Dakotaraptor

*Dakotaraptor steini*

Luis Rey Artwork

***Dakotaraptor*** (meaning "Dakota thief") is a potentially chimaeric genus of dromaeosaurs known from partial skeletons unearthed in South Dakota, USA. Its fossils have been found in the Maastrichtian-age Hell Creek Formation, dated to the very end of the Cretaceous (69-66 MYA) period, making it one of the last surviving dromaeosaurids, is said to have lived in North America during the latter part of the Cretaceous period. However the publications conducted that this dromaeosaurid genus, is very likely being a Chimera (meaning made up from different fossils and animals)

*Dakotaraptor* was about five to six metres long, which makes it one of the largest dromaeosaurids known. It had long arms. One of the lower arm bones shows quill knobs, proving that *Dakotaraptor* was feathered. It also had long rear legs with a very large sickle claw on the second toe; this claw could be used to kill relatively large plant-eating dinosaurs. *Dakotaraptor* lived in the same time and area as *Tyrannosaurus rex*.

## Discovery

In 2005, paleontologist Robert DePalma in Harding County, South Dakota discovered the skeleton of a large dromaeosaurid. Subsequently, the same site produced additional dromaeosaurid remains, as well as abundant other fossils in 2010 described by DePalma in his master's thesis. In 2015, the type species *Dakotaraptor steini* was named and described by Robert A. DePalma, David A. Burnham, Larry Dean Martin, Peter Lars Larson and Robert Thomas Bakker. The generic name combines a reference to South Dakota and the Dakota people with a Latin raptor, "plunderer". The specific name honours paleontologist Walter W. Stein. *Dakotaraptor* was one of eighteen dinosaur taxa from 2015 to be described in open access or free-to-read journals.

The holotype, PBMNH.P.10.113.T, was found in a sandstone layer of the upper Hell Creek Formation, dating from the late Maastrichtian. It consists of a partial skeleton, lacking the skull, of an adult individual. It contains a piece of a back vertebra, ten tail vertebrae, a furcula, both upper arm bones, both ulnae, both radii, the first and second right metacarpals, three claws of the left hand, a right thighbone, both shinbones, a left astragalus bone, a left calcaneum, the left second, third and fourth metatarsal, the right fourth metatarsal, and the second and third claw of the right foot. Apart from the remains of the holotype, in the site bones were discovered that also belonged to *Dakotaraptor* but which represented a more gracile morph. These included the specimens PBMNH.P.10.115.T: a right shinbone; PBMNH.P.10.118.T: a connected left astragalus and calcaneum; and KUVP 152429: a furcula. Additionally four isolated teeth were referred, specimens PBMNH.P.10.119.T, PBMNH.P.10.121.T, PBMNH.P.10.122.T, and PBMNH.P.10.124.T. These fossils are part of the collection of The Palm Beach Museum of Natural History. Other referred fossils are NCSM 13170, a furcula, and KUVP 156045, an isolated tooth.

*Dakotaraptor* may be a different species or "size-morph" of *Acheroraptor*, due to them being so much larger than adult *Acheroraptor* specimens. Phylogenetic tests show that the two fossils of *Dakotaraptor* and *Acheroraptor* are not related closely.

## Description

### Size

*Dakotaraptor* is exceptionally large for a dromaeosaurid; it has an estimated adult length of 5.5 m (18 ft). This approaches the size of the largest known dromaeosaurid, *Utahraptor*. *Dakotaraptor* however, does not have the proportions and adaptations of *Utahraptor*, but more closely resembles smaller dromaeosaurids like *Deinonychus*.

### Distinguishing traits

Apart from the large size, the description of 2015 indicated some additional distinguishing traits. On the fourth foot claw, the boss that serves as an attachment for the tendon of the flexor muscle is reduced in size. The "blood groove" on the outer side of the fourth claw of the foot, towards the tip is fully enclosed over half of its length, forming a bony tubular structure. The second and third claws of the foot have sharp keels at their undersides. The second foot claw, the "sickle claw", equals 29% of the thighbone length. On the shinbone, the crista fibularis, the crest that contacts the calfbone, is long and lightly built with a height that does not exceed 9% of the crest length. The upper edge of this crest ends in a hook. On the second metacarpal, of the two condyles that contact the finger, the inner one is almost as large as the outer one. The outer side of the second metacarpal has but a shallow groove for the ligament that connects it to the third metacarpal. When the arm is seen in a flat position, of the second metacarpal the edge between the wrist joint and the upper shaft is straight in top view. The teeth have fifteen to twenty denticles per five millimetres on the rear edges and twenty to twenty-seven denticles on the front edges.

### Skeleton

#### Vertebral column

The vertebrae of the back are highly pneumatised, filled with trabecular bone that shows many air spaces. On the middle tail vertebrae the front joint processes, the prezygapophyses, are extremely elongated with an estimated intact length of seventy centimetres, spanning about ten vertebrae. This stiffens the tail.

#### Forelimb

Like virtually all other theropods, *Dakotaraptor* possessed a furcula, or “wishbone”, as part of the shoulder girdle. Theropod wishbones are quite varied and often different from the strongly U-shaped furculae most modern birds possess. *Dakotaraptor* had a furcula that was U- to V-shaped, similar to many other dromaeosaurids such as *Velociraptor*, and even the large spinosaurid theropod *Suchomimus*. In 2015, a study by Victoria Megan Arbour e.a. claimed that the presumed *Dakotaraptor furculae* in fact represented a part of a turtle armour, the entoplastron of *Axestemys splendida*, a member of the Trionychidae.

The wing of *Dakotaraptor* was given much attention in the describing article. Here, “wing” is used as an anatomical descriptive term not related to its functionality, since *Dakotaraptor* was flightless. This is similar to the term “wing” for the same appendages in ostriches, emus, and other flightless birds. It is meant to express that arm is equipped with long feathers. Many of the wing bones were discovered (humerus, radius, ulna, two of the three metacarpal wrist bones, and parts of the finger digits), so the wing is very complete. The humerus, the upper arm bone, is relatively long and slender. It is somewhat bent to the inside. The most notable anatomical feature is the row of very prominent bumps along a ridge on the lower edge of the ulna, one of the forearm bones. These are called ulnar papillae, or quill knobs. In birds and some other theropod dinosaurs, these bumps were spots for reinforced attachment of the remiges, or wing feathers. When quill knobs are present, this is considered a strong indication that the animal had long remiges on the wings. Since quill knobs are rare in the fossil record, paleontologists mainly relied on phylogenetic bracketing to determine if a species was likely to have had wing feathers - if a relative on a "higher" branch of the evolutionary tree had the feathers, and one on a "lower" branch down had them also, then a species in the middle position likely did as well. *Dakotaraptor’s* quill knobs show that the animal unequivocally had prominent wing feathers, making it the largest dromaeosaurid with confirmed plumage of that type. The quill knobs of *Dakotaraptor* have a diameter of about eight to ten millimetres, which shows that these feathers were rather large. It was estimated that a complete series might include fifteen of these papillae ulnares. The ulna is thirty-six centimetres long and the other lower arm bone, the radius, measures thirty-two centimetres. The hand bones show that their joints allowed for little mobility. The wingspan of *Dakotaraptor* was estimated at 120 centimetres, not taking into account possible primary remiges longer than the hand.

The second metacarpal of the metacarpus of the hand, the bone that primary remiges attach to, also had a flat bony shelf as its dorsal surface. The shelf made a perfect spot for the primary feathers to lay across in their life-attachment.

#### Hindlimb

Overall, the hindlimb is built lightly and with long elements, contrary to the robust, stocky hind limbs of *Utahraptor*. *Dakotaraptor* more closely resembles the agile, springy smaller dromaeosaurids and would have been well-suited at running and pursuit predation. The length of the thighbone is 558 millimetres. It is relatively shorter and more lightly built than that of *Utahraptor*. To the contrary the shinbone is rather elongated. The holotype shinbone is with a length of 678 millimetres the longest dromaeosaurid tibia known. It is 22% longer than the thighbone, indicating a good running capability. The shinbone's cnemial crest has a sharp corner pointing to the front. Its fibular crest ends in a hook-shaped process pointing to above, a condition that is unique in the entire Theropoda. The astragalus and calcaneum, the upper ankle bones, are fused just as in *Bambiraptor*. The top of the calcaneum has but a small contact facet for the calfbone, indicating that this fibula must have had a very narrow lower end.

The metatarsus has an estimated length of thirty-two centimetres, which makes it rather long relative to the remainder of the hindlimb.

The foot claws of *Dakotaraptor* include a typical dromaeosaurid raptorial second claw, or "sickle claw", which was used for killing or holding down prey. It is large and robust with a diameter of sixteen centimetres and a length of twenty-four centimetres measured along the outer curve. This equals 29% of the length of the thighbone compared to 23% with *Deinonychus*. The claw is transversely flattened and has a droplet-shaped cross-section. The flexor tubercle, a large bump near the base, served as an attachment site for flexor muscles - the larger it was, the greater the slashing strength. *Dakotaraptor* has a flexor tubercle that is larger relative to overall claw size than it is in other discovered dromaeosaurids, potentially giving it the strongest slashing strength of any known member of this group. The flexor tubercle on the third claw of the foot is almost non-existent, very reduced in size compared to other dromaeosaurids, suggesting a more minimized use of that claw. As these are the bony cores of the claws, they would have been covered in a keratinous sheath that extended the "nail" and ended in a sharp tip. The third claw too is keeled but is much smaller with a tip to joint length of seven centimetres and a curve length of nine centimetres. The groove on its outer side towards the tip ends in a bone tunnel, a rare condition.

## Classification

Dakotaraptor was first recovered as a dromaeosaurid, being sister to *Dromaeosaurus*. Then, it was was found to be in clade with *Utahraptor* and *Achillobator*. It was suggested to separate, being a 4th instance of size increase in dromaeosaurs:

Hartman et al. (2019) recover Dakotaraptor as a unenlagiid, which would mark one of the northernmost occurrences if true, disregarding the Hell Creek unenlagiid:

Currie and Evans (2019) recovered a eudromaeosaur, noting that the holotype may not be a single individual:

The naming of *Dineobellator* found the same as above, but noted its chimeric status, and casted doubt upon this placement, finding it to be too basal.

## Paleobiology

The sickle claw was used to pin down prey, of which it had a greater capability due to a stronger flexor tubercle. The third foot claw was smaller and seems to have had no function. Robust and gracile morphs are present, with both represented by adult specimens. Individual variation or pathologies may have induced this, but sexual dimorphism is the strongest explanation. It is unknown which morph represents what sex. Large pennaceous feathers were present despite being flightless. Instead, they may have shielded eggs, displayed, intimidated or kept balance during prey restraint. It may have descended from an ancestor that had these, and none of these functions do not necessitate wing feathers.

## Paleoecology

*Dakotaraptor* lived in the Hell Creek Formation, which was populated by a wide variety of fauna; mosasaurs, hadrosaurs, ceratopsians, dromaeosaurs, tyrannosaurs, oviraptorosaurs, ornithomimosaurs, alvarezsaurs, fish, pterosaurs, small ornithischians, mammals, insects, birds, crocodilians, turtles and others. It lived up until the K-PG boundary. It may have been a pack hunter, in which case it was capable of taking larger prey. It may have been a pursuit predator.

## Appearance in other media

### *Jurassic Park*

* *Dakotaraptor* appears in Jurassic World: Alive as an epic creature obtainable with 150 DNA.

## References

1. ↑ https://www.researchgate.net/publication/308979112\_The\_furculae\_of\_the\_dromaeosaurid\_dinosaur\_Dakotaraptor\_steini\_are\_trionychid\_turtle\_entoplastra
2. ↑ https://peerj.com/preprints/1570/
3. ↑ https://twitter.com/LancianIdolatry/status/1392150985288544258
4. ↑ 4.0 4.1 4.2 4.3 https://en.wikipedia.org/wiki/Dakotaraptor
5. ↑ Cite error: Invalid `<ref>` tag; no text was provided for refs named `0:`
6. ↑ https://en.wikipedia.org/wiki/Paleobiota\_of\_the\_Hell\_Creek\_Formation