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Did Velociraptors have feathers?All dromaeosaurids had feathers, because we have the evidence that indicates that they did.In other dinosaurs, what feathers and what extent is still a little iffy. But for the dromaeosaurids, and the maniraptors in general, we have the evidence.In fact, I feel this answer would be best served by giving a scientifically accurate view of Velociraptor mongoliensis, which is my favorite dinosaur, so how can I pass up this opportunity to go over the science?We’ll start with the history of the animal’s discovery, then move onto the science of the animal, supplemented with peer-reviewed scientific papers describing the specimens, go over one fantastic specimen of note and the story of how the two animals died.Then we’ll cover some of the animal’s ecology, at least what we know.Finally, we’ll put it all together in the form of paleo-art, created by talented artists that work closely with the scientists.Why?Because I feel paleo-art is important, especially if the artist works with the paleontologists closely, because we all can see the fossils. What really captures the imagination is bringing these fascinating creatures back to life, if only in the forms of drawings and paintings.The science describes the details, the art brings those details to life.Anyhow, without further ado, let’s discuss Velociraptor in a scientifically accurate manner.HISTORY OF DISCOVERY:During an American Museum of Natural History expedition to the Outer Mongolian Gobi Desert, on 11 August 1923 Peter Kaisen recovered the first Velociraptor fossil known to science: a crushed but complete skull, associated with one of the raptorial second toe claws.This is the holotype specimen that they discovered:In 1924, museum president Henry Fairfield Osborn designated the skull and claw (which he assumed to come from the hand) as the holotype specimen of his new genus, Velociraptor.The name itself is derived from the Latin. “Velox” meaning “swift,” and “raptor,” meaning “thief,” or “plunderer.”In fact, raptor means snatching things so another way to explain the name is that Velociraptor, is literally a fast snatcher.Osborn named the type species V. mongoliensis after its country of origin, Mongolia.Earlier that year, Osborn had mentioned the animal in a popular press article, under the name "Ovoraptor djadochtari" (not to be confused with the similarly named dinosaur Oviraptor).However, because the name "Ovoraptor" was not published in a scientific journal or accompanied by a formal description, it is considered a nomen nudium, which is Latin for “naked name”, being a fancy Latin way of saying that the name has no meaning, and thus the name Velociraptor retains priority.In my opinion, it’s good that the name Velociraptor retains priority, because it both summarizes the animal, as well as the fact that it’s just a cool name that rolls off the tongue easily.While North American teams were shut out of communist Mongolia during the Cold War, expeditions by Soviet and Polish scientists, in collaboration with their Mongolian colleagues, recovered several more specimens of Velociraptor.The most famous is part of the famous "Fighting Dinosaurs" specimenHere’s the specimen in question:It shows a Protoceratops in a battle against a Velociraptor. We’ll discuss this beautiful specimen at length, later.This specimen is considered a national treasure of Mongolia, although in 2000 it was loaned to the American Museum of Natural History in New York City for a temporary display.All found in the Gobi Desert. The Gobi Desert is very fossil rich, and often preserves animals in three dimensions, rather than the typical being squished by tons and tons of rock that we see in other dinosaurs.As you can imagine, a lot of work was taken underway, considering these fossils are found in the middle of the freaking Gobi Desert!Which is why Mongolian paleontology is often an area of political, economic, and physical barriers.Because there’s loads of specimens of Velociraptor, I cannot possibly cover them all, just went over a few that I feel are most important.That’s due to several specimens discovered in the 1990’s that still are being formally described.So that’s the history of the discovery of this dinosaur. Let’s discuss the science.SIZE:Because of Jurassic Park basically taking a Deinonychus, a closely related relative to Velociraptor, and scaling it up to the size of Achillobator, a large dromaeosaurid:Many people look at the size of Velociraptor, and rightly think, “That’s it?!”But that’s kind of unfair, since Velociraptor is considered to be a medium sized dromaeosaurid.They’re not small. They’re around 2 meters long, and weigh in the neighborhood of 15–20 kg.That’s decently sized, if you ask me.So that kid in Jurassic Park was kinda on to something when he said that Velociraptor is a 6 ft turkey.Well yeah. I like to think of them as funny looking, flightless hawks, with a reptilian maw full of teeth, and long tails, because that’s kinda what they look like.HEAD SHAPE:Also because of the Velociraptors in Jurassic Park, many people look at a real Velociraptor skull, and are kinda confused.See, you were led to believe this:That looks like to me, more like a Deinonychus skull, not Velociraptor.Here’s the skull of Deinonychus.And here’s the skull of Velociraptor:Much narrower and much more gracile than you’ve been led to believe eh?I determine this by the snout. Velociraptor’s snout kinda swoops and has a concave, as the holotype skull I showed earlier does.Now, you might say, “But that means it didn’t have that strong of a bite!”Well, yeah. Dromaeosaurids in general don’t have that powerful of a bite, as they evidently used them for eating. Their mouths aren’t their primary weapon, or the way they killed things.Interesting thing about the skull, when reconstructing the animal, many artists put extra-oral tissues on the face, colloquially called lips.Now, lips on dinosaurs are somewhat of a contentious topic open for debate right now, but these two posts explain what the skulls can show:Making Lip of ItExposed teeth in dinosaurs, sabre-tooths and everything else: thoughts for artistsThe second one is by Mark Witton, a paleontologist, and he makes a pretty convincing case for extra-oral tissues.Basically put, the rougher the skull, the less flesh is covering it, and the smoother the skull, the more that flesh is covering it.Velociraptors seem to have had extra-oral tissues on their face. As to what extent, that’s up for debate.Since we’re talking about the flesh on the face, let’s talk about the stuff that would have poked out of the flesh.INTEGUMENT:We know from related animals that have been preserved with feather impressions that there’s reticulate scales near the front of the snout, feathers covering the eyes, and the rest of the head.This assertion is supported by this paper:Basal paravian functional anatomy illuminated by high-detail body outlineThe paper describes a related dinosaur, Anchiornis, that has preserved feather and skin impressions, and gives a great, accurate look into the anatomy of the creature.Sometimes you might see some dromaeosaurids depicted with fun crests on their heads, and while it’s not outright wrong, it is based on a mistake on part of the artists when looking at this Microraptor fossil, a dinosaur in the same family as Velociraptor:Paleontologists, being the type of people that they are, say that this weird feather crest that you see in the fossil, is just the weird way that the feathers were flattened. Apparently this poor Microraptor was squished.So, it’s not definitely wrong, but one ought to be cognizant of the science. :)Speaking of the Microraptor fossil for a bit, you may notice the clear feather impressions. Since Microraptor is in the exact same group as the other dromaeosaurids that we’ve mentioned, phylogenetic bracketing makes a very strong case that all of them were feathered, at least to the extent of Microraptor.In other fossils of Microraptor, you see the impressions a bit more clearly:And another dromaeosaurid, Sinornithosarus, shows clear evidence of feathers as well:A bit harder to see, but the animal is completely covered in feathers.In another fossil of this dinosaur, it also shows clear feather impressions:We also see it in another closely related animal, Zhenyuanlong:How does all of this apply to Velociraptor?Well, because if these three very closely related animals have these advanced feathers, then it’s reasonable to argue that Velociraptor, Deinonychus, Utahraptor, and Dakotaraptor were feathered as well.But for Velociraptor, things are even more dramatic. We have an ulna of a Velociraptor that shows clear quill knobs where the feathers were anchored, much like how we see in modern birds:So it would seem that Velociraptor had wings like a bird.It’s also not unreasonable to deduce that because it had wings, that the rest of the animal was covered with feathers as well.We know from Microraptor and Sinornithosaurus, that the tail would either have a fan, or a frond.That’s supported by this paper:Tail anatomy of Buitreraptor gonzalezorum (Theropoda, Unenlagiidae) and comparisons with other basal paraviansBecause of the ossified bony rods in the tail of Velociraptor, it would be reasonable to assume a tail fan, at least, since the rods seem to served to keep their tails up off the ground.Speaking of crests and feathers, this paper by Holliday, et al. discusses hollows in archosaur heads, which would of course, include Velociraptor:The Frontoparietal Fossa and Dorsotemporal Fenestra of Archosaurs and Their Significance for Interpretations of Vascular and Muscular Anatomy in DinosaursBasically, the paper is discussing some hollow holes on the top of the skulls of some dinosaurs, and since they aren’t sites for muscle attachment, the scientists theorize they might have served the purpose that they do in crocodylians and avians, in that they served as thermoregulation, since they seem to have been an area rich in blood.But, that thermoregulation is magnified if you add a dewlap, a snood, caruncular skin, like in a turkey, or wattles, and such.While I don’t think that Velociraptor had these structures, we should keep that in the back of our mind. :)So despite what Jurassic Park has led people to believe, Velociraptors, and its other closely related kin were basically grounded hawks with nasty claws, a reptilian maw full of teeth, and long tails.In fact, one of the references for dromaeosaurid reconstruction is birds of prey like eagles, hawks, and falcons.I mean are you willing to contend with those claws?I’m not. Listen to the eagle, maybe he’s had a hard day.And nobody hesitates to call them apex predators of the sky, and nobody thinks their feathers look stupid or ridiculous, so why do we assume it does for Velociraptor?VELOCIRAPTOR ECOLOGY:I hesitate to call Velociraptor’s “wings” as wings because that has the implication that they served some kind of aerodynamic function that you see in birds when they do Wing Assisted Incline Running, or use their wings other than for flying.The reason? Velociraptor didn’t have much in the way of up and down movement.Comparison of forelimb function between Deinonychus and Bambiraptor (Theropoda: Dromaeosauridae)So Velociraptor couldn’t use its wings for an aerodynamic function to make it more agile in running.Speaking of running, Velociraptor might not have been the most swift runner out there.Think more like an ambush predator. It stalks its prey, then waits for the moment, ambushes it, and kills it with its second toe claw.As far as the tail, you often see dromaeosaurids shown with a rigid, stiff tail, because the tail does have ossified bony rods in it.But we know from an articulated Velociraptor fossil, which as the tail bent in an S curve, that they could move their tails from side-to-side, but not up and down, demonstrating that the rods where helping to keep their tails up off the ground.Talking of the rods, they also seem to be ossified tendons, allowing good control of the tail, without the need of big ol’ muscles to facilitate control of the tail.A lot of the muscles that dinosaurs like Tyrannosaurus rex, and Allosaurus would have to allow them to move their tails are reduced in Velociraptor.So it seems that this might fill in the puzzle a bit more.What this means is that Velociraptor’s tail is not this rigid, muscular thing like in other theropods, but this spindly, feathered covered thing.This applies to dromaeosaurids in general, as they have reduced caudal musculature, so this makes some sort of sense.One particular muscle is of note, the caudis femoralis muscle that goes from the tail and into the back of the leg to allow for more efficient running, is reduced in dromaeosaurids in general.Further, their lower legs are quite short, even by dromaeosaurid standards.What implications does this have?Well, they might not have been as fast as you’ve been led to believe.See, in order to be a strong runner in the dinosaur sense, you’d need nice, strong muscles to facilitate it, and nice long legs to increase the length of your stride.Further, if you have an arctometatarsal, which is the middle metatarsal being squished by the other two metatarsals, like we see in T. rex:Where this foot structure improves the structure and rigidity of the foot, allowing the animal to regain energy on each stride, the better.But Velociraptor doesn’t have this.Here’s a Velociraptor foot:It doesn’t have an arctometatarsal, meaning it would lose out on potential energy with each stride.Further, the muscles that facilitate swift running are reduced in dromaeosaurids in general.And, in Velociraptor’s case, its legs are rather short, meaning it doesn’t have a long stride, reducing its speed, inherently.This means it’s not a swift runner.Think more like an ambush predator like a jaguar, and you have the right idea of what it was doing.All of this led to researchers Fowler, et al. to discuss the predatory ecology of dromaeosaurids, using Deinonychus as the example.I’ve referenced this study before in other answers relating to dromaeosaurids, but here’s the study in question:The Predatory Ecology of Deinonychus and the Origin of Flapping in BirdsThe idea is that the animal would stalk its prey, then ambush it and pin it down with its body weight, and using its hyper-extended second toe claw, and other claws of the feet to dig in to the animal’s flesh, pin it, then use its mouth to eat the animal alive.Indeed, a lot like hawks do!Further in the study, they determined the gripping articulation of Deinonychus’s foot as a model for the other dromaeosaurids, and found they had a great degree of articulation and power in grip strength.So while they certainly could kick an animal to death, it’s not as efficient, because they couldn’t transfer much of their strength through the claw, so their preferred method was using gravity and their weight to hold the animal down as they gripped really hard to hold the animal.That looks like something I’ve seen before:Nasty?You betcha. But it’s a ridiculously successful method to kill an animal, if you’re a natural predator.But, am I saying they never used their claw to kick and jam it into an animal’s hide to kill it?No.Behold, the Fighting Dinosaurs specimen that I linked before:This here Velociraptor is kicking this poor old Protoceratops right in the jugular with its left leg.“Did Velociraptor and its closely related cousins ever hunt in packs to bring a large herbivore like Protoceratops down?” people might ask.Well, doesn’t seem like it.This study by Xing, et al. shows a group of fossils of a group deinonychosaurians all moving in the same direction.Multiple parallel deinonychosaurian trackways from a diverse dinosaur track assemblage of the Lower Cretaceous Dasheng Group of Shandong Province, ChinaBut it’s quite a leap in logic to infer pack hunting behavior from this.Because pack hunting in general is a pretty sophisticated hunting strategy, and the fact that only few groups of mammals in the entirety of the animal kingdom do it, should be enough to paint a picture of how sophisticated it really is, and why most animals don’t engage in the behavior.Compounded with this is the fact that mammals, in general, are pretty smart.“Well, surely, Velociraptor and its closely related kin were smart, right?”Well, nobody’s sure of the level of intelligence for any extinct paleo-fauna, much less dromaeosaurids, and more specifically Velociraptor.See, inferring intelligence in living mammals has been proven tough to estimate, and that’s animals that have brains that we can study.With non-avian dinosaurs, the brains have rotted away due to millions of years of being fossilized and in the ground. We have no brains to go on.That said, we have studied the braincase of Velociraptor:Feathered DragonsAll we’ve been able to deduce is that Velociraptor’s brain was more devoted to smell, than in sight, like in birds.As far as problem solving ability, we may never be able to know, unfortunately.So lacking fossils, and lacking observing their behavior directly (we’re 65 million years too late), I’d say to be skeptical of the pack-hunting behavior for now.I’ve seen Velociraptor depicted with slit pupils, and that’s not entirely unreasonable, according to this paper:Nocturnality in Dinosaurs Inferred from Scleral Ring and Orbit MorphologyThe ring tells us two things:The focal length of the eye.And the size of the aperture, or the opening of the pupil.This gives us the animal’s F number. If it has a low F number, it’s adapted to low-light conditions, if it’s higher, it’s adapted to high light conditions.The paper demonstrates that Velociraptors had low F numbers, indicating their eyes were adapted to low-light conditions, much like the light during twilight, both dawn, and the evening.Thus we can say that Velociraptor was most active during twilight hours.Funnily enough Protoceratops, was not strictly nocturnal, as to summarize the fancy science speak, it was active when it was hungry.Thus, the Fighting Dinosaurs fossil, the researchers conclude, most plausibly occurred in the twilight hours of the morning, or the evening.What does this mean for Velociraptor?Well, it needs to be able to see well at night, but retain color vision so that they can see with a good amount of detail in the twilight hours of morning or evening.And so we need to find a modern, extant animal that’s closely related to Velociraptor needs to help fill in that gap for us.Here’s the animal in question:So a slit pupil for Velociraptor does make sense considering when it was most active, and considering the fact it needed to be able to see with a good deal of clarity in twilight hours of morning or evening.Going back to the Fighting Dinosaurs fossil:What exactly happened here?So my description is based on this analysis:Combattenti per l'EternitàIt’s twilight in the Gobi Desert.Evidently, this Protoceratops was stuck in unstable sand. Now considering that Protoceratops was heavier than Velociraptor, I surmise that Velociraptor may have only occasionally fed upon Protoceratops.Well, this Protoceratops was stuck in this unstable sand, and along comes a Velociraptor, and sees an opportunity that it couldn’t pass up.It leaps onto Protoceratops, attempting to dig its claws in.Well, Protoceratops is a bit scrappy himself, and bites onto Velociraptor’s arm. The Velociraptor, getting desperate, grabs the face of Protoceratops, and kicks its leg out into the throat of Protoceratops, killing it, but now Velociraptor is pinned down by this large herbivore and can’t move.Then either a sandstorm, or collapsing sand dune, buries the animals, so that we discover it some 75 million years later.PALEO-ART:Now comes the fun part, paleo-art!This gives a general look of the animal.Here’s one by the fantastic paleo-artist, Emily Willoughby. Do go and follow her, as her art is fantastic, and being a smart cookie herself, works very hard and closely with paleontologists to ensure she’s being fair to the science, as well as her artistic inspiration.Here’s a Velociraptor by Emily Willoughby again, showing it hunting its primary food source, small mammals.Here’s one from paleontologist Mark Witton, as he’s a gifted paleo-artist himself.Here’s another one from Mark Witton, showing what happened to their prey when they caught it.CONCLUSION:Whew! That was a lot to cover!Hope this answers the question sufficiently, and I hope I was comprehensive as well as succinct.For a summary, here’s this fantastic video from Your Dinosaurs Are Wrong, discussing Velociraptor:

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