology 24(1):26-40.
  2. Stein WE. 2002. SUNY-Binghamton Paleobiology Collection. URL: . Accessed April 13, 2016.
  3. Daeschler EB, Shubin NH .2007. New data on Hyneria lindae (Sarcopterygii, Tristichopteridae) from the Late Devonian of Pennsylvania, USA. Journal of Vertebrate Paleontology 27(S3): 65A.
  4. Shubin N. 2009. Your Inner Fish: A Journey into the 3.5-Billion-Year History of the Human Body. New York: Vintage. pp. 13. 
  5. Broom. 1913. On the Origin of the Cheiropterygium. Bulletin of the American Museum of Natural History 32: 459-464.


Antediluvian Beasts of the East: Dryptosaurus aquilunguis

Summer is the season which gives life to the animals of Cretaceous New Jersey. As the trees rustle in the cool breeze, a large bull Hadrosaurus minor grumbles to attract mates. His calls can be heard for miles, and can attract unwanted attention, including rival males. Hadrosaurus minor males frequently fight, sometimes killing one another in the heat of battle. Luckily for this male, only females appear from the forest, made interested by the hadrosaur’s distinctive bellows. The females will graze in close proximity to the male until they are ready to mate. However, this time of plenty will soon end, giving way to the cold, wet months of winter.

The vast expanses of the New Egypt wilderness host more then just dinosaurs. Crocodilians and marine reptiles frequent the area during the warm summer, feasting on the schools of fish which shoot up the deltaic plains of the coastal lowlands. Among them is the 14 foot long Thoracosaurus, a long-snouted crocodilian specializing in fish-hunting. These slender predators follow the large schools of Enchodus fish up and down the eastern shoreline of Appalachia. In winter, the cold-blooded thoracosaurs will leave the deltas of the Navesink for warmer hunting sites elsewhere. Other predators of the deltas, such as the mosasaur Halisaurus, will eek out a living a few miles off the coast of Appalachia, where they will be exposed to larger predators.

For other animals of the New Egypt wilderness, the coming winter months pose more of a threat. Many of the large herbivorous dinosaurs of the region will be forced into the nearby Appalachian mountains, which in the west eclipse their summer home of plenty. To reach the plateau within the mountains, the large herbivores of the region must embark on a long trek across 50 or more miles of treacherous terrain, and many will not survive.

Fall, and the calls of another large animal fill the New Egypt wilderness. Lambeosaurines, some 26 feet in length, are forming gigantic migrating herds. Males of this dinosaur are adorned will colorful head crests which they use to deter rivals and predators. The already forming heavy proto-feather coats of these hadrosaurs signal the coming of winter. They must store up fat to survive to cold, or else they will never see spring. Juveniles are especially at risk, and often will not even survive the first days of the trek up the mountains.

For another type of animal, the coming of winter is an advantage. The top predators of the region have been busy all summer raising the next generation, but now they stir from their nests. The most common species here is Dryptosaurus aquilunguis. Packs of these killers will torment the herds of herbivorous dinosaurs making their way through mountain passes. Unlike the hadrosaurs, whose bristly and somewhat sparse coats provide only a little protection from the cold, Dryptosaurus aquilunguis sport a coat of feathers similar in structure to the downy feathers of birds. Their warm coat provides a huge advantage when cold weather sets in, allowing them to thrive in the cold winter. Their padded feet, armed at the tip with large claws, give them ample traction during their climb up the sometimes rocky faces of the Appalachian mountains.

Winter finally shows its ugly head to the animals of the Appalachian wilderness. Although there will be little or no snow, long-lasting storms will drench the deltas with ice-cold water and temperatures will drop to an average of 2 degrees Celsius. The small animals of the deltas, like salamanders and frogs, will cling on to life here as best they can, but the large animals will now have to leave. The hadrosaurs start to march up the mountains, and are followed by the hungry Dryptosaurus aquilunguis, which seek to prey on the duck-billed dinosaurs. These predators hunt and kill dozens of hadrosaurs during the start of winter, feasting on the remains of the weary herbivores. Winter, however, will even test the might of the dryptosaurs, as the worst is yet to come.

Mid-December, and the battalions of migrating dinosaurs find themselves among the cold forests of the plateau they’ve set on out for. Even in these forests, insulated by mountains all around them, the height of winter is desolate, and sleet, hail, and rain are common sights. For the groups of dryptosaurs, these are the days which put them to the test. Among the trees, a mating pair cuddles for warmth. The hadrosaurs are faring even worse then their predators, and many browse among the bodies of their brethren which have succumbed to the cold. The hadrosaurs would fare even worse back in the coastal lowlands, where the constant cold weather kills the plants of the deltas. The pine trees here barely sustain these majestic titans, which, in turn, allow the dryptosaurs to survive.

As the hadrosaurs browse, the mating pair of dryptosaurs find a native resident of these alpine woodlands. A nodosaur, decked in armor from nose to tail, rips bark off the sleet-covered conifers as the theropods watch from behind a grove of dead magnolia trees. The female dryptosaur cautiously approaches the armored herbivore, which grunts to deter her. Taking notice, the dryptosaur backs off, deciding not to mess with an unknown herbivore. The Dryptosaurus aquilunguis will have to settle for hadrosaur meat.

Late January, and signs of spring bring hope to the weary dinosaurs. The dryptosaur pair has survived, and have been tracking a small group of ornithomimosaurs for a few days. They finally strike, killing a large ornithomimosaur using their immense hand claws. These weapons are only found on dryptosaurs, giving them another advantage over the other lineages of large predator that live in the Navesink ecosystem. These theropods have braved the harsh Appalachian winter, and are rewarded by warmer weather.

This same rise in temperature signals the herds of hadrosaurs to migrate back to the lowlands, where they will be joined by groups of leptoceratopsids which have migrated back to their summer home from the south. The hadrosaurs seem joyful as they march through the same mountain passes they faced months earlier.

As the herds of dinosaurs march back, the fading winter lashes out once more. A sudden downpour of sleet and snow rivets the migrating animals, who have already lost some of their winter coats. The storm has come from the cold, high peaks of the northern Appalachians, where winter always lurks. The hadrosaurs, however, will survive, due to the pounds of conifer matter they ingested while staying in the high plateau. The dryptosaur pair escape the storm undaunted, feeding on the carcass of a dead pterosaur which fell from the skies during one of the harsh storms of mid-winter.

The hadrosaurs have finally reached the promised land. They have braved the harsh wintry months, and feast on the warm, succulent plants of the deltas. For the dryptosaur pair, the end of winter gives way to another event. The spring is the mating season of the dryptosaurs, and the female Dryptosaurus aquilunguis of the pair will soon lay a clutch of eggs. By the middle of summer, the dryptosaur pair will welcome chicks. When again the onset of fall signals the migration of the hadrosaurs, these chicks, alongside their parents, will face the sleet storms of deep winter, and, if they survive, will join their parents in being the most versatile predators of the Cretaceous.

Dryptosaurus aquilunguis remains one of the most important dinosaurs in the history of paleontology. The remains of this large predator, discovered during the late 1800s, would set the foundation for the concept that dinosaurs were extremely active, adaptable animals, contrasting with the view of them as lumbering “evolutionary failures”. To completely understand the role Dryptosaurus aquilunguis played in what would eventually be known as the Dinosaur Renaissance, we must first venture to the time when the D. aquilunguis type specimen was discovered and uncovered from a marl deposit in Barnsboro, New Jersey.

It’s the late 1800s, and the partial skeleton of a predatory dinosaur is removed from New Jersey sediment. The excavation is under the watch of Edward Drinker Cope, a young paleontologist and comparative anatomist living in Haddonfield, NJ (Gallagher, 1997).  Incredibly, the theropod specimen is more well-preserved than any other found before it, provoking much excitement among Edward Cope and his team. Cope envisioned this new killer as an active hunter, running and leaping across the ancient New Jersey environment. Cope named the animal Laelaps aquilunguis, the “eagle-clawed catcher of all its quarry”, alluding to the dog Laelaps, which, in Greek mythology, always caught whatever it pursued. The success of the discovery of Laelaps prompted Cope to continue his excavation and exploration of the animals of the New Jersey marl, and more fascinating discoveries followed.

Cope had won the lottery. He knew Laelaps was related to the European Megalosaurus (another predatory dinosaur only known from scrappy remains) and since the former was known from a specimen far more complete then the latter, Cope could continue his mentor’s (influential paleontologist Joseph Leidy) work in finding out what these ancient creatures actually looked like in life. Cope’s Laelaps started to make scientists wonder if dinosaurs were active animals rather then the slow moving-behemoths they had been thought to be previously. It turned out Cope was correct in thinking that dinosaurs were very active animals, and the legacy of his “leaping Laelaps lives on today.

All this excitement attracted another prominent young scientist by the name of Othniel Charles Marsh. Marsh and Cope had known each other previously, but now Marsh had a more devious agenda. Marsh wanted some of the New Jersey marl specimens for himself, and during his stay started to establish his own group of marl diggers (Gallagher, 1997). Marsh and his newly-found henchmen started to collect the best specimens from the marl, much to Cope’s suspicion. Marsh also took it upon himself to publicly humiliate Cope any chance he got to,with  one of the most famous incidents concerning the plesiosaur Elasmosaurus. Marsh even took away Cope’s prize animal: Laelaps. Pointing out that the genus name in question  was already given to an arthropod, Marsh re-described Cope’s pride and joy as Dryptosaurus aquilunguis, the “eagle-clawed tearing lizard” (Gallagher, 1997), a name which still stands today. Cope was devastated, and continued to use Laelaps in his papers to refer to carnivorous dinosaurs (Gallagher, 1997). In his later years, Cope  would work with the artist Charles Knight to create a restoration of two Dryptosaurus fighting each other, a piece of paleoart which still sets the standard for paleontological illustration. 

Although Cope’s Laelaps might not have lasted, his theories on the ecology of the dinosaur did. Future discoveries would paint a dynamic picture of dinosaurs as active animals, asserting that his vision of Dryptosaurus as an active animal was true. Today, Dryptosaurus and its relatives (tyrannosauroids) are known to be fast, powerful predators just as Cope had envisioned. However, today the remains of Dryptosaurus are little compared to the material we have of other dinosaurs, and so some deduction and guesswork is required to understand the species ecologically, anatomically, etc.

The holotype of Dryptosaurus aquilunguis was unearthed from the New Egypt Formation (Brusatte et. al., 2011), a Maastrichtian-age (~72-65 MYA) deposit in what would become New Jersey. The New Egypt Formation is especially important as it provides a glimpse into one of the last Mesozoic Appalachian ecosystems. In fact, just above the New Egypt lies the Hornerstown Formation, a deposit which records the faunal change from a latest Cretaceous ecosystem including animals such as Mosasaurus sp. to a Cenozoic ecosystem recovering from the K-T extinction event. However, the Hornerstown does not contain the diverse dinosaurian fauna present in the New Egypt. Within the latter formation, Dryptosaurus aquilunguis had access to a wide range of dinosaur prey, including Hadrosaurus minor, another unnamed “duck-billed” dinosaur genus, and lambeosaurine hadrosaurs. Furthermore, ornithomimosaurs of the taxon Coelosaurus antiquus were also present within the New Egypt ecosystem, providing another food source for the aptly-named “tearing lizard”.

Although many of the dinosaurs and other animals present in the ecosystem of Dryptosaurus aquilunguis were related to forms in western North America and Asia, the morphology of these Appalachian forms differed from their relatives which lived far away. This is likely due to the unique faunal interchange which had defined the fauna of Appalachia ever since its start during the Cenomanian. Before the breakup of the North American landmass during the Late Cretaceous, an almost homogenous fauna could be found across the continent. On the east coast, the presence of a dinosaurian fauna almost completely homogenous to that of the west (Deinonychus sp., Tenontosaurus sp. , titanosauriformes, nodosauridae, allosauroidea) is found within the Arundel Formation of Maryland (Weishampel, 2006). Surprisingly, the Arundel Formation also shows the beginnings of changes in the North American fauna. The remains of an indeterminate ornithomimosaur and more surprisingly a tooth assigned to an indeterminate neoceratopsian have also been unearthed (Weishampel, 2006).


Deinonychus, a likely component of both eastern and western North American dinosaurian fauna. Photo by the author, 2015.

Skip ahead a couple million years, and Cenomanian Appalachia bears a “chimaera fauna”: a mix between taxa belonging to groups which had lived across the continent before the breakup of Laramidia and Appalachia and taxa which had evolved on or migrated to Appalachia. The best record of this time on Appalachia comes from Texas, where the Cenomanian Woodbine Formation preserves the remains of both animals with relatives found in the Arundel (nodosauridae, allosauroidea, dromaeosauridae) (Main, 2013)(Lee, 1997), but also those which do not have close relatives within the earlier formation (Protohadros) (i. e. Main, 2013). However, Protohadros does seem to have close relatives in the west (Prieto-Marquez & Norell, 2010)(Wenhao & Godefroit, 2012), suggesting that faunal interchange between Laramidian and Appalachian animals did occur at least in one direction.

Travel to the Campanian, and Alabama preserves a dinosaurian fauna seemingly created by the mixing of endemic and foreign fauna. Here, a derived non-tyrannosaurid tyrannosauroid (Appalachiosaurus) and a close relatives of hadrosaurids (Lophorhothon) appear, albeit not in the same formation. Finally, the Maastrichtian and Campanian sediments of New Jersey, North and South Carolina, Delaware, and Missouri preserve a terrestrial fauna almost completely unique from one anywhere else on the globe. Giant primitive hadrosaurids (Hypsibema, Ornithotarsus, “Parrosaurus”) are found across the continent, along with oddly primitive hadrosauroids (Lophorhothon, ?Claosaurus), hadrosaurines, which varied greatly in size (Hadrosaurus foulkii, Hadrosaurus minor), and leptoceratopsids (Longrich, 2016). 


Hadrosaurus foulkii fossils displayed at the Philadelphia Academy of Natural Sciences, Drexel University. Photo from Wikipedia.

Along with these animals, which seemingly arose on Appalachia after the splitting of the North American continent, forms found on eastern North America during the Early Cretaceous days of the Arundel Formation are still found within the ecosystems of Appalachia. Ornithomimids and nodosaurs are present in New Jersey, the Carolinas, and Alabama. Dromaeosaurs are also present in Alabama and possibly the Carolinas and New Jersey. Finally, primitive tyrannosaurs (Dryptosaurus macropus, Dryptosaurus aquilunguis, Appalachiosaurus montgomeriensis), echoes from the Early Cretaceous when their kind were just starting to become large predators, are the big-game hunters of Appalachia. The presences of a distinct fauna on Appalachia is supported by Longrich (2016).

The distinct fauna of Appalachia might also explain the intriguing morphology of some of material we have of Dryptosaurus aquilunguis. The massive hands tipped with huge claws which the “tearing lizard” possessed point to the prospect of this Appalachian tyrannosauroid employing a different style of attack than that of its western counterparts. Brusatte et. al. (2011) pointed out that the forelimb morphology of D. aquilunguis was “unlike that of any other tyrannosauroids”. The 2011 study also concluded Dryptosaurus had lost some grasping ability in comparison with more basal tyrannosaurs. Furthermore, unlike the tyrannosaurids of the west which possessed large, robust skulls (i.e. Tyrannosaurus, Daspletosaurus), Dryptosaurus seems to have possessed a more delicately constructed skull and, also unlike western tyrannosaurs, ziphodont teeth (Brusatte et. al., 2011).

Clearly Dryptosaurus was employing a different hunting style than other large tyrant dinosaurs, and the reason for this was perhaps due to the different types of prey the tyrannosaurs of the east and west were hunting. These features are, however, similar to those found in allosauroids, which are sometimes thought of as hunters of large game such as sauropods and non-hadrosaur ornithopods. The hadrosaurs of the east coast can be thought of as analogous to these other herbivorous dinosaur groups as they did possess large size in many cases and also seem to have not possessed any obvious defensive features (spikes, horns, armor).


The skeleton of Tyrannosaurus rex, a tyrannosaurid tyrannosauroid showing the robust skull morphology and small arms associated with derived tyrannosaurs. Photo by the author, 2015.

Almost the exact opposite occurs in many groups of large herbivores in the west, where ceratopsids boasted a menagerie of different horns and frills, ankylosaurs were coated in a tough layer of bony scutes and plates, and pachycephalosaurs sported domed, spiky heads. The tyrannosaurids of the west lived in a land where the “plain” hadrosaurs were in well-armed company. Dryptosaurus seems to have lived in an ecosystem where large animals with obviously defensive features were less common (a leptoceratopsid is known from the Tar Heel Formation of the Carolinas (Longrich, 2016), and nodosaurid material from the Campanian and Maastrichtian New Jersey has also been recovered (Weishampel, 2006), and that’s pretty much it as far as Appalachian dinosaur with defensive capabilities go). Dryptosaurus likely would have hunted and killed hadrosaurs by slicing flesh with its teeth and large claws, thereby inducing blood loss.

Dryptosaurus aquilunguis remains one of the most astounding dinosaurs ever found. The incredible history of its collection combined with its peculiar anatomical features stand as a hallmark of paleontological discovery in eastern North America as well as in the rest of the word. The “tearing lizard” truly is an amazing dinosaur, a hallmark of the fantastic animal oddities which evolved on the forgotten continent of Appalachia.



Dryptosaurus aquilunguis rests within the coastal forest that will one day become the New Egypt Formation. Also pictured are the ornithomimosaur Coelosaurus antiquus, an unnamed lambeosaurine hadrosaur, a dromaeosaurid, and enantiornithine birds. Foliage includes Metasequoia sp. (Dawn Redwood), Liriodendron (Tulip Trees, leaves pictured), Picea sp. (Spruce Tree), rosaceae sp., and Humulus sp. Illustration by the author. Colored pencils on paper, 2016.

For more on the dinosaurs of Cretaceous Appalachia, see these articles:

Antediluvian Beasts of the East: Hypsibema crassicauda

Antediluvian Beasts of the East: Hypsibema missouriense

Antediluvian Beasts of the East: Ornithotarsus immanis

PaleoNews #18 (National Fossil Day 2015 Special Edition) 


  1. Gallagher WB. 1997. When Dinosaurs Roamed New Jersey. New Brunswick: Rutgers University Press. pp. 34-39.
  2. Brusatte SL, Benson RBJ, Norell MA. 2011. The Anatomy of Dryptosaurus aquilunguis (Dinosauria: Theropoda) and a Review of its Tyrannosauroid Affinities. American Museum Novitates 3717: 1-53.
  3. Weishampel DB. 2006. Another look at the dinosaurs of the East Coast of North America. In: Coletivo Arqueológico-Paleontológico Salense, eds: Actas III Jornadas Dinosaurios Entorno. Salas de los Infantes: Burgos. pp. 129-168.
  4. Main D. 2013. Appalachian Delta Plain Paleoecology of the Cretaceous Woodbine Formation at the Arlington Archosaur Site in North Texas. D. Phil. Dissertation: University of Texas.
  5. Lee Y. 1997. The Archosauria from the Woodbine Formation (Cenomanian) in Texas. Journal of Paleontology 71(6): 1147–1156.
  6. Prieto-Marquez A & Norell MA. 2010. Anatomy and Relationships of Gilmoreosaurus mongoliensis (Dinosauria: Hadrosauroidea) from the Late Cretaceous of Central Asia. American Museum Novitates 3694: 1–52.
  7. Wenhao W & Godefroit P. 2012. Anatomy and Relationships of Bolong yixianensis, an Early Cretaceous Iguanodontoid Dinosaur from Western Liaoning, China. In: Godefroit P, eds: Bernissart Dinosaurs and Early Cretaceous Terrestrial Ecosystems. Bloomington: Indiana University Press. pp. 293–333.
  8. Longrich NR. 2016. A ceratopsian dinosaur from the Late Cretaceous of eastern North America, and implications for dinosaur biogeography. Cretaceous Research 57: 199-207.


The Home of Some Antediluvian Beasts of the East : Ramanessin Brook

The coastal seas surrounding Appalachia left behind some of the best preserved fossil specimens in all of the East Coast. One state in particular is famous for its Cretaceous Marine fossils: New Jersey. Every year, private collectors and museum paleontologists alike flock to New Jersey to uncover ancient bones left behind by creatures which lived around 70 million years ago. One of the most famous site is Ramanessin Brook, where not only Cretaceous marine fossils but also Pleistocene fossils can be found.

Commonly, fossils from the Navesink and Wenonah/ Mt. Laurel Formations are found in the brook, providing a glimpse into the ancient past of New Jersey. Fossils dating from the Pleistocene Ice Age are also found in the brook, giving this site a bit of variation in its fossil contents. Ramanessin Brook is known for one thing in particular: shark teeth. Exquisitely preserved shark teeth are a dime-a-dozen when it comes to this place. Although seemingly nothing compared to localities in the west, sites like Ramanessin brook offer us a lot of information on the ancient sea life of the Western Atlantic Ocean.

Scapanorhynchus texanus teeth collected Ramanessin Brook, NJ.  Photo by the author, 2015.

Check out these pretty chompers! Scapanorhynchus texanus teeth collected Ramanessin Brook, NJ. Photo by the author, 2015.

Invertebrate fossils are also found in the sediments of the brook. Partial ammonite shells, belemnite guards, and even giant oyster shells are pretty common at the brook. Here are a selection of invertebrate fossils:

An ammonite carapace collected at Ramanessin Brook. Note that this is a sliver of the whole shell of the ammonite. This carapace fragment most likely belongs to Placenticeras sp. (meeki?), but might also be that of a Sphenodiscus sp. Photo by the author, 2015.

An ammonite carapace collected at Ramanessin Brook. Note that this is a sliver of the whole shell of the ammonite. This carapace fragment most likely belongs to Placenticeras sp. (meeki?), but might also be that of a Sphenodiscus sp. Photo by the author, 2015.

Here is a piece of matrix with some miscellaneous shell fragments and shell imprints. Boy was this tricky to prep! Photo by the author, 2015.

Here is a piece of matrix with some miscellaneous shell fragments and shell imprints. Boy was this tricky to prep! Photo by the author, 2015.

Vertebrate body fossils are even more common then those of invertebrates at Ramanessin Brook. By far the most common are shark teeth. During this trip, shark teeth from at least 5 different species of shark were collected. Common teeth include those of Archaeolamna kopingensis kopingensis Squalicorax kaupi, and Scapanorhynchus texanus. What’s especially interesting is that S. texanus, unlike its modern relative, lived in coastal waters. S. texanus is also the largest known species of Scapanorhynchus. 

Scapanorhynchus texanus lateral tooth. Photo by the author, 2015.

Scapanorhynchus texanus lateral tooth. Photo by the author, 2015.

Porbeagle sharks are represented by a couple genera, namely Archaeolamna kopingensis kopingensis and Cretodus borodini. These were both mid-sized sharks, with A. kopingensis kopingensis reaching 10 feet in length and C. borodini peaking at 7 feet from snout to tail.

Archaeolamna kopingensis kopingensis teeth. Photo by the author, 2015.

Archaeolamna kopingensis kopingensis teeth. Photo by the author, 2015.

The largest sharks in these waters were Squalicorax pristodontus. Only one tooth of this shark was found during my trip. These guys could reach up to 20 feet in length. Even these sharks, however, weren’t the largest predators in the sea, with large mosasaurs like Mosasaurus and Prognathodon also calling the west Atlantic home (Gallagher, 2005).

The ecosystem the Navesink and Mt. Laurel/ Wenonah Formations represent is a coastal sea. As evidenced by the shark teeth above, there was a high diversity of sharks in the Western Atlantic during the Late Campanian/ Early Maastrichtian.

These fossils are also a tricky task to identify in some circumstances. Not much has been published recently on Eastern US deposits (one of the reasons I started the series Antediluvian Beasts) , and so to accurately identify these fossils, we must look at the old literature (I’m taking literature by Joseph Leidy) and the new(ish). I will have much more to say about this site in the future, but for now I wanted to give a brief introduction. For a little more on the animals of Ramanessin Brook, refer to these past articles:

Antediluvian Beasts of The East: Scapanorhynchus texanus 


1.Gallagher, W. B. . 2005. “Recent mosasaur discoveries from New Jersey and Delaware, USA: stratigraphy, taphonomy and implications for mosasaur extinction.”  Netherlands Journal of Geosciences 84(3):241-245