Reconstruction of Typothorax coccinarum based on skeletons from eastern New Mexico
Image by Matt Celeskey

These skeletons allowed us to revise previously published reconstructions of this aetosaur. In particular, we now have good evidence of the total number of rows of armor, the arrangement of the scutes on the belly, new insight into the appendages (particularly the shoulder girdle and feet), a more domelike carapace based on extremely wide and gently curved paramedian scutes, and the very first aetosaur reconstruction to sport cloacal spikes.

This has been a fun project to be involved with, and I thank Dr. Andy Heckert (a former coworker at the NMMNH, now at Appalachian State University) for inviting me to assist in reconstructing this armor-plated Triassic reptile.

Update 5/21: At the request of commenter dmaas, I’m uploading a detail of the head of the reconstruction. Clicking on the thumbnail will bring it up at more than twice the size of the original painting.

• Reference: Heckert, A. B., Lucas, S. G., Rinehart, L. F., Celeskey, M. D., Spielmann, J. A., and Hunt, A. P. (2010) Articulated skeletons of the aetosaur Typothorax coccinarum Cope (Archosauria: Stagonolepididae) from the Upper Triassic Bull Canyon Formation (Revueltian: Early-Mid Norian), eastern New Mexico, USA. Journal of Vertebrate Paleontology, vol. 30, no. 3, pp. 619–642.
• Elsewhere on the web:
  • Society of Vertebrate Paleontology press release (also at Appalachian State University)
  • Discovery News: Dino-Era Reptiles: Part Cow, Armadillo, Crocodile (lots of quotes from myself, the other authors & others)
  • Critical review at Chinleana: I am so disappointed in the Typothorax description in the new issue of JVP

No time to treat this with more than a passing mention, but a letter in today’s Nature presents a new, Middle Triassic silesaurid from Tanzania named Asilisaurus kongwe (“ancient ancestor lizard”). This adds another continent to the known range of the silesaurids, a group of plant-eating proto-dinosaurs previously found in Europe, South America and North America. It also extends the age of this group back 10 million years, which makes it not only the oldest-known silesaurid, but the oldest known reptile on the bird side of the bird-crocodile split.

Neat stuff, and particularly interesting after looking at Eucoelophysis…

For more information:

• The Nature article: Ecologically distinct dinosaurian sister group shows early diversification of Ornithodira
• Figures from the article
• Graphics and information from the research team
• Coverage at Chinleana

Less-well publicized but just as interesting is the PLoS Biology’s “primer” article on Studying Function and Behavior in the Fossil Record by Michael Benton. It provides an overview of three lines of evidence that can lead to testable hypotheses about ancient behavior: empirical evidence, comparison with modern animals, and biomechanical modeling. For each of these approaches, examples are pulled from paleontological (mostly dinosaur) research over the past decade. There is, I suspect, a lot more that could be said on the topic of inferring behavior from fossils, but this brief is a useful companion piece to the Sanajeh paper and I’d certainly recommend taking a look at it.

In other news of fossilized behavior, my posting will be a bit petrified this week as I push forward on another project—more Coelophysis posts are in the works, however, and I should have one or more to put up next week…

Outside of the inside of its mouth, however, little is known about the appearance of Ptychodus. The researchers behind the new study propose that it might have been something like a modern nurse shark (Ginglystoma cirratum), which also finds most of its food in the along the bottoms of tropical seas.

I was happy to read this because several years ago I had a similar thought and worked up a sketch of a nurse shark-like Ptychodus, complete with big pectoral fins, little eyes and speculative whisker-like barbels for sensing prey beneath the sediments.

The enigmatic durophage Ptychodus as a nurse shark analogue.

About a year after I put together that sketch, my friend Mary Sundstrom expanded on the sketch to create a dynamic, shell-crunching reconstruction for a web project at the day job:

Painting of Ptychodus based on the previous sketch, by Mary Sundstrom, 2005.

• References: Shimada, Kenshu, Everhart, Michael J., Decker, Ramo and Decker, Pamela D. 2010. A new skeletal remain of the durophagous shark, Ptychodus mortoni, from the Upper Cretaceous of North America: an indication of gigantic body size. Cretaceous Research, vol. 31, pp. 249–254. doi:10.1016/j.cretres.2009.11.005
• Shimada, Kenshu, Rigsby, Cynthia K. and Kim, Sun H., 2009. Partial skull of Late Cretaceous durophagous shark, Ptychodus occidentalis (Elasmobranchii: Ptychodontidae), from Nebraska, U.S.A. Journal of Vertebrate Paleontology, vol. 29 (2), pp. 336–349. doi: 10.1671/039.029.0226

Quick note: New readers might want to look at previous installments of the Paleobiology of Coelophysis (Parts 1 & 2) series before diving into this post.

In order to collect data from other specimens of Coelophysis bauri, some members of the research team (notably Larry Rinehart and Andy Heckert) visited several other museums to study the Whitaker quarry blocks in their care. After a trip to the Royal Tyrrell Museum of Paleontology in Alberta, Larry shared observations and photographs from one specimen that seemed a bit out of place compared to the other Coelophysis we had seen.

TMP 84-63-33, highlighted against the rest of the Tyrrell Museum block
Original photograph courtesy of Larry Rinehart

Specimen TMP 84-63-33 is the most easily seen skeleton on the Tyrrell Museum block. Although the front and back ends of the animal are missing, most of the middle is well-preserved, particularly the two hindlimbs. At first glance, it looks much like any other Coelophysis from the quarry.

Some details, however, led us to wonder about this. For instance, while Coelophysis has five sacral vertebrae connected to its hip, TMP 84-63-33 appears to have only four. Several features on the proximal end of the femur (that is, the part of the thigh bone that connects to the hip bones) also seemed notably different from what we saw on Coelophysis, and these features suggested an alternative identification.

Focus on the Femora
or, A Discomfiting Object Inserted in the Acetabulum

The proximal end of the left femur of TMP 84-63-33
articulated within the acetabulum (hip socket)
Photograph courtesy of Larry Rinehart

Some of the odd femoral features include:

1. A well-defined groove on top of the head of the femur.
2. The head of the femur is offset but appears to be completely rounded off. There is no sign of the hook-like prong seen in Coelophysis and other theropods.
3. A distinct crest-like trochanter (or bump of bone) on the front (anterior) side of the femur toward the outside (lateral) edge, which has not been reported in Coelophysis.

Of these features, the first and (particularly) the third were a close match for features seen on NMMNH P-22298, the holotype specimen of Eucoelophysis baldwini. Eucoelophysis (“True Coelophysis” or “True Hollow Form”) also lacks a hook-like prong on the head of its femur, although it is quite different in overall shape from the rounded, offset femur head of TMP 84-63-33. This might be a real difference, or it might be due to the badly weathered condition of the Eucoelophysis holotype. In the end, based primarily on the similarities noted here, we assigned TMP 84-63-33 to Eucoelophysis sp.

Comparison of the femoral heads of Eucoelophysis baldwini, TMP 84-63-33, and Coelophysis bauri. Click for larger view. This figure is not from Rinehart et al. 2009, but drawn up as a visual aid to this post.

That’s where we left things for publication—a genus-level assignment based on some key characters that TMP 84-63-33 and NMMNH P-22298 have in common. Things get more interesting when you add a little background and some other fossils, so I’ll take the opportunity to explore some of those tangents here.

Eucoelophysis vs. Coelophysis: An Example of Interspecific Digression
or, Reflections on a Hollow Form of Truth

When Eucoelophysis was first described, it was considered to be a theropod dinosaur closely related to Coelophysis (Sullivan and Lucas 1999). However, a pair of later studies (Ezcurra 2006, Nesbitt et al. 2007) concluded that Eucoelophysis was a “non-dinosaurian dinosauriform”—not only was it not particularly close to Coelophysis, it lacked the requisite anatomical features needed to be included in the Dinosauria proper. Its closest companion in these dinosaur hinterlands appeared to be Silesaurus, a beaked, herbivorous reptile known from excellent skull and skeletal material from the Late Triassic of Poland (Dzik 2003).

The idea that Eucoelophysis might be a Silesaurus-style dinosauriform has received support from new and newly-recognized discoveries of other Silesaurus-like fossils in Late Triassic rocks from Arizona and New Mexico (Parker et al. 2006, Irmis et al. 2007). These include some blocky, angled femur heads whose overall shape is similar to that of both Silesaurus and the shape preserved in the Eucoelophysis holotype. And at least one of these femur heads (PEFO 34357) appears to have a Eucoelophysis-style anterolateral trochanter (=the dorsolateral trochanter noted by Nesbitt et al. 2007).

TMP 84-63-33, on the other hand, looks a lot more like Coelophysis than Silesaurus in many observable parts of its anatomy, especially the bones of its pelvis and feet. I wouldn’t say that our assignment of TMP 84-63-33 to Eucoelophysis reaffirms close relationship between Eucoelophysis and Coelophysis. But if this identification holds, then it doesn’t appear to do much for a Eucoelophysis-Silesaurus connection, either.

One last osteological nubbin of interest: the lesser trochanter (also referred to as the cranial or anterior trochanter) is a prong of bone that, in the animals we’re discussing, sits just below the head of the femur on the front-facing side. This trochanter is slender and crest-like on both Eucoelophysis and TMP 84-63-33. On observed and reported specimens of Coelophysis bauri, the lesser trochanter is thick, blocky, and connected to a well-developed shelf of bone that wraps around the outside of the femur. In other coelophysoids, both forms of lesser trochanter have been found within the same species—such as in the African species Coelophysis rhodesiensis, where the two different shapes may represent a difference between males and females (Raath 1990).

Comparison of the proximal left femurs of specimens mentioned in this post, in anterior (front) view, resized to similar widths. Inset shows silhouettes to scale. Redrawn from various sources. Click for larger view.

When a couple more femur heads are added to the previous figure, I begin to see a gradation of forms between the block-headed, slender-trochantered dinosauriform femora through to the hooked femur heads and robust trochanters of Coelophysis bauri. Note that I do not suggest that this shows any sort of evolutionary sequence. Instead, the continuum of shapes and features is probably due to a mix of phylogenetic differences, sexual dimorphism, age- and size-related changes, individual variation, and preservation quality.

The trick is to figure out what sort of meaningful divisions might be found within this femoral spectrum. In Rinehart et al. 2009, we made one division based on similarities between the femora TMP 84-63-33 and Eucoelophysis. I suspect that the wealth of fossils from the Whitaker quarry will have more to reveal on the topic, both from close evaluation of femur variation in the large Coelophysis population, and from comparing those variations with data gleaned from other parts of the skeleton.

Next time: Data Gleaned from other Parts of the Skeleton
With far less talk of femur variation, and perhaps even some actual paleobiology!

Previously:

Part I: Introduction
Part II: Other Critters in the Quarry

• Main Reference: Rinehart, Larry F., Lucas, Spencer G., Heckert, Andrew B., Spielmann, Justin A. and Celeskey, Matthew D., 2009. The Paleobiology of Coelophysis bauri (Cope) from the Upper Triassic (Apachean) Whitaker quarry, New Mexico, with detailed analysis of a single quarry block. New Mexico Museum of Natural History and Science Bulletin 45, 260pp. Abstract [Rich text file]
• Additional References: Dzik, Jerzy, 2003. A beaked herbivorous archosaur with dinosaur affinities from the Early Late Triassic of Poland. Journal of Vertebrate Paleontology, vol. 23 (3), pp. 556–574.
• Ezcurra, Martín D., 2006. A review of the systematic position of the dinosauriform archosaur Eucoelophysis baldwini Sullivan & Lucas, 1999 from the Upper Triassic of New Mexico, USA. Geodiversitas, vol. 28 (4), pp. 649–684. [PDF]
• Irmis, Randall B., Nesbitt, Sterling J., Padian, Kevin, Smith, Nathan D., Turner, Alan H., Woody, Daniel and Downs, Alex, 2007. A Late Triassic dinosauromorph assemblage from New Mexico and the rise of dinosaurs. Science, vol. 317, no. , pp. 358–361. doi: 10.1126/science.1143325
• Nesbitt, Sterling J., Irmis, Randall B. and Parker, William G., 2007. A critical re-evaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology, vol. 5 (2), pp. 209–243. doi: 10.1017/S1477201907002040
• Parker, William G., Irmis, Randall B. and Nesbitt, Sterling J., 2006. Review of the Late Triassic dinosaur record from Petrified Forest National Park, Arizona. Museum of Northern Arizona Bulletin 62, pp. 160–161.
• Raath, Michael A. 1990. Morphological variation in small theropods and its meaning in systematics: evidence from Syntarsus rhodesiensis in Dinosaur Systematics: Perspectives and Approaches, Kenneth Carpenter and Philip J. Currie, eds. Cambridge University Press. pp. 91–105.
• Sullivan, Robert M. and Lucas, Spencer G., 1999. Eucoelophysis baldwini, a new theropod dinosaur from the Upper Triassic of New Mexico, and the status of the original types of Coelophysis. Journal of Vertebrate Paleontology, vol. 19 (1), pp. 81–90.

• Love in the Time of Chasmosaurs: a daily dose of Mesozoic megafauna. Highly recommended. This week’s pick: An interview with pterosaur researcher/artist Mark Witton.
• Pterosaur.net Blog: posts from Mark Witton and the rest of the team at Pterosaur.net. Pick: John Conway’s discussion of the Pot-bellied Pteranodon.
• Jim Robins Zoo.Art: art of a paleontological nature. Pick: The multi-view rendering of Caudipteryx zoui.
• Theropod Database: Mickey Mortimer’s in-depth examinations of theropod anatomy and phylogenetics. Pick: A handsome little picture of the first Asian alvarezsaur fossil ever cataloged.
• Morbid Anatomy shows off an admission certificate for an 1869 course on Practical Anatomy. Instructor: Joseph Leidy, who named the first American dinosaurs.
• Laelaps describes how Arctodus simus, the Pleistocene “short-faced bear,” got a makeover.
• More fossil pigments: the plumage color patterns of Anchornis. In 3D!

• New basal alvarezsaur: Haplocheirus sollers.
• What’s an alvarezsaur? Dave Hone’s got it covered.
• Why is a basal one noteworthy? Jonah Choiniere, of Haplocheirus-description team, puts it in context and highlights some of its unique features.
• Fossil pigment 1: Belemnite ink
• Fossil pigment 2: Chestnut-striped coelurosaurs
• Happy Family: Proboscidea
• Medical Texts: A long, painful decline (courtesy bioephemera)
• The Wonderful World: Compellingly awful

• 
  Bistahieversor sealeyi
  Image by Mary Sundstrom and myself, for the
  New Mexico Museum of Natural History and Science.

• New Dinosaur: Bistahieversor sealeyi
• Pronounced: bis-tah-he-ee-VER-sor SEE-lee-eye
• Name means: Sealey’s Bisti destroyer (Paul Sealey discovered the fossils in the Bisti/De-na-zin Wilderness Area)
• Named by: Carr and Williamson 2010
• Relations: Tyrannosauroid, a group that includes Tyrannosaurus rex and its not-too-distant relatives
• Location: Northwest New Mexico, United States of America
• Age: Late Cretaceous (Campanian), ~73,000,000 years old
• Length: ~9 meters (29 feet)
• Info: Two fairly complete skeletons of a new type of New Mexican dinosaur are helping to fill in gaps in the evolution and distribution of tyrannosauroids, the group of dinosaurs that includes T. rex and its fairly close relatives.
• Tyrannosauroid fossils have been known from southwestern North America for over 100 years, but mostly in bits and pieces—isolated bones and teeth or, at best, incomplete skeletons of uncertain identity (see Carr & Williamson 2000 for a good overview). Traditionally, researchers have assigned these fossils to well-known tyrannosauroid genera like Albertosaurus or Daspletosaurus, whose more complete remains were originally found further north in Wyoming, Montana, and Alberta.
• Thanks to recent discoveries, the southwest now has a tyrannosaur all its own. Bistahieversor is the new name given to a complete skull and (mostly unprepared) skeleton from the Bisti badlands of northwestern New Mexico. Its skull and jaws display a healthy list of detailed anatomical characters that distinguish it from all other tyrannosauroids, including a complex joint between the nasal and frontal bones on top of its skull, and a unique hole above its eye. Other tyrannosaur remains from northwest New Mexico, including the partial skull and skeleton of a juvenile, appear to be specimens of Bistahieversor as well.
• Images:
  NMMNH P-27469, holotype skull and jaw of Bistahieversor sealeyi
  Photograph by David Baccadutre, New Mexico Museum of Natural History and Science.

• 
  NMMNH P-25049, incomplete skull and skeleton of juvenile Bistahieversor sealeyi
  Both these specimens are on display in the New Mexico’s Seacoast hall of the NMMNHS.

• Main Reference: Carr, Thomas D. and Williamson, Thomas E., 2010 Bistahieversor sealeyi, gen. et sp. nov., a new tyrannosauroid from New Mexico and the origin of deep snouts in Tyrannosauroidea. Journal of Vertebrate Paleontology, vol. 30 (1), pp. 1–16. doi: 10.1080/02724630903413032
• Additional Reference: Carr, Thomas D. and Williamson, Thomas E., 2000 A review of Tyrannosauridae (Dinosauria, Coelurosauria) from New Mexico. New Mexico Museum of Natural History and Science Bulletin 17, pp. 113–145.
• Elsewhere on the web:
  • Dinochick Blogs: Williamson and Carr introduce the destroyer – Bistahieversor sealeyi: Part 1 Part 2
  • Theropoda: Il Distruttore di Bisti (Carr & Williamson 2010) [english]
  • SVP & Paleo News: PRESS RELEASE – New Species of Tyrannosaur Discovered in Southwestern U.S.
  • National Geographic News: New “Destroyer” Dinosaur Found, was T. rex Relative
  • Brett Booth reconstructed Bistahieversor for Draw a Dinosaur Day

Although fossils of the Triassic theropod Coelophysis bauri are by far the most numerous vertebrate remains preserved in blocks from the Whitaker quarry, several other animals are known from the site as well. Many have just been uncovered or described within the past ten years, and in the course of preparing the NMMNH block (and examining other blocks for comparison), several new fossils have come to light. A brief, annotated list of other fauna known from the quarry appears below:

Invertebrates: As mentioned last time, ostracods (Darwinula sp.) and conchostracans (Shipingia) were found in a sandy layer below the bones, remnants of temporary ponding at the site prior to the Coelophysis burial.

Synorichthys chased by Chinlea, sculpted by Gary Staab

Fish: Schaeffer (1967) reported paleonisciform and coelacanth fish in association with Coelophysis at Ghost Ranch. Both were found in the NMMNH block above the invertebrate layer and just below the lowest tetrapod bones: scaly little redfieldiid paleonisciforms, tentatively assigned to Synorichthys, and bits of fin, scale, and skull from the large coelacanth Chinlea sorenseni. As these fish would have been too large to thrive in the type of ephemeral pond indicated by the invertebrates, we posited that floodwaters washed them in to the site from a larger body of water.

Whitakersaurus bermani: This diminutive sphenodontian (my tiny-tuatara-based restoration at the left) is known from pieces of the upper and lower jaw found within 2 centimeters of the edge of the NMMNH block. The largest piece of the holotype, an incomplete right dentary preserving nineteen tooth-positions, is about 5 millimeters long (Heckert et al. 2008).

Drepanosaurs: Harris & Downs (2002) reported the first drepanosaur material from the quarry—a well-preserved (but isolated) shoulder girdle from the block at the Ruth Hall Museum of Paleontology. In a new review of the drepanosaurs, Renesto et al. (2010) assign that shoulder girdle to the genus Drepanosaurus, and describe a partially articulated but generically indeterminate foot on the edge of the NMMNH block (pictured below).

NMMNH P-57651, the foot of a small drepanosaur, preserved portion roughly 5cm long

Vancleavea campi: By far the most complete specimens of this armor-coated reptile are two beautifully articulated skeletons from the Ruth Hall Museum block, recently described by Nesbitt et al. (2009). Remains of a partial, disarticulated skeleton are associated with some characteristic Vancleavea armor in a mostly unprepared fossil removed from the NMMNH block.

Vancleavea sculpture by Phil Bircheff at the Ruth Hall Museum of Paleontology.

Phytosaurs: An ~80cm long phytosaur skull from the Whitaker/Coelophysis quarry (the holotype of Redondasaurus bermani Hunt and Lucas 1993) is housed at the Carnegie Museum. The skull and lower jaws of a juvenile phytosaur were found in the NMMNH block, currently exposed in left lateral/palatal view on a partially prepared jacket removed from the main block (photo below).

NMMNH P-44920, juvenile phytosaur (Redondasaurus?) skull and jaws, left lateral/palatal view. Photo courtesy of Larry Rinehart.

Postosuchus kirkpatricki: The Carnegie Museum of Natural History and the Ruth Hall Museum of Paleontology both have specimens of this rauisuchian on Whitaker/Coelophysis quarry blocks. These were partially figured in Long and Murry (1995, figs 145–146).

Effigia sculpture by Phil Bircheff
at the Ruth Hall Museum of Paleontology.

Effigia okeefeae: The type specimens of this bipedal suchian were recently discovered in jackets pulled from the quarry during the early excavations by the American Museum (Nesbitt & Norell 2006, Nesbitt 2007). A scapula and coracoid found in the NMMNH block might belong to this animal.

Hesperosuchus agilis: One of the best-preserved specimens of this early crocodylomorph is an articulated skull and partial skeleton now at the Carnegie Museum (Clark et al. 2000). Only a few armor scutes are known from the NMMNH block.

To the best of my knowledge, this little bestiary lists pretty much all the non-Coelophysis animals identified from Ghost Ranch Whitaker quarry fossils. All of them, that is, with one interesting exception, noticed while reviewing specimens on other Coelophysis blocks. Its story will make up the bulk of the next post.

Next time: Truly, Coelophysis?
or, The Mysterious Canadian

Previously:
Introduction

• Main Reference: Rinehart, Larry F., Lucas, Spencer G., Heckert, Andrew B., Spielmann, Justin A. and Celeskey, Matthew D., 2009. The Paleobiology of Coelophysis bauri (Cope) from the Upper Triassic (Apachean) Whitaker quarry, New Mexico, with detailed analysis of a single quarry block. New Mexico Museum of Natural History and Science Bulletin 45, 260pp. Abstract [Rich text file]
• Additional References: Clark, James M., Sues, Hans-Dieter and Berman, David S., 2000. A new specimen of Hesperosuchus agilis from the Upper Triasic of New Mexico and the interrelationships of basal crocodylomorph archosaurs. Journal of Vertebrate Paleontology, vol. 20 (4), pp. 683–704. doi: 10.1671/0272-4634(2000)020[0683:ANSOHA]2.0.CO;2
• Harris, Jerald D. and Downs, Alex, 2002. A drepanosaurid pectoral girdle from the Ghost Ranch (Whitaker) Coelophysis quarry (Chinle Group, Rock Point Formation, Rhaetian), New Mexico. Journal of Vertebrate Paleontology, vol. 22 (1), pp. 70–75. doi: 10.1671/0272-4634(2002)022[0070:ADPGFT]2.0.CO;2 [PDF]
• Heckert, Andrew B., Lucas, Spencer G., Rinehart, Larry F. and Hunt, Adrian P., 2008. A new genus and species of sphenodontian from the Ghost Ranch Coelophysis quarry (Upper Triassic: Apachean), Rock Point Formation, New Mexico, USA. Palaeontology, vol. 51, pp. 827–845. doi: 10.1111/j.1475-4983.2008.00786.x [PDF]
• Hunt, Adrian P. and Lucas, Spencer G., 1993. A new phytosaur (Reptilia: Archosauria) genus from the uppermost Triassic of the western United States and its biochronological significance. New Mexico Museum of Natural History and Science Bulletin 3, pp. 193–196.
• Long, Robert A. and Murry, Phillip A., 1995. Late Triassic (Carnian and Norian) tetrapods from the southwestern United States. New Mexico Museum of Natural History and Science Bulletin 4, 254pp. [link to PDF]
• Nesbitt, Sterling J. 2007. The anatomy of Effigia okeeffeae (Archosauria, Suchia), theropod convergence, and the distribution of related taxa. Bulletin of the American Museum of Natural History, vol. 302, pp. 1–84.
• Nesbitt, Sterling J. and Norell, Mark A., 2006. Extreme convergence in the body plans of an early suchian (Archosauria) and ornithomimid dinosaurs (Theropoda). Proceedings of the Royal Society B, vol. 273, no. 1590, pp. 1045–1048. doi: 10.1098/rspb.2005.3426
• Nesbitt, Sterling J., Stocker, Michelle R., Small, Brian J. and Downs, Alex, 2009. The osteology and relationships of Vancleavea campi (Reptilia: Archosauriformes). Zoological Journal of the Linnean Society, vol. 157 (4), pp. 814–864. doi: 0.1111/j.1096-3642.2009.00530.x
• Renesto, Silvio, Spielmann, Justin A., Lucas, Spencer G. and Tarditi Spagnoli, Georgio, 2010. The taxonomy and paleobiology of the Late Triassic (Carnian-Norian: Adamanian-Apachean) drepanosaurs (Diapsida: Archosauromorpha: Drepanosauromorpha). New Mexico Museum of Natural History and Science Bulletin 46, 81pp.
• Schaeffer, Bobb, 1967. Late Triassic fishes from the western United States. Bulletin of the American Museum of Natural History, vol. 135 (6), pp. 285–342.