Raptors &#;small- to medium-sized feathered dinosaurs equipped with single, long, curving hind claws on their hind feet&#;were among the most fearsome predators of the Mesozoic Era . On the following slides, you'll find pictures and detailed profiles of over 25 raptors, ranging from A (Achillobator) to Z (Zhenyuanlong).
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01
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Achillobator
Matt Martyniuk/Wikimedia Commons/CC BY 2.5
Achillobator was named after the hero of Greek myth (its name is actually a combination of Greek and Mongolian, "Achilles warrior"). Not much is known about this central Asian raptor, whose oddly shaped hips set it slightly apart from others of its kind.
02
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Adasaurus
Karkemish/Wikimedia Commons/CC BY 3.0
Name
Adasaurus (Greek for "Ada lizard"); pronounced AY-dah-SORE-us
Habitat
Woodlands of central Asia
Historical Period
Late Cretaceous (75-65 million years ago)
Size and Weight
About 5 feet long and 50-75 pounds
Diet
Meat
Distinguishing Characteristics
Tall skull; short claws on hind feet; probable feathers
Adasaurus (named after an evil spirit from Mongolian mythology) is one of the more obscure raptors to be unearthed in central Asia, much less well-known than its close contemporary Velociraptor. To judge by its limited fossil remains, Adasaurus had an unusually tall skull for a raptor (which doesn't necessarily mean that it was smarter than others of its kind), and the single, oversized claws on each of its hind feet were positively puny compared to those of Deinonychus or Achillobator. About the size of a large turkey, Adasaurus preyed on the smaller dinosaurs and other animals of late Cretaceous central Asia.
03
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Atrociraptor
FunkMonk/Wikimedia Commons/CC BY-SA 3.0
Name
Atrociraptor (Greek for "cruel thief"); pronounced ah-TROSS-ih-rap-tore
Habitat
Woodlands of North America
Historical Period
Late Cretaceous (70 million years ago)
Size and Weight
About three feet long and 20 pounds
Diet
Meat
Distinguishing Characteristics
Small size; short snout with backward-curving teeth
It's amazing how a mere name can color our view of a long-extinct dinosaur. For all intents and purposes, Atrociraptor was very similar to Bambiraptor&#;both were puny, albeit dangerous, raptors with sharp teeth and ripping hind claws&#;but judging by their names you'd probably want to pet the latter and run away from the former. Whatever the case, Atrociraptor was certainly deadly for its size, as demonstrated by its backward-curving teeth&#;the only conceivable function of which would have been to tear off jagged chunks of meat (and prevent live prey from escaping).
04
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Austroraptor
ESV/Wikimedia Commons/CC BY-SA 3.0
Name
Austroraptor (Greek for "southern thief"); pronounced AW-stroh-rap-tore
Habitat
Woodlands of South America
Historical Period
Late Cretaceous (70 million years ago)
Size and Weight
About 16 feet long and 500 pounds
Diet
Meat
Distinguishing Characteristics
Large size; narrow snout; short arms
As with all types of dinosaurs, paleontologists are unearthing new raptors all the time. One of the latest to join the flock is Austroraptor, which was "diagnosed" in based on a skeleton dug up in Argentina (hence the "austro," meaning "south," in its name). To date, Austroraptor is the largest raptor yet discovered in South America, measuring a full 16 feet from head to tail and probably weighing in the neighborhood of 500 pounds&#;proportions that would have given its North American cousin, Deinonychus, a run for its money, but would have made it no match for the nearly one-ton Utahraptor that lived tens of millions of years earlier.
05
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Balaur
Jaime Headden/Wikimedia Commons/CC BY 4.0
Name
Balaur (Romanian for "dragon"); pronounced BAH-lore
Habitat
Woodlands of eastern Europe
Historical Period
Late Cretaceous (70-65 million years ago)
Size and Weight
About three feet long and 25 pounds
Diet
Meat
Distinguishing Characteristics&#;
Muscular build; double claws on hind feet
Its full name, Balaur bondoc, makes it sound like the supervillain from a James Bond movie, but if anything this dinosaur was even more interesting: an island-dwelling, late Cretaceous raptor with a host of weird anatomical features. First, unlike other raptors, Balaur sported two oversized, curved claws on each of its hind feet, rather than one; and second, this predator cut an unusually squat, muscular profile, very unlike its lithe, speedy cousins like Velociraptor and Deinonychus. In fact, Balaur possessed such a low center of gravity that it may have been capable of tackling much larger dinosaurs (especially if it hunted in packs).
Why did Balaur occupy a position so far outside the raptor norm? Well, it seems that this dinosaur was restricted to an island environment, which can produce some strange evolutionary results&#;witness the "dwarf" titanosaur Magyarosaurus, which only weighed a ton or so, and the comparably shrimpy duck-billed dinosaur Telmatosaurus. Clearly, Balaur's anatomical traits were an adaptation to the limited flora and fauna of its island habitat, and this dinosaur evolved in its strange direction thanks to millions of years of isolation.
06
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Bambiraptor
Ballista/Wikimedia Commons/CC BY-SA 3.0
Its warm, fuzzy name invokes images of gentle, furry forest creatures, but the fact is Bambiraptor was as vicious as a pit bull&#;and its fossil has yielded valuable clues about the evolutionary relationship between dinosaurs and birds.
07
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Buitreraptor
FunkMonk/Wikimedia Commons/CC BY-SA 3.0
Name
Buitreraptor (combination Spanish/Greek for "vulture thief"); pronounced BWEE-tray-rap-tore
Habitat
Plains of South America
Historical Period
Late Cretaceous (90 million years ago)
Size and Weight
About four feet long and 25 pounds
Diet
Small animals
Distinguishing Characteristics
Long, narrow snout; smooth teeth; probably feathers
Only the third raptor ever to be discovered in South America, Buiteraptor was on the small side, and the lack of serrations on its teeth indicate that it fed on much smaller animals, rather than ripping into the flesh of its fellow dinosaurs. As with other raptors, paleontologists have reconstructed Buitreraptor as covered with feathers, connoting its close evolutionary relationship to modern birds. (By the way, this dinosaur's odd name stems from the fact that it was unearthed, in , in the La Buitrera area of Patagonia&#;and since Buitrera is Spanish for "vulture," the moniker seemed appropriate!)
08
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Changyuraptor
Emily Willoughby/Wikimedia Commons/CC BY-SA 4.0
Name
Changyuraptor (Greek for "Changyu thief"); pronounced CHANG-yoo-rap-tore
Habitat
Woodlands of Asia
Historical Period
Early Cretaceous (125 million years ago)
Size and Weight
About three feet long and 10 pounds
Diet
Small animals
Distinguishing Characteristics
Four wings; long feathers
As is often the case when a brand-new dinosaur is discovered, there has been a lot of speculation about Changyuraptor, not all of which is warranted. Specifically, the media have been touting the hypothesis that this raptor&#;a relative of the much smaller, and also four-winged, Microraptor&#;was capable of powered flight. While it's true that the tail feathers of Changyuraptor were a foot long, and may have served some navigational function, it may also be the case that they were strictly ornamental and only evolved as a sexually selected characteristic.
Another clue that Changyuraptor's aerial bona-fides are being overstated is that this raptor was fairly large, about three feet from head to tail, which would render it much less airworthy than Microraptor (after all, modern turkeys have feathers, too!). At the very least, though, Changyuraptor should shed new light on the process by which the feathered dinosaurs of the early Cretaceous period learned to fly.
09
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Cryptovolans
Stephen A. Czerkas/Prehistoric Wiki
Name
Cryptovolans (Greek for "hidden flyer"); pronounced CRIP-toe-VO-lanz
Habitat
Woodlands of Asia
Historical Period
Early Cretaceous (130-120 million years ago)
Size and Weight
About three feet long and 5-10 pounds
Diet
Meat
Distinguishing Characteristics
Long tail; feathers on front and hind limbs
True to the "crypto" in its name, Cryptovolans has occasioned its share of disputes among paleontologists, who aren't quite sure how to classify this early Cretaceous feathered dinosaur. Some experts believe that Cryptovolans is actually a "junior synonym" of the better known Microraptor, a four-winged raptor that made a big splash in paleontology circles a couple of years ago, while others maintain that it deserves its own genus, mainly because of its longer-than-Microraptor tail. Adding to the mystery, one scientist insists that Cryptovolans not only merits its own genus but was more evolved toward the bird end of the dinosaur-bird spectrum than even Archaeopteryx&#;and thus should be considered a prehistoric bird rather than a feathered dinosaur!
10
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Dakotaraptor
Emily Willoughby/Wikimedia Commons/CC BY-SA 4.0
The late Cretaceous Dakotaraptor is only the second raptor ever to be discovered in the Hell Creek formation; the type fossil of this dinosaur bears unmistakable "quill knobs" on its front limbs, meaning it almost certainly possessed winged forearms. See an in-depth profile of Dakotaraptor
11
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Deinonychus
Emily Willoughby/Wikimedia Commons/CC BY-SA 4.0
The "Velociraptors" in Jurassic Park were actually modeled after the Deinonychus, a fierce, man-sized raptor distinguished by the huge claws on its back feet and its grasping hands&#;and that wasn't nearly as smart as it has been depicted in the movies.
Read More
Mesozoic Feathered Dinosaurs: Tiny Insectivores to Predators
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Dromaeosauroides
FunkMonk/Wikimedia Commons/CC BY-SA 3.0
Name
Dromaeosauroides (Greek for "like Dromaeosaurus"); pronounced DROE-may-oh-SORE-oy-deez
Habitat
Woodlands of northern Europe
Historical Period
Early Cretaceous (140 million years ago)
Size and Weight
About 10 feet long and 200 pounds
Diet
Meat
Distinguishing Characteristics
Large head; curved claws on hind feet; probably feathers
The name Dromaeosauroides is quite a mouthful and has probably rendered this meat-eater less well-known to the public than it rightfully should be. Not only is this the only dinosaur ever to be discovered in Denmark (a couple of fossilized teeth recovered from the Baltic Sea island of Bornholm), but it's also one of the earliest identified raptors, dating to the early Cretaceous period, 140 million years ago. As you may have guessed, the 200-pound Dromaeosauroides was named in reference to the better-known Dromaeosaurus ("running lizard"), which was much smaller and lived tens of millions of years later.
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Dromaeosaurus
Yinan Chen/Wikimedia Commons/Public Domain
Name
Dromaeosaurus (Greek for "running lizard"); pronounced DRO-may-oh-SORE-us
Habitat
Plains of North America
Historical Period
Late Cretaceous (75 million years ago)
Size and Weight
About six feet long and 25 pounds
Diet
Meat
Distinguishing Characteristics
Small size; powerful jaws and teeth; probably feathers
Dromaeosaurus is the eponymous genus of dromaeosaurs, the smallish, speedy, bipedal, probably feather-covered dinosaurs better known to the general public as raptors. Still, this dinosaur differed from more famous raptors like Velociraptor in some important respects: the skull, jaws, and teeth of Dromaeosaurus were relatively robust, for instance, a very tyrannosaur-like trait for such a small animal. Despite its standing among paleontologists, Dromaeosaurus (Greek for "running lizard") isn't very well represented in the fossil record; all we know of this raptor amounts to a few scattered bones unearthed in Canada in the early 20th century, mostly under the supervision of the buccaneering fossil-hunter Barnum Brown.
Analysis of its fossils reveals that Dromaeosaurus was a more formidable dinosaur than Velociraptor: its bite may have been three times as powerful (in terms of pounds per square inch) and it preferred to disembowel its prey with its toothy snout, rather than the single, oversized claws on each of its hind feet. The recent discovery of a closely related raptor, Dakotaraptor, lends added weight to this "teeth first" theory; like Dromaeosaurus, this dinosaur's hind claws were relatively inflexible, and wouldn't have been of much use in close-quarters combat.
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Graciliraptor
FunkMonk/Wikimedia Commons/CC BY-SA 3.0
Name
Graciliraptor (Greek for "graceful thief"); pronounced grah-SILL-ih-rap-tore
Habitat
Woodlands of Asia
Historical Period
Early Cretaceous (125 million years ago)
Size and Weight
About three feet long and a few pounds
Diet
Meat
Distinguishing Characteristics
Small size; feathers; large, single claws on hind feet
Discovered in China's famous Liaoning fossil beds&#;the final resting place of a huge variety of small, feathered dinosaurs from the early Cretaceous period&#;Graciliraptor is one of the earliest and smallest raptors yet identified, measuring only about three feet long and weighing a couple of pounds soaking wet. In fact, paleontologists speculate that Graciliraptor occupied a position close to the "last common ancestor" of raptors, troodontids (feathered dinosaurs closely related to Troodon), and the first true birds of the Mesozoic Era, which probably evolved around this time. Though it's unclear whether it was similarly equipped, Graciliraptor also seems to have been closely related to the famous, four-winged Microraptor, which arrived on the scene a few million years later.
15
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Linheraptor
Smokeybjb/Wikimedia Commons/CC BY-SA 3.0
Name
Linheraptor (Greek for "Linhe hunter"); pronounced LIN-heh-rap-tore
Habitat
Plains of central Asia
Historical Period
Late Cretaceous (85-75 million years ago)
Size and Weight
About six feet long and 25 pounds
Diet
Meat
Distinguishing Characteristics
Long legs and tail; bipedal posture; probably feathers
The amazingly well-preserved fossil of Linheraptor was discovered during an expedition to the Linhe region of Mongolia in , and two years of preparation have revealed a sleek, probably feathered raptor that prowled the plains and woodlands of late Cretaceous central Asia in search of food. Comparisons to another Mongolian dromaeosaur, Velociraptor, are inevitable, but one of the authors of the paper announcing Linheraptor says it's best compared to the equally obscure Tsaagan (yet another, similar raptor, Mahakala, has been found in these same fossil beds).
16
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Luanchuanraptor
FunkMonk/Wikimedia Commons/CC BY-SA 3.0
Name
Luanchuanraptor (Greek for "Luanchuan thief"); pronounced loo-WAN-chwan-rap-tore
Habitat
Woodlands of Asia
Historical Period
Late Cretaceous (70 million years ago)
Size and Weight
About 3-4 feet long and 5-10 pounds
Diet
Meat
Distinguishing Characteristics
Small size; bipedal posture; probably feathers
As obscure as it is, the tiny, probably feathered Luanchuanraptor occupies an important place in the dinosaur record books: it was the first Asian raptor to be discovered in eastern rather than northeastern China (most dromaeosaurs from this part of the world, like Velociraptor, lived further west, in modern-day Mongolia). Other than that, Luanchuanraptor seems to have been a fairly typical "dino-bird" for its time and place, possibly hunting in packs to overwhelm the bigger dinosaurs that counted as its prey. Like other feathered dinosaurs, Luanchuanraptor occupied an intermediate branch on the tree of bird evolution.
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Microraptor
CoreyFord/Getty Images
Microraptor fits uneasily into the raptor family tree. This tiny dinosaur had wings on both its front and back limbs, but it probably wasn't capable of powered flight: rather, paleontologists picture it gliding (like a flying squirrel) from tree to tree.
18
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Neuquenraptor
PaleoGeekSquared/Wikimedia Commons/CC BY-SA 4.0
Name
Neuquenraptor (Greek for "Neuquen thief"); pronounced NOY-kwen-rap-tore
Habitat
Woodlands of South America
Historical Period
Late Cretaceous (90 million years ago)
Size and Weight
About six feet long and 50 pounds
Diet
Meat
Distinguishing Characteristics
Large size; bipedal posture; feathers
If only the paleontologists who discovered it had gotten their act together, Neuquenraptor might stand today as the first identified raptor from South America. Unfortunately, this feathered dinosaur's thunder wound up being stolen by Unenlagia, which was discovered in Argentina a few months later but, thanks to a canny bit of analytical work, named first. Today, the weight of the evidence is that Neuquenraptor was actually a species (or specimen) of Unenlagia, characterized by its unusually large size and its propensity for flapping its arms (but not actually flying).
19
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Nuthetes
Mark Witton/Wikimedia Commons/CC BY 2.0
Name
Nuthetes (Greek for "monitor"); pronounced noo-THEH-teez
Habitat
Woodlands of western Europe
Historical Period
Early Cretaceous (145-140 million years ago)
If you want to learn more, please visit our website Animatronic Birds.
Size
About six feet long and 100 pounds
Diet
Meat
Distinguishing Characteristics
Small size; bipedal posture; possibly feathers
As problematic genera go, Nuthetes has proven a tough nut to crack. It took over a decade after its discovery (in the mid-19th century) for this dinosaur to be classified as a theropod. The question was exactly what kind of theropod: was Nuthetes a close relative of Proceratosaurus, an ancient forebear of Tyrannosaurus Rex, or a Velociraptor-like dromaeosaur? The problem with this last category (which has only reluctantly been accepted by paleontologists) is that Nuthetes dates to the early Cretaceous period, over 140 million years ago, which would make it the earliest raptor in the fossil record. The jury, pending further fossil discoveries, is still out.
20
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Pamparaptor
Eloy Manzanero/Wikimedia Commons/CC BY-SA 3.0
Name
Pamparaptor (Greek for "Pampas thief"); pronounced PAM-pah-rap-tore
Habitat
Plains of South America
Historical Period
Late Cretaceous (90-85 million years ago)
Size and Weight
About two feet long and a few pounds
Diet
Meat
Distinguishing Characteristics
Small size; bipedal posture; feathers
Argentina's Neuquen province, in Patagonia, has proven to be a rich source of dinosaur fossils dating to the late Cretaceous period. Originally diagnosed as a juvenile of another South American raptor, Neuquenraptor, Pamparaptor was elevated to genus status on the basis of a well-preserved hind foot (sporting the single, curved, elevated claw characteristic of all raptors). As dromaeosaurs go, the feathered Pamparaptor was on the tiny end of the scale, only measuring about two feet from head to tail and weighing a few pounds soaking wet.
21
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Pyroraptor
Conty/Wikimedia Commons/CC BY 3.0
Name
Pyroraptor (Greek for "fire thief"); pronounced PIE-roe-rap-tore
Habitat
Plains of western Europe
Historical Period
Late Cretaceous (70 million years ago)
Size and Weight
About 8 feet long and 100-150 pounds
Diet
Meat
Distinguishing Characteristics
Large, sickle-shaped claws on feet; probably feathers
As you may have guessed from the last part of its name, Pyroraptor belongs to the same family of theropods as Velociraptor and Microraptor: the raptors, which were distinguished by their speed, viciousness, single-clawed hind feet and (in most cases) feathers. Pyroraptor ("fire thief") didn't earn its name because it actually stole fire, or even breathed fire, in addition to the usual array of raptor weapons: the more prosaic explanation is that the only known fossil of this dinosaur was discovered in , in southern France, after a forest fire.
22
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Rahonavis
Bernard Sandler via FunkMonk/Wikimedia Commons/CC BY 2.0
Name
Rahonavis (Greek for "cloud bird"); pronounced RAH-hoe-NAY-viss
Habitat
Woodlands of Madagascar
Historical Period
Late Cretaceous (75 million years ago)
Size and Weight
About one foot long and one pound
Diet
Probably insects
Distinguishing Characteristics
Small size; feathers; single curved claw on each foot
Rahonavis is one of those creatures that triggers enduring feuds among paleontologists. When it was first discovered (an incomplete skeleton unearthed in Madagascar in ), researchers assumed it was a type of bird, but further study showed certain traits common to dromaeosaurs (better known to the general public as raptors). Like such undisputed raptors as Velociraptor and Deinonychus, Rahonavis had a single huge claw on each hind foot, as well as other raptor-like features.
What is the current thinking about Rahonavis? Most scientists agree that raptors counted among the early ancestors of birds, meaning that Rahonavis might be a "missing link" between these two families. The trouble is, it wouldn't be the only such missing link; dinosaurs may have made the evolutionary transition to flight multiple times, and only one of these lineages went on to spawn modern birds.
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Saurornitholestes
Emily Willoughby/Wikimedia Commons/CC BY 3.0
Name
Saurornitholestes (Greek for "lizard-bird thief"); pronounced sore-OR-nith-oh-LESS-tease
Habitat
Plains of North America
Historical Period
Late Cretaceous (75 million years ago)
Size and Weight
About five feet long and 30 pounds
Diet
Meat
Distinguishing Characteristics
Sharp teeth; large claws on feet; probably feathers
If only Saurornitholestes had been given a manageable name, it might be as popular as its more famous cousin, Velociraptor. Both these dinosaurs were excellent examples of late Cretaceous dromaeosaurs (better known to the general public as raptors), with their slight, agile builds, sharp teeth, relatively large brains, big-clawed hind feet, and (probably) feathers. Tantalizingly, paleontologists have discovered a wing bone of the huge pterosaur Quetzalcoatlus with a Saurornitholestes tooth embedded inside it. Since it's unlikely that a 30-pound raptor could have taken down a 200-pound pterosaur all by itself, this can be taken as evidence that either a) Saurornitholestes hunted in packs or b) more likely, a lucky Saurornitholestes happened upon an already-dead Quetzalcoatlus and took a bite out of the carcass.
24
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Shanag
FunkMonk/Wikimedia Commons/CC BY-SA 3.0
Name
Shanag (after the Buddhist "Cham Dance"); pronounced SHAH-nag
Habitat
Plains of central Asia
Historical Period
Early Cretaceous (130 million years ago)
Size and Weight
About three feet long and 10-15 pounds
Diet
Meat
Distinguishing Characteristics
Small size; feathers; bipedal posture
During the early Cretaceous period, 130 million years ago, it was difficult to distinguish one small, feathered dinosaur from the next&#;the boundaries separating raptors from "troodontids" from plain-vanilla, bird-like theropods were still in flux. As far as paleontologists can tell, Shanag was an early raptor closely related to the contemporary, four-winged Microraptor, but also shared some characteristics with the line of feathered dinosaurs that went on to spawn the late Cretaceous Troodon. Since all we know of Shanag consists of a partial jaw, further fossil discoveries should help determine its exact place on the dinosaur evolutionary tree.
25
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Unenlagia
Sergey Krasovskiy
Name
Unenlagia (Mapuche for "half-bird"); pronounced OO-nen-LAH-gee-ah
Habitat
Plains of South America
Historical Period
Late Cretaceous (90 million years ago)
Size and Weight
About six feet long and 50 pounds
Diet
Meat
Distinguishing Characteristics
Large size; flapping arms; probably feathers
Although it was unmistakably a dromaeosaur (what ordinary folks call a raptor), Unenlagia has raised some puzzling issues for evolutionary biologists. This feathered dinosaur was distinguished by its very limber shoulder girdle, which gave its arms a broader range of motion than comparable raptors&#;so it's only a short step to imagining that Unenlagia actually flapped its feathered arms, which might well have resembled wings.
The puzzlement pertains to the fact that Unenlagia was clearly too big, about six feet long and 50 pounds, to take to the air (by way of comparison, flying pterosaurs with comparable wingspans weighed much less). This raises the prickly question: could Unenlagia have spawned a (now-extinct) line of flying, feathered descendants similar to modern birds, or was it a flightless relative of the first, genuine birds that preceded it by tens of millions of years?
26
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Utahraptor
Emily Willoughby/Wikimedia Commons/CC BY-SA 3.0
Utahraptor was by far the biggest raptor that ever lived, which raises a serious conundrum: this dinosaur lived tens of millions of years before its more famous descendants (like Deinonychus and Velociraptor), during the middle Cretaceous period!
27
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Variraptor
Abujoy/Wikimedia Commons/CC BY-SA 3.0
Name
Variraptor (Greek for "Var River thief"); pronounced VAH-ree-rap-tore
Habitat
Woodlands of western Europe
Historical Period
Late Cretaceous (85-65 million years ago)
Size and Weight
About seven feet long and 100-200 pounds
Diet
Meat
Distinguishing Characteristics
Long arms; long, lightly built skull with numerous teeth
Despite its impressive name, the French Variraptor occupies a place on the second tier of the raptor family, since not everyone accepts that this dinosaur's scattered fossil remains add up to a convincing genus (and it's not even clear exactly when this dromaeosaur lived). As it has been reconstructed, Variraptor was slightly smaller than the North American Deinonychus, with a proportionately lighter head and longer arms. There's also some speculation that (unlike most raptors) Variraptor may have been a scavenger rather than an active hunter, though the case for that would certainly be bolstered by more convincing fossil remains.
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Velociraptor
LEONELLO CALVETTI/Getty Images
Velociraptor wasn't a particularly big dinosaur, though it did have a mean disposition. This feathered raptor was about the size of a large chicken, and there's no evidence that it was anywhere near as smart as it has been depicted in the movies.
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Zhenyuanlong
Emily Willoughby/Wikimedia Commons/CC BY-SA 4.0
Name
Zhenyuanlong (Chinese for "Zhenyuan's dragon"); pronounced zhen-yan-LONG
Habitat
Woodlands of Asia
Historical Period
Early Cretaceous (125 million years ago)
Size and Weight
About five feet long and 20 pounds
Diet
Meat
Distinguishing Characteristics
Relatively large size; short arms; primitive feathers
There's something about Chinese bonebeds that lend themselves to spectacularly preserved fossil specimens. The latest example is Zhenyuanlong, announced to the world in and represented by a nearly complete skeleton (lacking only the hind part of the tail) complete with the fossil imprint of wispy feathers. Zhenyuanlong was fairly large for an early Cretaceous raptor (about five feet long, which places it in the same weight class as the much later Velociraptor), but it was hobbled by a relatively short arm-to-body ratio and it was almost certainly unable to fly. The paleontologist who discovered it (no doubt seeking press coverage) has called it the "fluffy feathered poodle from hell."
This article is about the dinosaur. For the doom metal group, see Deinonychus (band)
Not to be confused with Deinocheirus or Deinosuchus
Deinonychus ([1] dy-NON-ih-kəs; from Ancient Greek δεινός (deinós) 'terrible', and &#;νυξ (ónux), genitive &#;νυχος (ónukhos) 'claw') is a genus of dromaeosaurid theropod dinosaur with one described species, Deinonychus antirrhopus. This species, which could grow up to 3.4 meters (11 ft) long, lived during the early Cretaceous Period, about 115&#;108 million years ago (from the mid-Aptian to early Albian stages). Fossils have been recovered from the U.S. states of Montana, Utah, Wyoming, and Oklahoma, in rocks of the Cloverly Formation and Antlers Formation, though teeth that may belong to Deinonychus have been found much farther east in Maryland.
Paleontologist John Ostrom's study of Deinonychus in the late s revolutionized the way scientists thought about dinosaurs, leading to the "dinosaur renaissance" and igniting the debate on whether dinosaurs were warm-blooded or cold-blooded. Before this, the popular conception of dinosaurs had been one of plodding, reptilian giants. Ostrom noted the small body, sleek, horizontal posture, ratite-like spine, and especially the enlarged raptorial claws on the feet, which suggested an active, agile predator.[2]
"Terrible claw" refers to the unusually large, sickle-shaped talon on the second toe of each hind foot. The fossil YPM preserves a large, strongly curved ungual. In life, archosaurs have a horny sheath over this bone, which extends the length. Ostrom looked at crocodile and bird claws and reconstructed the claw for YPM as over 120 millimetres (4.7 in) long.[2] The species name antirrhopus means "counter balance", which refers to Ostrom's idea about the function of the tail. As in other dromaeosaurids, the tail vertebrae have a series of ossified tendons and super-elongated bone processes. These features seemed to make the tail into a stiff counterbalance, but a fossil of the very closely related Velociraptor mongoliensis (IGM 100/986) has an articulated tail skeleton that is curved laterally in a long S-shape. This suggests that, in life, the tail could bend to the sides with a high degree of flexibility.[3] In both the Cloverly and Antlers formations, Deinonychus remains have been found closely associated with those of the ornithopod Tenontosaurus. Teeth discovered associated with Tenontosaurus specimens imply they were hunted, or at least scavenged upon, by Deinonychus.
Discovery and naming
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Fossilized remains of Deinonychus have been recovered from the Cloverly Formation of Montana and Wyoming[2] and in the roughly contemporary Antlers Formation of Oklahoma,[4] in North America. The Cloverly formation has been dated to the late Aptian through early Albian stages of the early Cretaceous, about 115 to 108 Ma.[5][6] Additionally, teeth found in the Arundel Clay Facies (mid-Aptian), of the Potomac Formation on the Atlantic Coastal Plain of Maryland may be assigned to the genus.[7]
The first remains were uncovered in in southern Montana near the town of Billings. The team leader, paleontologist Barnum Brown, was primarily concerned with excavating and preparing the remains of the ornithopod dinosaur Tenontosaurus, but in his field report from the dig site to the American Museum of Natural History, he reported the discovery of a small carnivorous dinosaur close to a Tenontosaurus skeleton, "but encased in lime difficult to prepare."[8] He informally called the animal "Daptosaurus agilis" and made preparations for describing it and having the skeleton, specimen AMNH , put on display, but never finished this work.[9] Brown brought back from the Cloverly Formation the skeleton of a smaller theropod with seemingly oversized teeth that he informally named "Megadontosaurus". John Ostrom, reviewing this material decades later, realized that the teeth came from Deinonychus, but the skeleton came from a completely different animal. He named this skeleton Microvenator.[9]
Cast of the holotype foot YPM from two anglesA little more than thirty years later, in August , paleontologist John Ostrom led an expedition from Yale's Peabody Museum of Natural History which discovered more skeletal material near Bridger. Expeditions during the following two summers uncovered more than 1,000 bones, among which were at least three individuals. Since the association between the various recovered bones was weak, making the exact number of individual animals represented impossible to determine properly, the type specimen (YPM ) of Deinonychus was restricted to the complete left foot and partial right foot that definitely belonged to the same individual.[10] The remaining specimens were catalogued in fifty separate entries at Yale's Peabody Museum although they could have been from as few as three individuals.[10]
Later study by Ostrom and Grant E. Meyer analyzed their own material as well as Brown's "Daptosaurus" in detail and found them to be the same species. Ostrom first published his findings in February , giving all the referred remains the new name of Deinonychus antirrhopus. The specific name "antirrhopus", from Greek &#;ντίρροπος, means "counterbalancing" and refers to the likely purpose of a stiffened tail.[11] In July , Ostrom published a very extensive monograph on Deinonychus.[10]
Though a myriad of bones was available by , many important ones were missing or hard to interpret. There were few postorbital skull elements, no femurs, no sacrum, no furcula or sternum, missing vertebrae, and (Ostrom thought) only a tiny fragment of a coracoid. Ostrom's skeletal reconstruction of Deinonychus included a very unusual pelvic bone&#;a pubis that was trapezoidal and flat, unlike that of other theropods, but which was the same length as the ischium and which was found right next to it.[10]
Further findings
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In , Ostrom published another monograph on the shoulder of Deinonychus in which he realized that the pubis that he had described was actually a coracoid&#;a shoulder element.[12] In that same year, another specimen of Deinonychus, MCZ , was discovered and excavated in Montana by Steven Orzack during a Harvard University expedition headed by Farish Jenkins. This discovery added several new elements: well preserved femora, pubes, a sacrum, and better ilia, as well as elements of the pes and metatarsus. Ostrom described this specimen and revised his skeletal restoration of Deinonychus. This time it showed the very long pubis, and Ostrom began to suspect that they may have even been a little retroverted like those of birds.[13]
Reconstructed skeleton of specimen AMNH , with outdated hand posture The similarity of the forelimbs (left) with those of Archaeopteryx (right) led John Ostrom to revive the link between dinosaurs and birdsA skeleton of Deinonychus, including bones from the original (and most complete) AMNH specimen, can be seen on display at the American Museum of Natural History,[14] with another specimen (MCZ ) on display at the Museum of Comparative Zoology at Harvard University. The American Museum and Harvard specimens are from a different locality than the Yale specimens. Even these two skeletal mounts are lacking elements, including the sterna, sternal ribs, furcula, and gastralia.[15]
Even after all Ostrom's work, several small blocks of lime-encased material remained unprepared in storage at the American Museum. These consisted mostly of isolated bones and bone fragments, including the original matrix, or surrounding rock in which the specimens were initially buried. An examination of these unprepared blocks by Gerald Grellet-Tinner and Peter Makovicky in revealed an interesting, overlooked feature. Several long, thin bones identified on the blocks as ossified tendons (structures that helped stiffen the tail of Deinonychus) turned out to actually represent gastralia (abdominal ribs). More significantly, a large number of previously unnoticed fossilized eggshells were discovered in the rock matrix that had surrounded the original Deinonychus specimen.[16]
In a subsequent, more detailed report, on the eggshells, Grellet-Tinner and Makovicky concluded that the egg almost certainly belonged to Deinonychus, representing the first dromaeosaurid egg to be identified.[8] Moreover, the external surface of one eggshell was found in close contact with the gastralia suggesting that Deinonychus might have brooded its eggs. This implies that Deinonychus used body heat transfer as a mechanism for egg incubation, and indicates an endothermy similar to modern birds.[17] Further study by Gregory Erickson and colleagues finds that this individual was 13 or 14 years old at death and its growth had plateaued. Unlike other theropods in their study of specimens found associated with eggs or nests, it had finished growing at the time of its death.[18]
Implications
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Ostrom's description of Deinonychus in has been described as the most important single discovery of dinosaur paleontology in the mid-20th century.[19] The discovery of this clearly active, agile predator did much to change the scientific (and popular) conception of dinosaurs and opened the door to speculation that some dinosaurs may have been warm-blooded. This development has been termed the dinosaur renaissance. Several years later, Ostrom noted similarities between the forefeet of Deinonychus and that of birds, an observation which led him to revive the hypothesis that birds are descended from dinosaurs.[20] Forty years later, this idea is almost universally accepted.
Because of its extremely bird-like anatomy and close relationship to other dromaeosaurids, paleontologists hypothesize that Deinonychus was probably covered in feathers.[21][22][23] Clear fossil evidence of modern avian-style feathers exists for several related dromaeosaurids, including Velociraptor and Microraptor, though no direct evidence is yet known for Deinonychus itself.[24][25] When conducting studies of such areas as the range of motion in the forelimbs, paleontologists like Phil Senter have taken the likely presence of wing feathers (as present in all known dromaeosaurs with skin impressions) into consideration.[26]
Description
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Life restorationBased on the few fully mature specimens,[27] Paul estimated that Deinonychus could reach 3.3&#;3.4 meters (10 ft 10 in &#; 11 ft 2 in) in length, with a skull length of 410 millimeters (16 in), a hip height of 0.87 meters (2.9 ft) and a body mass of 60&#;73 kg (132&#;161 lb).[28][29] Campione and his colleagues proposed a higher mass estimate of 100 kg (220 lb) based on femur and humerus circumference.[30] The skull was equipped with powerful jaws lined with around seventy curved, blade-like teeth. Studies of the skull have progressed a great deal over the decades. Ostrom reconstructed the partial, imperfectly preserved skulls that he had as triangular, broad, and fairly similar to Allosaurus. Additional Deinonychus skull material and closely related species found with good three-dimensional preservation[31] show that the palate was more vaulted than Ostrom thought, making the snout far narrower, while the jugals flared broadly, giving greater stereoscopic vision. The skull of Deinonychus was different from that of Velociraptor, however, in that it had a more robust skull roof, like that of Dromaeosaurus, and did not have the depressed nasals of Velociraptor.[32] Both the skull and the lower jaw had fenestrae (skull openings) which reduced the weight of the skull. In Deinonychus, the antorbital fenestra, a skull opening between the eye and nostril, was particularly large.[31]
Size compared with a humanDeinonychus possessed large "hands" (manus) with three claws on each forelimb. The first digit was shortest and the second was longest. Each hind foot bore a sickle-shaped claw on the second digit, which was probably used during predation.[10]
No skin impressions have ever been found in association with fossils of Deinonychus. Nonetheless, the evidence suggests that the Dromaeosauridae, including Deinonychus, had feathers.[24] The genus Microraptor is both older geologically and more primitive phylogenetically than Deinonychus, and within the same family.[33] Multiple fossils of Microraptor preserve pennaceous, vaned feathers like those of modern birds on the arms, legs, and tail, along with covert and contour feathers.[24] Velociraptor is geologically younger than Deinonychus, but even more closely related. A specimen of Velociraptor has been found with quill knobs on the ulna. Quill knobs are where the follicular ligaments attached, and are a direct indicator of feathers of modern aspect.[25]
Classification
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Deinonychus antirrhopus is one of the best known dromaeosaurid species,[34] and also a close relative of the smaller Velociraptor, which is found in younger, Late Cretaceous-age rock formations in Central Asia.[35][36] The clade they form is called Velociraptorinae. The subfamily name Velociraptorinae was first coined by Rinchen Barsbold in [37] and originally contained the single genus Velociraptor. Later, Phil Currie included most of the dromaeosaurids.[38] Two Late Cretaceous genera, Tsaagan from Mongolia[35] and the North American Saurornitholestes,[28] may also be close relatives, but the latter is poorly known and hard to classify.[35] Velociraptor and its allies are regarded as using their claws more than their skulls as killing tools, as opposed to dromaeosaurines like Dromaeosaurus, which have stockier skulls.[28] Phylogenetically, the dromaeosaurids represent one of the non-avialan dinosaur groups most closely related to birds.[39] The cladogram below follows a analysis by paleontologists Robert DePalma, David Burnham, Larry Martin, Peter Larson, and Robert Bakker, using updated data from the Theropod Working Group. This study currently classifies Deinonychus as a member of the Dromaeosaurinae.[40]
Size of Deinonychus (6) compared with other dromaeosauridsA study of the dromaeosaurid Kansaignathus recovered Deinonychus as a velociraptorine rather than a dromaeosaurine, with Kansaignathus being an intermediate basal form more advanced than Deinonychus but more primitive than Velociraptor.[41][42] The cladogram below showcases these newly described relationships:
A study in however, reclassified Deinonychus as a basal member of Dromaeosaurinae again.[43]
Paleobiology
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Predatory behavior
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Interpretation of a Deinonychus preying on a Zephyrosaurus in manner suggested by Fowler et al. ()Deinonychus teeth found in association with fossils of the ornithopod dinosaur Tenontosaurus are quite common in the Cloverly Formation. Two quarries have been discovered that preserve fairly complete Deinonychus fossils near Tenontosaurus fossils. The first, the Yale quarry in the Cloverly of Montana, includes numerous teeth, four adult Deinonychus and one juvenile Deinonychus. The association of this number of Deinonychus skeletons in a single quarry suggests that Deinonychus may have fed on that animal, and perhaps hunted it. Ostrom and Maxwell have even used this information to speculate that Deinonychus might have lived and hunted in packs.[44] The second such quarry is from the Antlers Formation of Oklahoma. The site contains six partial skeletons of Tenontosaurus of various sizes, along with one partial skeleton and many teeth of Deinonychus. One tenontosaur humerus even bears what might be Deinonychus tooth marks. Brinkman et al. () point out that Deinonychus had an adult mass of 70&#;100 kg (150&#;220 lb), whereas adult tenontosaurs were 1&#;4 metric tons. A solitary Deinonychus could not kill an adult tenontosaur, suggesting that pack hunting is possible.[4]
A study by Roach and Brinkman has called into question the cooperative pack hunting behavior of Deinonychus, based on what is known of modern carnivore hunting and the taphonomy of tenontosaur sites. Modern archosaurs (birds and crocodiles) and Komodo dragons typically display little cooperative hunting; instead, they are usually either solitary hunters, or are drawn to previously killed carcasses, where much conflict occurs between individuals of the same species. For example, in situations where groups of Komodo dragons are eating together, the largest individuals eat first and will attack smaller Komodos that attempt to feed; if the smaller animal is killed, it is cannibalized. When this information is applied to the tenontosaur sites, it appears that what is found is consistent with Deinonychus having a Komodo or crocodile-like feeding strategy. Deinonychus skeletal remains found at these sites are from subadults, with missing parts consistent with having been eaten by other Deinonychus.[45] On the other hand, a paper by Li et al. describes track sites with similar foot spacing and parallel trackways, implying gregarious packing behavior instead of uncoordinated feeding behavior.[46] Contrary to the claim crocodilians do not hunt cooperatively, they have actually been observed to hunt cooperatively,[47][48] meaning that the notion of infighting, competition for food and cannibalism ruling out cooperative feeding may actually be a false dichotomy.
Foot (MOR 747) in flexionIn , Manning and colleagues interpreted dromaeosaur claw tips as functioning as a puncture and gripping element, whereas the expanded rear portion of the claw transferred load stress through the structure.[49] They argue that the anatomy, form, and function of the foot's recurved digit II and hand claws of dromaeosaurs support a prey capture/grappling/climbing function. The team also suggest that a ratchet-like &#;&#;locking&#;&#; ligament might have provided an energy-efficient way for dromaeosaurs to hook their recurved digit II claw into prey. Shifting body weight locked the claws passively, allowing their jaws to dispatch prey. They conclude that the enhanced climbing abilities of dromaeosaur dinosaurs supported a scansorial (climbing) phase in the evolution of flight.[49] In , Denver Fowler and colleagues suggested a new method by which Deinonychus and other dromaeosaurs may have captured and restrained prey.[50] This model, known as the "raptor prey restraint" (RPR) model of predation, proposes that Deinonychus killed its prey in a manner very similar to extant accipitrid birds of prey: by leaping onto its quarry, pinning it under its body weight, and gripping it tightly with the large, sickle-shaped claws. Like accipitrids, the dromaeosaur would then begin to feed on the animal while still alive, until it eventually died from blood loss and organ failure. This proposal is based primarily on comparisons between the morphology and proportions of the feet and legs of dromaeosaurs to several groups of extant birds of prey with known predatory behaviors. Fowler found that the feet and legs of dromaeosaurs most closely resemble those of eagles and hawks, especially in terms of having an enlarged second claw and a similar range of grasping motion. However, the short metatarsus and foot strength would have been more similar to that of owls. The RPR method of predation would be consistent with other aspects of Deinonychus's anatomy, such as their unusual jaw and arm morphology. The arms were likely covered in long feathers, and may have been used as flapping stabilizers for balance while atop struggling prey, along with the stiff counterbalancing tail. Its jaws, thought to have had a comparatively weak bite force,[51] might be used for saw motion bites, like the modern Komodo dragon which also has a weak bite force, to finish off its prey if its kicks were not powerful enough.[52]
Bite force
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Bite force estimates for Deinonychus were first produced in , based on reconstructed jaw musculature. This study concluded that Deinonychus likely had a maximum bite force only 15% that of the modern American alligator.[51] A study by Paul Gignac and colleagues attempted to estimate the bite force based directly on newly discovered Deinonychus tooth puncture marks in the bones of a Tenontosaurus. These puncture marks came from a large individual, and provided the first evidence that large Deinonychus could bite through bone. Using the tooth marks, Gignac's team were able to determine that the bite force of Deinonychus was significantly higher than earlier studies had estimated by biomechanical studies alone. They found the bite force of Deinonychus to be between 4,100 and 8,200 newtons, greater than living carnivorous mammals including the hyena, and equivalent to a similarly-sized alligator.[53]
However, this estimate has come into question, as it was based on bite marks rather than a Deinonychus skull. A recent study used a Deinonychus skull for their estimate and calculated 706 Newtons.[54]
Gignac and colleagues also noted, however, that bone puncture marks from Deinonychus are relatively rare, and unlike larger theropods with many known puncture marks like Tyrannosaurus, Deinonychus probably did not frequently bite through or eat bone. Instead, they probably used their strong bite force for defense or to capture prey, rather than for feeding.[53]
A study by Tse, Miller, and Pittman et al., focusing on the skull morphology and bite forces of various dromaeosaurids discovered that Deinonychus, the largest taxon examined, had a skull that was well adapted to hunting of large vertebrates and delivering powerful bites to prey alongside Dromaeosaurus, to which it was compared. In this study, Deinonychus represented the most extreme specializations compared to other dromaeosaurids when it came to its adaptations. The same study also revealed that Deinonychus' skull was less resistant to bite forces than that of Velociraptor, which apparently was engaging in more scavenging behavior, suggesting high bite force resistance was more common in dromaeosaurid taxa that were obtaining food through scavenging more than engaging in active predation. it is also suggested in these findings that Deinonychus may have fed by using neck-driven pullback movements to dismember carcasses when feeding, akin to modern varanid lizards.[55]
Limb function
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Cast in climbing postureDespite being the most distinctive feature of Deinonychus, the shape and curvature of the sickle claw varies between specimens. The type specimen described by Ostrom in has a strongly curved sickle claw, while a newer specimen described in had a claw with much weaker curvature, more similar in profile with the 'normal' claws on the remaining toes.[13] Ostrom suggested that this difference in the size and shape of the sickle claws could be due to individual, sexual, or age-related variation, but admitted he could not be sure.
There is anatomical[2] and trackway[56] evidence that this talon was held up off the ground while the dinosaur walked on the third and fourth toes.
Ostrom suggested that Deinonychus could kick with the sickle claw to cut and slash at its prey.[2] Some researchers even suggested that the talon was used to disembowel large ceratopsian dinosaurs.[57] Other studies have suggested that the sickle claws were not used to slash but rather to deliver small stabs to the victim.[58] In , Manning and colleagues ran tests on a robotic replica that precisely matched the anatomy of Deinonychus and Velociraptor, and used hydraulic rams to make the robot strike a pig carcass. In these tests, the talons made only shallow punctures and could not cut or slash. The authors suggested that the talons would have been more effective in climbing than in dealing killing blows.[59] In , Manning and colleagues undertook additional analysis dromaeosaur claw function, using a numerical modelling approach to generate a 3D finite element stress/ strain map of a Velociraptor hand claw.[49] They went on to quantitatively evaluate the mechanical behavior of dromaeosaur claws and their function. They state that dromaeosaur claws were well-adapted for climbing as they were resistant to forces acting in a single (longitudinal) plane, due to gravity.
Ostrom compared Deinonychus to the ostrich and cassowary. He noted that the bird species can inflict serious injury with the large claw on the second toe.[2] The cassowary has claws up to 125 mm (4.9 in) long.[60] Ostrom cited Gilliard () in saying that they can sever an arm or disembowel a man.[61] Kofron ( and ) studied 241 documented cassowary attacks and found that one human and two dogs had been killed, but no evidence that cassowaries can disembowel or dismember other animals.[62][63] Cassowaries use their claws to defend themselves, to attack threatening animals, and in agonistic displays such as the Bowed Threat Display.[60] The seriema also has an enlarged second toe claw, and uses it to tear apart small prey items for swallowing.[64] In , a study suggested that the sickle claw would likely have been used to pin down prey while biting it, rather than as a slashing weapon.[50]
Hand bones of MOR 747Biomechanical studies by Ken Carpenter in confirmed that the most likely function of the forelimbs in predation was grasping, as their great lengths would have permitted longer reach than for most other theropods. The rather large and elongated coracoid, indicating powerful muscles in the forelimbs, further strengthened this interpretation.[65] Carpenter's biomechanical studies using bone casts also showed that Deinonychus could not fold its arms against its body like a bird ("avian folding"), contrary to what was inferred from the earlier descriptions by Jacques Gauthier[66] and Gregory S. Paul in .[28]
Studies by Phil Senter in indicated that Deinonychus forelimbs could be used not only for grasping, but also for clutching objects towards the chest. If Deinonychus had feathered fingers and wings, the feathers would have limited the range of motion of the forelimbs to some degree. For example, when Deinonychus extended its arm forward, the 'palm' of the hand automatically rotated to an upward-facing position. This would have caused one wing to block the other if both forelimbs were extended at the same time, leading Senter to conclude that clutching objects to the chest would have only been accomplished with one arm at a time. The function of the fingers would also have been limited by feathers; for example, only the third digit of the hand could have been employed in activities such as probing crevices for small prey items, and only in a position perpendicular to the main wing.[26] Alan Gishlick, in a study of Deinonychus forelimb mechanics, found that even if large wing feathers were present, the grasping ability of the hand would not have been significantly hindered; rather, grasping would have been accomplished perpendicular to the wing, and objects likely would have been held by both hands simultaneously in a "bear hug" fashion, findings which have been supported by the later forelimb studies by Carpenter and Senter.[67] In a study conducted by Bruce Rothschild and other paleontologists, 43 hand bones and 52 foot bones referred to Deinonychus were examined for signs of stress fracture; none were found.[68] The second phalanx of the second toe in the specimen YPM has a healed fracture.[69]
Parsons and Parsons have shown that juvenile and sub-adult specimens of Deinonychus display some morphological differences from the adults. For instance, the arms of the younger specimens were proportionally longer than those of the adults, a possible indication of difference in behavior between young and adults.[70] Another example of this could be the function of the pedal claws. Parsons and Parsons have suggested that the claw curvature (which Ostrom [] had already shown was different between specimens[13]) maybe was greater for juvenile Deinonychus, as this could help it climb in trees, and that the claws became straighter as the animal became older and started to live solely on the ground.[71] This was based on the hypothesis that some small dromaeosaurids used their pedal claws for climbing.[59]
Flight
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In a paper, it was reported after further analysis of immature fossils that the open and mobile nature of the shoulder joint might have meant that young Deinonychus were capable of some form of flight.[72]
Speed
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Restoration of a walking individualDromaeosaurids, especially Deinonychus, are often depicted as unusually fast-running animals in the popular media, and Ostrom himself speculated that Deinonychus was fleet-footed in his original description.[10] However, when first described, a complete leg of Deinonychus had not been found, and Ostrom's speculation about the length of the femur (upper leg bone) later proved to have been an overestimate. In a later study, Ostrom noted that the ratio of the femur to the tibia (lower leg bone) is not as important in determining speed as the relative length of the foot and lower leg. In modern fleet-footed birds, like the ostrich, the foot-tibia ratio is .95. In unusually fast-running dinosaurs, like Struthiomimus, the ratio is .68, but in Deinonychus the ratio is .48. Ostrom stated that the "only reasonable conclusion" is that Deinonychus, while far from slow-moving, was not particularly fast compared to other dinosaurs, and certainly not as fast as modern flightless birds.[13]
The low foot to lower leg ratio in Deinonychus is due partly to an unusually short metatarsus (upper foot bones). The ratio is actually larger in smaller individuals than in larger ones. Ostrom suggested that the short metatarsus may be related to the function of the sickle claw, and used the fact that it appears to get shorter as individuals aged as support for this. He interpreted all these features&#;the short second toe with enlarged claw, short metatarsus, etc.&#;as support for the use of the hind leg as an offensive weapon, where the sickle claw would strike downwards and backwards, and the leg pulled back and down at the same time, slashing and tearing at the prey. Ostrom suggested that the short metatarsus reduced overall stress on the leg bones during such an attack, and interpreted the unusual arrangement of muscle attachments in the Deinonychus leg as support for his idea that a different set of muscles was used in the predatory stroke than in walking or running. Therefore, Ostrom concluded that the legs of Deinonychus represented a balance between running adaptations needed for an agile predator, and stress-reducing features to compensate for its unique foot weapon.[13]
In his study of Canadian dinosaur footprints, Richard Kool produced rough walking speed estimates based on several trackways made by different species in the Gething Formation of British Columbia. Kool estimated one of these trackways, representing the ichnospecies Irenichnites gracilis (which may have been made by Deinonychus), to have a walking speed of 10.1 kilometers per hour (6 miles per hour).[73]
Eggs
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Artist's impression of an individual in brooding positionThe identification, in , of a probable Deinonychus egg associated with one of the original specimens allowed comparison with other theropod dinosaurs in terms of egg structure, nesting, and reproduction. In their examination of the specimen, Grellet-Tinner and Makovicky examined the possibility that the dromaeosaurid had been feeding on the egg, or that the egg fragments had been associated with the Deinonychus skeleton by coincidence. They dismissed the idea that the egg had been a meal for the theropod, noting that the fragments were sandwiched between the belly ribs and forelimb bones, making it impossible that they represented contents of the animal's stomach. In addition, the manner in which the egg had been crushed and fragmented indicated that it had been intact at the time of burial, and was broken by the fossilization process. The idea that the egg was randomly associated with the dinosaur was also found to be unlikely; the bones surrounding the egg had not been scattered or disarticulated, but remained fairly intact relative to their positions in life, indicating that the area around and including the egg was not disturbed during preservation. The fact that these bones were belly ribs (gastralia), which are very rarely found articulated, supported this interpretation. All the evidence, according to Grellet-Tinner and Makovicky, indicates that the egg was intact beneath the body of the Deinonychus when it was buried. It is possible that this represents brooding or nesting behavior in Deinonychus similar to that seen in the related troodontids and oviraptorids, or that the egg was in fact inside the oviduct when the animal died.[8]
Examination of the Deinonychus egg's microstructure confirms that it belonged to a theropod, since it shares characteristics with other known theropod eggs and shows dissimilarities with ornithischian and sauropod eggs. Compared to other maniraptoran theropods, the egg of Deinonychus is more similar to those of oviraptorids than to those of troodontids, despite studies that show the latter are more closely related to dromaeosaurids like Deinonychus. While the egg was too badly crushed to accurately determine its size, Grellet-Tinner and Makovicky estimated a diameter of about 7 cm (2.8 in) based on the width of the pelvic canal through which the egg had to have passed. This size is similar to the 7.2 cm (2.8 in) diameter of the largest Citipati (an oviraptorid) eggs; Citipati and Deinonychus also shared the same overall body size, supporting this estimate. Additionally, the thicknesses of Citipati and Deinonychus eggshells are almost identical, and since shell thickness correlates with egg volume, this further supports the idea that the eggs of these two animals were about the same size.[8]
A study published in November by Norell, Yang and Wiemann et al., indicates that Deinonychus laid blue eggs, likely to camouflage them as well as creating open nests. The study also indicates that Deinonychus and other dinosaurs that created open nests likely represent an origin of color in modern bird eggs as an adaptation both for recognition and camouflage against predators.[74][75][76]
Life cycle
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A study on Deinonychus tooth isotopes suggests precociality in the genus. The isotopes examined for different aged specimens indicates that adults and juveniles had different diets across the various age groups. As the data suggests that Deinonychus had a more typical reptilian set of life stages, the examinations also have been stated to indicate a lack of complex, cooperative social behavior found in mammalian terrestrial pack-hunters such as wolves.[77]
Paleoenvironment
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Mounted skeletons of Deinonychus and the contemporary Zephyrosaurus (left), Natural History Museum of DenmarkGeological evidence suggests that Deinonychus inhabited a floodplain or swamplike habitat.[34] The paleoenvironment of both the upper Cloverly Formation and the Antlers Formation, in which remains of Deinonychus have been found, consisted of tropical or sub-tropical forests, deltas and lagoons, perhaps similar to the environment of modern-day Louisiana.[78][79] Other animals Deinonychus shared its world with include herbivorous dinosaurs such as the nodosaurid Sauropelta and the ornithopods Zephyrosaurus and Tenontosaurus. In Oklahoma, the ecosystem of Deinonychus also included the large theropod Acrocanthosaurus, the huge sauropod Sauroposeidon, the crocodilians Goniopholis and Paluxysuchus, and the gar Lepisosteus.[79] If the teeth found in Maryland are those of Deinonychus, then its contemporaries would include the sauropod Astrodon and the poorly-known nodosaur Priconodon. The middle portion of the Cloverly Formation ranges in age from 115 ± 10 Ma near the base[5] to 108.5 ± 0.2 Ma near the top.[6]
Cultural significance
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Hypothetical feathered model adjacent to an earlier, featherless model from the Westphalian Museum of Natural HistoryDeinonychus were featured prominently in Harry Adam Knight's novel Carnosaur and its film adaption, and Michael Crichton's novels Jurassic Park and The Lost World and their film adaptations, directed by Steven Spielberg. Crichton ultimately chose to use the name Velociraptor for these dinosaurs, rather than Deinonychus. Crichton had met with John Ostrom several times during the writing process to discuss details of the possible range of behaviors and life appearance of Deinonychus. Crichton at one point apologetically told Ostrom that he had decided to use the name Velociraptor in place of Deinonychus for his book, because he felt the former name was "more dramatic". Despite this, according to Ostrom, Crichton stated that the Velociraptor of the novel was based on Deinonychus in almost every detail, and that only the name had been changed.[80]
The Jurassic Park filmmakers followed suit, designing the film's models based almost entirely on Deinonychus instead of the actual Velociraptor, and they reportedly requested all of Ostrom's published papers on Deinonychus during production.[80] As a result, they portrayed the film's dinosaurs with the size, proportions, and snout shape of Deinonychus.[2] The 20-foot-long (6.1 m) Utahraptor is commonly considered to be a close match to the film's dinosaurs, which are much larger than either Deinonychus or Velociraptor were in life.[81][82]
See also
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References
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