From the Dartmouth College motto adapted from the Gospel of Mark and
subtitle of Merle Graffam's treatise entitled Fossils from the Tropic Shale
Although some dinosaurs may have spent time feeding in open water and possibly a few may have become strongly amphibious as implied by some trackways, it’s a common misconception that dinosaurs colonized the seas. If so, what were the bones of a terrestrial dinosaur – a new species of therizinosaur - doing amongst the marine fauna of the Late Cretaceous Tropic Shale, at least 60 miles from the nearest dry land at the time?
|Artist Victor Leshyk’s portrayal of the proto-feathered, Late Cretaceous dinosaur Nothronychus graffami dining upon mangroves growing marginal to the Western Interior Seaway. The therizinosaur is thought capable of balancing tripodally on its massive pelvis while raking in tree branches with its long slender claws, which it passed to its toothless beak.|
From the Museum of Northern Arizona. Visit Victor here. Visit MNA here.
WELCOME TO THE MOON
If you’ve ever visited the badlands outside of the tiny southern Utah town of Big Water, you know the meaning of the word “barren”. The landscape consists of a coarse, brownish sandstone bedrock covered by a monotonous repetition of eroding, low-slung, blue-gray mounds of fine mud turned-to-shale against a backdrop of buff-colored, sandstone cliffs littered at the base with dislodged blocks of stone. Little grows and nothing moves, other than the wind and the imperceptible forces of gravity and erosion that are incessantly at work.
The region is so drab and desolate that locals call it ‘The Moon.’ To geologists and paleontologists – who are of the same ilk - it’s all hauntingly beautiful and exciting beyond anything imaginable, not just for its appearance but for the story of its formation and the bounty of lifeforms that are preserved. Personally, I couldn’t wait to get out there with geologist and acclaimed author Wayne Ranney, and Merle Graffam - namesake of the dinosaur Nothronychus graffami.
INTRODUCING MERLE GRAFFAM
Indeed, there’s nary a soul in sight on The Moon unless you stumble upon Merle - retired commercial artist, Big Water resident, Bureau of Land Management Park Ranger at the Big Water Visitor Center, and amateur paleontologist par excellence. Merle takes regular walks on the Moon, combing the ancient seabed for marine fossils with the intuition, trained eye, laser focus and insatiable curiosity of a seasoned field expert.
CREATURES OF, ABOVE AND ALONGSIDE THE SEA
Merle knows The Moon and the fossils preserved within in it, all creatures of a long gone sea - megafaunal marine and brackish-water invertebrates such as oysters, gastropods, solitary corals, inoceramid bivalves and ammonites, and marine vertebrates such as fish, rays and sharks, turtles, crocodilians and an occasional short-necked plesiosaur. Above the sea soared pterosaurs and early avians with toothed-beaks. As bleak and depauperate as the landscape looks now, at one time, The Moon was the site of a thriving marine ecosystem.
|Creatures above and within the Late Cretaceous Western Interior Seaway|
Adapted from nd.gov
Terrestrial deposits marginal to the sea preserve extensive skeletal remains and trackways that attest to a diverse dinosaur fauna that plied the shoreline's habitats, while diminutive, insectivorous mammals hid in the shadows amongst the gymnosperms and newly-evolved angiosperm plants. It takes considerable imagination to view this ancient land and seascape while standing on the landscape of The Moon.
AS LUCK WOULD HAVE IT
On one of Merle's lunar constitutionals in 2000, he made an unsuspecting discovery that would change his life. What's more, it would rewrite a portion of dinosaur phylogeny, offer a new perception of dietary plasticity amongst theropods and expand our knowledge of biodiversity within the Cretaceous ecosystem.
Merle discovered a small toe-bone - a phalange - in the Tropic Shale that eventually would lead to the remains of a spectacular dinosaur skeleton at the crest of a small hillock of eroding marine sediments of the Tropic Shale. It proved to be the most complete therizinosaurid yet discovered.
|Very excited at the excavation site, Merle exclaimed, "Here's the spot!"|
LET'S GET OUR BEARINGS
With a population barely of 475, the settlement of Big Water is a tiny speck on the map (green laser dot) located in Kane County on Highway 89 in southernmost Utah near the Arizona border. On maps from the late 50’s and early 60’s, it's called Glen Canyon City and housed workers who built the nearby Glen Canyon Dam.
The name Big Water seems a misnomer, since the high desert and badlands are as dry as a bone with an average rainfall of barely six inches a year. The nearest “big water” is a slender arm of Lake Powell called Wahweap Bay about 10 miles down the highway to the southeast, where a trip downstream leads to the dam that impounds the Colorado River. So where’s all the water at Big Water?
|Wayne Ranney aims his laser-pointer at Big Water on a topographical relief map at the Big Water Visitor Center. The highway takes you down and across the Glen Canyon Dam that impounds Lake Powell.|
The three sections of the Monument record sedimentation throughout the Mesozoic. The centrally-located Kaiparowits Plateau section is exemplified by plateaus, buttes and mesas carved in rocks acquired in the Cretaceous when the region was situated along the western shore of an extensive inland body of shallow water called the Western Interior Seaway.
|Three sections of the Grand Staircase-Escalante National Monument in southern Utah with Big Water on Highway 89 near the Arizona border.|
Modified from Utah Geological Association, Second Edition DVD.
The Sevier front consisted of a fault zone, an active volcanic arc, low-angle thrust slices and a broad foreland basin. The retroarc basin - so called because it was 140-200 km cratonward of the thrust front - was an asymmetric depression created in response to the load superimposed by the east-advancing wedge of thrust sheets that downwarped the lithosphere. In response to ongoing Sevier thrusting, the foreland migrated eastward and continued to rapidly subside. The basin received massive amounts of detritus delivered by rivers across alluvial plains from the encroaching front from the west and southwest.
BIRTH OF AN INLAND SEA
In the Early Cretaceous (Aptian to Albian), the basin began to flood with marine waters from the north and south, connecting the Boreal and Tethyan seas. By the Late Cretaceous, long arms of the sea converged forming an inland epicontinental sea (epi is Greek for above). Nearest the front, deep-water sediments pass upward into shallow-water sediments recorded with conglomerates that pass into sandstones and shales, which in turn pass into carbonate marine sediments well to the east.
|Late Cretaceous oblique, north view of the asymmetric Western Interior Seaway illustrating the subducting Farallon slab, the Sevier orogen and Western Interior Seaway.|
Modified from Wikipedia
"WHENCE THE FLOOD COMETH" - GEOLOGICAL NOT BIBLICAL
The development of the inland sea occurred by active subsidence of the foreland but was assisted at a time of eustasy (global high seas). Sea level changes are affected by the volume of water contained in the ocean basins and the volume of water displaced from the basins. For example, melting polar ice adds to the basins causing glacioeustasy, and shifting plates and shallowing basins removes water called tectonoeustasy. It's a rather simplistic scenario but not far from reality.
Pangaea's aridity during the Triassic and Jurassic - demonstrated by widespread eolian sandstones and evaporites in the west - was replaced by a humid, subtropical climate in the Cretaceous, as North America drifted out of lower, equatorial latitudes. Concurrently, as Atlantic seafloor spreading increased, the ocean basin shallowed, displacing vast quantities of water, while extrusive continental volcanics associated with rifting elevated temperatures 18°F (10°C) higher than average.
Submitting to the global greenhouse, melting polar ice further drove seas higher. Low-lying regions - coasts, interior lowlands and cratonic platforms - drowned worldwide including the subsiding basin of the Sevier foreland. And in its wake, the seas left vast sequences of sedimentary rocks. The great flood is known as the Zuni transgression - the greatest of six major high water events of the Phanerozoic. As an aside, our modern world with rising seas is in a state of Holocene (post-Pleistocene) glacioeustasy. Now back to the Cretaceous Seaway!
|The six major transgressions of the Phanerozoic Eon with the Cretaceous Zuni highlighted.|
Modified from Earth System History, Second Edition, 2005 and msubillings.edu.
THE WESTERN INTERIOR SEAWAY
At its zenith in the Late Cretaceous, the Western Interior Seaway in places was almost 300 meters deep. Inland seas are built on buoyant continental platforms and are relatively shallow compared to deeper-denser ocean basins. The sea connected the Arctic Ocean and Hudson Bay with the Gulf of Mexico, and stretched from Utah in the west to the western Appalachians in the east. It split North America into two massive landmasses - eastern Appalachia and western Laramidia - and divided the terrestrial ecosystem forcing it to pursue independent courses of evolution, as did the resident faunal populations riding on Pangaea's drifting continental siblings.
Also in the Late Cretaceous, Laramidia formed an arctic land-based connection with northeast Asia called Beringia. The loosely defined region in the vicinity of the Bering Strait has intermittently persisted through Recent times. During the Pleistocene, an Ice House climatic condition created regressive global seas exposing the land bridge; whereas during the Greenhouse conditions of the Cretaceous, the land was devoid of polar ice, having formed tectonically from a series of accretionary events. Like the Pleistocene connection that allowed the passage of Paleo-Indians and mammalian megafauna (the Asian saber-toothed cat comes to mind), the Cretaceous bridge (up to a 1,000 miles wide) likely allowed faunal and floral exchange in a similar manner in both directions.
|Approximate extent of the Beringia Land Bridge|
Adapted from ic.arizona.edu
In the Late Cretaceous, Laramidia experienced a major evolutionary radiation of dinosaurs possibly related to new biomes generated by the Sevier front and foreland, and may have been infused by immigrant fauna that migrated across the bridge from Asia (or vice-versa). The relevance of Cretaceous paleography will become relevant in our forthcoming discussion of therizinosaurs from Laurasia (the combined landmasses of North America and Eurasia that formed Pangaea with Gondwana of the Southern Hemisphere).
TRANGRESSIONS, REGRESSIONS AND DEPOSITIONAL SEQUENCES OF THE VACILLATING SEA
During the Late Cretaceous for nearly 25 million years, the Western Interior Seaway dominated paleography and sedimentation over a vast area of the Southwest. At least two major and numerous minor transgression-regression sequences - called cyclothems - are recorded in the rock record.
Marine waters advanced onto the continent's downwarping interior, rising and falling with starts and stops while the shoreline shifted to and fro from east to west. As the sea advanced onto land, the sandy shore was buried by new, higher shores, while previously deeper muds migrated as well. Terrestrial deposits met marine that vied for space in an overlapping, alternating geometry, all related to the whim of the vacillating sea. When the sea eventually reversed its direction, the opposite layered architecture was deposited as newer shorelines formed on previously deeper muds called a transgressive-regressive sequence - visible stratigraphically.
The west part of the GSENM was elevated by Sevier tectonics before sediments were deposited in coastal areas ahead of the encroaching inland sea from the east. All Upper Jurassic and a good part of Middle Jurassic rocks were removed by erosion before Cretaceous sediments were deposited.
THE FIRST TRANSGRESSIVE SEQUENCE
Initial alluvial plain and coastal plain deposits were met by the sea's rapidly-rising westward advance called the Greenhorn Cyclothem (late Cenomanian to middle Turonian). Deposited in the sea's first transgression in the early Late Cretaceous about 95 million years ago came coarse, yellow-brown beach sands of the shallow marine Dakota Formation, deposited on either the Morrison Formation (east) or the Entrada Sandstone (west). The Dakota contains a record of shallow brackish and marine water environments, lush coastal swamps and sandy expanses incised by rivers and streams emptying into the sea.
In deeper waters, the Dakota grades into dark, organic-rich Mancos Shale - called the Tropic Shale regionally - and consists of exceptionally fossiliferous blue-gray silts and muds formed about 93 million years ago. The type section crops out around the town of Tropic, Utah, about 50 miles to the northwest. Elsewhere in Utah, Tropic stratigraphic equivalents have been referred to the Tununk Member of the Mancos Shale, the Tropic equivalent in most of the Southwest.
On top of the sequence with the sea retreated to the east lies the four-membered Straight Cliffs Formation, an overall regressive sequence rich in coal that followed the previous marine incursion about 85 million years ago. The sea returned again bringing with it another sequence of deposits, seen elsewhere on the Kaiparowits Plateau and in the Grays Cliffs of the Grand Staircase.
The aforementioned lithologies are conformable and form a classic transgressive-regressive sequence that documents the greatest widespread rise in sea level of the Cretaceous recognized worldwide. In summary, the foreland basin's sedimentary infill represents a record Sevier orogen tectonics, flexural subsidence, weathering and sedimentation and eustatic sea level change.
|The dissected landscape rocks of The Moon of Big Water preserve Upper Cretaceous transgressive Dakota sandstone, shale and some coal buried beneath eroding gray Mancos muds and regressive cliff-forming Straight Cliffs sands and coals.|
"HEY DAVE! WHAT'S THIS?"
In 2000, at the conclusion of a large plesiosaur excavation in the Tropic, Merle turned to Dr. Dave Gillette - Utah's former state paleontologist and current Colbert Curator of Vertebrate Paleontology at the Museum of Northern Arizona in Flagstaff. Pulling a bone from his pocket, Merle uttered the now famous phrase "Hey Dave! What's this?"
Dave recognized the toe bone, but it was clearly not from a plesiosaur, the large marine reptile found with increasing frequency on the Tropic seabed thanks to Merle's keen eye of discovery. The bone was too small to be from a hadrosaur, a terrestrial, duck-bill dinosaur found in large numbers along the shoreline far to the west.
Stumped by the implication of a dinosaur bone so far from land, they later returned to the site, found more bones and initiated an excavation. The dinosaur's identity was a mystery well into the dig. According to Dave, “We weren’t thinking ‘therizinosaur’ at first, because at that time they were known only from Asia. From that first toe bone, we thought maybe we had a big ‘raptor’ (an agile, hunting dinosaur). But when we found peculiar bones of the massive hips, we knew we had a sickle-claw dinosaur. They were like nothing we’d ever seen.”
|The active therizinosaur excavation site in the Tropic Shale. A project can require the removal of up to 20 tons of overburden and take a thousand hours of field and laboratory time. |
Photo by Dave Gillette
For Merle's discovery and contribution, Graffam became the species namesake. Following the dinosaur's reconstruction, the therizinosaur was featured in an exhibit at the Museum of Northern Arizona from 2007-2009 and at the Carl Hayden Visitor Center at the Glen Canyon Dam in 2012. The actual bones of N. graffami are in storage at the Natural History Museum of Utah, Salt Lake City. Several casts are on display such as the one inside the entrance to the Museum of Northern Arizona in Flagstaff.
For half a century, therizinosaurs have remained a poorly known and understood group of theropod dinosaurs with an extremely unusual combination of anatomical features. That's changed largely in the last decade with new discoveries in Cretaceous deposits in Mongolia, China and western North America.
Unlike earlier theropod dinosaurs that exhibit predatory morphological adaptations and carnivorous inclinations, therizinosaurs exhibit the characteristics inherent of herbivores. These are thought to include: tightly-packed, leaf-shaped cheek teeth (as opposed to elongate, typically Theropod meat-cutting teeth), an inset tooth row (suggesting fleshy cheeks necessary for plant mastication) in tandem with a rostral rhampotheca (keratinous, toothless bird-like beak to facilitate an herbivorous diet), a massive, highly derived pelvis (to accommodate a large gut synonymous with plant digestion), the development of large load-bearing hind limbs (to support a large abdomen), the loss of cursorial hind-limb adaptions (typical of predatory, swift carnivorous theropods) and an increased vertebral count (long neck speculated to increase browsing range similar to sauropods).
Therizinosaurs, especially more derived forms such as Nothronychus graffami, are thought to have been slow, large waddling, pot-bellied creatures rather than the quick, graceful gaited members of related Theropod predators. In spite of their likely herbivory, the group is thought to possess defensive capabilities with its powerful claws.
The tail was short and unnecessary for its mode of non-predatory diminished speed and thought to have provided upright, tripodal support for plant consumption. Unlike most theropods, the pes was curiously tetradactyl (four toes, which is a throwback to the ancestral dinosaur condition) with blunt unguals (claws), while its manus was tridactyl (three fingers) with elongated, recurved claws - the distinctive anatomical feature that gives the clade its name. Therizinosaur means "sickle-claw reptile". These Therizinosauroid features became increasing expressed from basal forms through more derived forms.
What's more, being members of Theropoda, the entire clade was thought to possess rudimentary proto-feathers - integumentary-derived structures such as hair, scales and nails. Please visit my daughter's post on feathers here for the evolution of this dinosaurian structure.
|An imaginative interpretation of a proto-feathered adult therizinosaur accompanied by juveniles|
Used with permission by artist Damir G. Martin. Visit him here.
Taking their singular, albeit bizarre morphology and fragmentary fossil record into account, it comes as no surprise that therizinosaurs have endured a convoluted taxonomic history within Dinosauria and have been variously assigned to nearly all of its major subclades.
At one time, the family Therizinosauridae was referred to as the now-outmoded, group Segnosauria (segnis means slow in Latin) based on their heavy bodies, short legs, and sloth-like claws with a comparable lifestyle. They have been variously regarded as gigantic turtles, aberrant theropods and sauropodomorphs. Based on their retroverted (opisthopubic) pelvis (posteroventrally-directed pubis bone which was aimed backwards as in ornithischian, bird-hipped dinosaurs), they were considered to be phylogenetic intermediates between herbivorous prosauropods and early ornithischians.
UNEQUIVOCAL PHYLOGENETIC RESOLUTION
With increasing numbers of discoveries in Asia and North America from the Cretaceous, the diversity of therizinosaurs has begun to exhibit remarkable growth. Yet, a significant impediment to ascertaining phylogenetic relationships has been the paucity of both ancestral and transitional forms. Speculation is gradually being replaced with resolution.
Therizinosaurs are now considered to be unequivocal descendants not only of theropods but of the coelurosaurian clade and maniraptoran subclade with a sister relationship with Oviraptorosauria (see below). Thus, therizinosaurs are Saurischian, Theropodal, Tetanurian, Coelurosaurian, Maniraptoran dinosaurs and members of the family Therizinosauridea.
Therizinosaurs also share the tree with more highly derived Avialae (birds) and possess avian-associated characters such as a pneumatic-skeleton (hollow light-weight bones that facilitate a high rate of respiration, and later, powered flight), a pygostyle (shortened tail with fused vertebrae), feathers (for thermo-regulation, sexual dimorphism and brooding) and an avian-trending brain (for enhanced sight, sound and mechano-reception).
UNCLEAR FAMILY INTERRELATIONSHIPS
Interrelationships between specific therizinosaur taxa remains less clear. Until recently, the fossil record was restricted to Asia. With discoveries in North America such as Nothronychus mckinleyi (the first undisputed North American therizinosaurid from the Upper Cretaceous of New Mexico) and Falcarius utahensis (the third therizinosaur discovered in America, the most morphologically primitive therizinosaur yet discovered and a sister taxon to the clade of Therizinosauroidea from the Lower Cretaceous Cedar Mountain Formation of east-central Utah), we can begin to ask questions about origination, geographic and stratigraphic range, and even potential faunal mixing between immigrant and endemic clades.
|One of many parsimonious phylogenies proposed for therizinosaurs|
50% Majority-rule consensus tree
Modified from Pu et al, 2013.
QUESTIONS WITHOUT ANSWERS
Did basalmost Falcarius utahensis originate in North America and certain populations expand into Asia? With recent finds in China such as Eshanosaurus deguchiianus from the Early Jurassic, the presence of derived coelurosaurian lineages including therizinosaurians is being pushed back earlier. Did therizinosaurs differentiate from coelurosaurian ancestors before the breakup of Pangaea into Eurasia and North America and/or did they migrate across the tectonic Beringia Land Bridge that was established in the Early Cretaceous between northeast Asia and northwest North America? Was there more than one dispersal event? At various times, Asian endemic therizinosaurids show faunal similarities with North American forms. Did endemic forms mix with immigrant forms? Do North American therizinosaur taxa exhibit an Asian affinity or vice versa?
VEGETARIANS WITHIN A FAMILY OF CARNIVORES
Based on therizinosaur's osteological anatomy and soft tissue reconstructions, taking into account the habitats in which they likely thrived, and using animal analogues such as the sloth, certain dietary assumptions have been made about therizinosaurs and the Coelurosaurian clade in which they belong - once thought to have been obligate carnivores. In a little over a decade, doubt has been shed on that notion, raising the possibility or even likelihood that "dietary diversification was more commonplace among 'predatory' dinosaurs than previously appreciated" (Zanno, 2009).
In fact, therizinosaurs are the most widely regarded candidate for herbivory among theropods. Dietary plasticity and facultative (capable of rather than restricted to) herbivory (omnivory) is thought to have afforded the group the potential to invade and exploit ecospaces early in evolution for survival.
|With a similar body shape and large claws on their front feet, Nothronychus graffami is shown as a bipedal browser analogous to the Giant Ground Sloth of the Ice Age.|
Artist Victor Leshyk
Wayne Ranney and I met Merle bright and early at the Visitor Center in Big Water for a tour of the Moon. Crossing the dry streambed of Wahweap Creek, we travelled on a planar surface of Tropic mudstone, occasionally bouncing around on the hummocky, eroded terrain. The easiest places to build roads on the Colorado Plateau, although the most difficult places to maintain them, are on the Mancos-Tropic Shale. The soft rock weathers readily, forming broad, flat expanses and easy routes to get from here to there.
Our first destination was the very spot of Merle's once-in-a-lifetime discovery. After a surprisingly short drive from the Visitor Center, we left our vehicle and began to ascend a large, loose mound of gray mudstone and claystone to a noticeably beveled off area.
Standing at the site of the former dig, let's let Merle tell the story. Although he's recounted the details many times in well over a decade since the find, it's clear that his enthusiasm hasn't diminished one iota. I'm filming, while Wayne is interjecting commentary.
|Merle and Wayne debate the mysterious circumstances of the terrestrial therizinosaur's burial at sea.|
BURIAL AT SEA - BUT BY WHAT MEANS?
One important question that has plagued paleontologists is how the dinosaur came to be buried in marine mud 60 miles out to sea with the nearest shore confirmed geologically near present-day Cedar City? To date, no definitive answer exists, although theories include "bloat and float" - having died on or near land and washed out to sea buoyed by decomposing body-gases followed by burial- and "lost at sea" - a less plausible scenario of having been caught by a flood or storm, floated out to sea alive, perhaps attacked by predators, and eventually buried nearly fully articulated. The skeleton of N. graffami was located in a supine position, belly-up, implying that it settled to its final resting place in the Tropic mud while buoyed by gases in a typical death pose.
I've seen the same "bloat and float" taphonymous (mode of fossilization) entity in New Jersey where terrestrial duck-billed dinosaur remains (the state fossil) have been found buried in glauconitic sands of the flooded Late Cretaceous continental shelf on Jersey farmland.
As the Tropic continues to erode, a few small osseous remnants of N. graffami have weathered to the surface since its excavation over a decade ago. It's that fact, among many others, that keeps Merle coming back to The Moon. There's always something new to be found on the ever-changing landscape. Come back the next day, and a new discovery will be awaiting you. What appears to be a static landscape is entirely the opposite!
HEADING DOWN DRY WAHWEAP CREEK
|Yours truly with Merle and his shirt|
• First Definitive Therizinosaurid From North America by James I. Kirkland and Douglas G. Wolfe, 2001.
• Fossils from the Tropic Shale by Merle H. Grafam, 2000. Personal copy from the author.
• Geological Evolution of the Colorado Plateau of Eastern Utah and Western Colorado by Robert Fillmore, 2011.
• The Geology of the Grand Staircase in Southern Utah by the Geological Society of America, 2002.
• The Pectoral Girdle and Forelimb of the Primitive Therizinosaiuriod Falcarius Utahensis by Lindsay A. Zanno, 2006.
• Therizinosaur – Mystery of the Sickle-Claw Dinosaur by David D. Gillette, Ph.D., Plateau, Museum of Northern Arizona, 2007.
• Vertebrate Paleontology by Michael J. Benton, 2005.