Tyrannosaurid skin impressions

[amargasaurus cazaui]

It has been pointed out for me multiple times but people who know theropods far better than I do but.....

It isn't so much that the xixian likely had anything unique going on featherwise, meaning there were likely feathered equivalents in other environments.......for instance in Hell Creek consider Anzou, most likely feathered...BUT.....
Xixia basin belongs to the the area known as the Jehol biota, a massive area that contains a lagerstatten type preversavtional area....there are few of them in the world and they produce the most exquisite feathered fossils known.......the area of Germany where some of the archaeopteryx fossils were found is another. It has been often repeated that is not so much a matter of if the dinosaurs were feathered, as the preservational bias against feathers in areas of the world makes it unlikely we find them.
    Another thing to consider is this...when many of the theropod fossils from the North American continent were found, those paleos had no concept they could or should be feathered and could has missed the feather impressions altogether...we do not know. If they were not looking for feathered Tyrannosaurus it is quite possible they chopped and removed bones rich in feather  type impressions and failed to understand what they were seeing....
A good analogy would be the story told of a paleo from the late 1800's that passed through an area and made note of layers and layers of "clam shells" fossilized....later on a hunch Horner retraced the mans steps and found instead all those clam shells were the remains of fossil dinosaur eggs, the first found in this hemisphere...but incorrectly identified as clams.
    I asked many times why we seldom find feathers here or for that matter eggshell, or fossil eggs......it happens but nearly as commonly as other environments. The answer seems to always come up preservation...the area was heavily volcanic and had high acidic content which may well have served to dissolve fragile eggshell,......or feathers for that matter.
Something was definitely happening differently here regarding preservation as opposed to the xixia.....

[Sim]

Avialans are what people often consider birds.  Archaeopteryx and Anchiornis have been classified within this group.  When you've been saying "avialan", do you perhaps actually mean "paravian"?  Paravians are all the avialans, dromaeosaurids, troodontids and scansoriopterygids: https://en.wikipedia.org/wiki/Paraves

Wikipedia says:

https://en.wikipedia.org/wiki/Bird
Aves can mean those advanced archosaurs with feathers (alternately Avifilopluma)
Aves can mean those that fly (alternately Avialae)
Aves can mean all reptiles closer to birds than to crocodiles (alternately Avemetatarsalia)
Aves can mean the last common ancestor of all the currently living birds and all of its descendants (a "crown group", in this sense synonymous with Neornithes)

Avialae isn't always synonymous with "bird/Aves". The clearest one to use is #4, and it's been more or less the gold standard since it was proposed (at least among ornithology).

https://en.wikipedia.org/wiki/Avialae
Avialae can also be used in reference to "the theropod group that includes all taxa closer to Passer than to Dromaeosaurus" which is a lot better than "those that fly" which can include everything from dromaeosaurs to modern birds but not all of each.

As I said in my post, "Avialans are what people often consider birds."  In my experience this is true, and it refers to people as a whole, which is why I often see the following considered birds: Archaeopteryx, Hesperornis, Ichthyornis, Confuciusornis, Balaur.  I only said what I said to try and make clearer what I was explaining to Neosodon, and I specifically said "Avialans are what people often consider birds.", so I clearly didn't say it's always synonymous with "bird".

Phylogenetic groups are often defined in more than one way, with some definitions becoming outdated, and some as you pointed out creating weird unnatural groups.  I don't think it's worthwhile getting too concerned about this, which is what I think is happening here.  My understanding is Avialae is generally considered the natural group that contains all the species closest to and including extant birds from the point where dromaeosaurids and troodontids won't be part of the group.  The definition by Jacques Gauthier of Avialae based on those that fly is not one I've seen followed, I imagine because it can create unnatural groups and is unreliable (it's not clear whether some species could fly and at what point or points flight evolved), which makes it a pointless definition.

As mentioned on the Avialae Wikipedia page:

Gauthier[9] (page 34) identified four conflicting ways of defining the term "Aves", which is a problem because the same biological name is being used four different ways. Gauthier proposed a solution, number 4 below, which is to reserve the term Aves only for the crown group, the last common ancestor of all living birds and all of its descendants. He assigned other names to the other groups.
[...]
Gauthier's proposals have been adopted by many researchers in the field of paleontology and bird evolution, though the exact definitions applied have been inconsistent. Avialae, initially proposed to replace the traditional fossil content of Aves, is sometimes used synonymously with the vernacular term "bird" by these researchers.[7]

The point from which something is considered a "bird" is still inconsistent among people, including specialists.  And as hinted by the last sentence in the above quote, avialan = bird is still widely used, in my experience being the most common meaning of "bird" or "avialan" which is why I said what I did in my explanation to Neosodon - I intended to use a simple explanation over a complicated one that could be confusing.

[Dinoguy2]

My theory on non avialan feather evolution is that they evolved in small dinosaurs that lived in cold environments. The perfect example is the Yixian Formation. All of the early feathered coelosaurs I know of lived there such as Sinosauropteryx, Sinocalliopteryx and Dilong. This makes it pretty clear why Yutyrannus was feathered as it also lived at the same time in the same formation and was also a coelosaur. There was something about the Yixian that pushed feather growth. Most likely the cold climate.

Some of this is contradicted by known information though.  The Yixian is a Cretaceous environment, and the more basal coelurosaurs from this formation e.g. the tyrannosauroids, compsognathids and therizinosaur all only have primitive filamentous feathers.  However, they coexisted with dromaeosaurids, troodontids, avialans and oviraptorosaurs which already had advanced pennaceous feathers which shows feathers had been evolving long before the Yixian.  As proof of this, feathers are known from Jurassic dinosaurs such as scansoriopterygids, Archaeopteryx, Anchiornis and Juravenator, and all but the last one already have pennaceous feathers, so the evolution of feathers would've started long before these as well.

The chances of a large dinosaur living in a hot climate being feathered is almost zero. The chances of a small dinosaur living in a cold climate being feathered is very high. Tyrannosaurus was a very large dinosaur living in a moderate climate so even without the fossil evidence of scales the chances of it being feathered is still very low.

Keep in mind this theory only applies to non avialans and can not be used on dromeosaurs and troodontids. But every feathered non avialan was either small or lived in a cold climate.

Dakotaraptor was a large non-avialan that was definitely feathered as shown by its preserved quill knobs, and it didn't live in a cold environment as it's from within the upper 20m of the Hell Creek Formation, where based on the info from the supplementary material, the annual average temperature was 23 degrees Celsius.

It's not like Hell Creek was an open plain, it had a lot of marshes but was still very forested. I'm not sure why the conifer forests of the Yixian were any more suited to large feathered animals than the deciduous and palm forests of Hell Creek. Scales would probably be better suited to moving though thick brush than feathers.

The impression I got reading the paper was that the Yixian was more densely forested than Hell Creek and the other tyrannosaurid environments it referred to.  My thinking is a denser forest means less sunlight and more shade, and as a result the environment is generally cooler even if the actual temperature isn't different to that of a less forested environment.

I'm not claiming feathers first evolved in the Yixian. I'm just saying that is where they first have a strong showing in the fossil record in non avialans and were they may have been introduced into The Tyrannosaur family tree.

Scansoriopterygids, Archaeopteryx and Anchiornis were avialans I think. They all used there feathers for locomotion. Juravenator was a small non avialan that lived in northern Europe which may have been cool at the time. Dakotaraptor is also an avialan too. All dromeosaurids are avialans I think.

Basically I think downy/filamentous feathers evolved to help small dinosaurs survive in cool environments. The first feathery dinosaurs like Juravenator may have evolved their feathers in cool mountainous environments like the Yixian.

The environment of Juravenator was anything but cool. It's from the Solnhofen archipelago, which was a very hot, arid environment surrounded by coral and sponge reefs in very warm water. But, of course Juravenator kind of proves the point: It was 99% covered in scales, with apparently just a sparse covering of the thinnest little short, stubbly hair-like feathers you can imagine. It would have actually looked a lot like that Schoutan painting again.

This is a pretty great match for the feathering of Juravenator:



Interestingly, Aerodactylus is from the same environment and it has a very similar amount and length of pycnofibres, though covering more of the body and minus scales. It COULD be that species with "protofeathers", when they lose them for temperature reasons, generally end up with a "pycnofibre"-like short stubble pelt on limited parts of the body.

[Sim]

Adding to what Dinoguy2 said, Archaeopteryx and Compsognathus are also from the Solnhofen archipelago.  Fossils of Archaeopteryx show it was extensively feathered, with large remiges and rectrices, basically it's feathering is bird-like.  Compsognathus however doesn't have any feathers preserved even though it's from the same place, and actually has scales preserved on its tail similar to Juravenator.  This makes me think several things:

- Compsognathus most likely had reduced feathering like Juravenator, rather than extensive feathering like Sinosauropteryx and Sinocalliopteryx which both lived in the relatively cold Yixian Formation.

- Compsognathus and Juravenator could be interpreted as showing these animals have such reduced feathering due to the combination of living in a hot environment and only having filamentous feathers.  Archaeopteryx however, could be interpreted as showing that an animal living in the same hot environment but which has pennaceous feathers, still retains its extensive feathering.

[Faelrin]

I thought Juravenator was supposed to look something like this? https://en.wikipedia.org/wiki/File:Juravenator_by_Tom_Parker.png The article even mentions a study done under UV light where more extensive filaments were discovered that was similar to compsognathids. I don't think I'd be able to read this paper to see what exactly they mean however since it's one of those that's behind a paywall.

The more I read the more I am doubting Tyrannosaurus would have been covered like the Saurian T. rex, let alone anything more extensively (though it seems the T. rex done for the autopsy show might still be within the realms of possibility, in regards to its feathering, maybe).

I hope the rest of what I'm posting is on topic. Now as much as I like the Safari and CollectA versions, I think out of curiosity I'd like to see a figure done now with minimal feathering (just on the back and sparse I guess), and in regards to this evidence (regardless if they are scales, skin, whatever). Considering how small those impressions would be on such a much larger animal, they wouldn't need to be visible or sculpted on a much smaller figure, although I wonder if that would mean it would feel textureless in those areas as a result, beyond skin folds I guess.

Actually that brings me to another question that I think I asked before, but why were these so small? If these are actually scales, in what ways would these be beneficial to T. rex at this size?

[Neosodon]

I thought Juravenator was supposed to look something like this? https://en.wikipedia.org/wiki/File:Juravenator_by_Tom_Parker.png The article even mentions a study done under UV light where more extensive filaments were discovered that was similar to compsognathids. I don't think I'd be able to read this paper to see what exactly they mean however since it's one of those that's behind a paywall.

The more I read the more I am doubting Tyrannosaurus would have been covered like the Saurian T. rex, let alone anything more extensively (though it seems the T. rex done for the autopsy show might still be within the realms of possibility, in regards to its feathering, maybe).

I hope the rest of what I'm posting is on topic. Now as much as I like the Safari and CollectA versions, I think out of curiosity I'd like to see a figure done now with minimal feathering (just on the back and sparse I guess), and in regards to this evidence (regardless if they are scales, skin, whatever). Considering how small those impressions would be on such a much larger animal, they wouldn't need to be visible or sculpted on a much smaller figure, although I wonder if that would mean it would feel textureless in those areas as a result, beyond skin folds I guess.

Actually that brings me to another question that I think I asked before, but why were these so small? If these are actually scales, in what ways would these be beneficial to T. rex at this size?

The examination under UV revealed a more extensive covering of filament-like structures, similar in anatomy to the primitive feathers of other compsognathids, including Sinosauropteryx.
https://en.wikipedia.org/wiki/Juravenator

Juravenator may be more feathered than once thought but it was still very small. For a non paravian dinosaur to evolve a downy feathers coating it has to be both small and living in a cool environment. For a non paravian to loose it's feather coating it has to be both big and living in a hot environment. Since Juravenator was so small it was able to keep much of it's feather coating despite the warm environment.

This was discussed in another thread but I agree. The safari T. rex was well sculpted but in an attempt to look modern but they went a little overboard by giving it what looks like pennaceous feathers on the neck and a overly heavy coat of feathers. A am very much looking forward to see how safari and collecta incorporate this latest research into their next Tyrannosaurus. Safari Giganotosaurus does not have visible scales yet it still looks awesome.

As to why the scales are so small I haven't got a clue. Is their any reason they would need to be big? If the purpose of scales are to protect from the sun and against abrasions small scales would still do the job although they may not be as effective against abrasions as large ones.

[stargatedalek]

These animals also lived on the coastline, and potentially even caught prey underwater or swam frequently. It doesn't seem unreasonable to suggest that these cases might be more behavioral than environmental, especially since neither Juravenator or Compsognathus is particularly large.

[WarrenJB]

Actually that brings me to another question that I think I asked before, but why were these so small? If these are actually scales, in what ways would these be beneficial to T. rex at this size?

I might be barking up the wrong tree, using too small a sample size etc., but if they are reticula, they don't seem to get much bigger on modern bird feet, regardless of the size of the bird. Compare chicken and turkey feet to big ratites.
If that was the case, I don't know how analogous avian (avialan?  ) pedal reticula would be to possible tyrannosaurid reticula from other, higher parts of the body. Some foot scales on some large birds are bigger, but they seem to be part of a gradation from tarsal scutes to smaller reticula. (Does that make them smaller scutes, or bigger reticula?) I guess this isn't much of an issue for scales on the flank or tail, even with dispersed feature scales. Would it make a difference if such reticula were derived from hairlike rather than pennaceous feathers, too?

As for the benefit, if the point is to aid heat loss, I guess it's enough to have the feathers out of the way. I can see Neosodon's POV that they might've protected against abrasion, though I wonder how they'd hold up against the kind of force that a Tyrannosaurus rex could bring to bear, even if just accidentally scraping against trees 'n' junk. Though I guess you could say the same for big ratites?

[Neosodon]

As for the benefit, if the point is to aid heat loss, I guess it's enough to have the feathers out of the way. I can see Neosodon's POV that they might've protected against abrasion, though I wonder how they'd hold up against the kind of force that a Tyrannosaurus rex could bring to bear, even if just accidentally scraping against trees 'n' junk. Though I guess you could say the same for big ratites?

I cannot say how effective Tyrannosaurus's mini scales would be if it was accidentally scraping against trees 'n' junk. But there are other uses. Wouldn't they block UV rays and insect bites too?

[HD-man]

Adding to what Dinoguy2 said, Archaeopteryx and Compsognathus are also from the Solnhofen archipelago.  Fossils of Archaeopteryx show it was extensively feathered, with large remiges and rectrices, basically it's feathering is bird-like.  Compsognathus however doesn't have any feathers preserved even though it's from the same place, and actually has scales preserved on its tail similar to Juravenator.

Actually, to quote Naish, "The small bumps reported on the bones of the Canjuers Compsognathus could be just about anything (they remind me of the pathological tubercles present on some Messel bird bones) and cannot be assumed to be integumentary: they're preserved on the bones, which makes it highly doubtful that they're scales" ( http://scienceblogs.com/tetrapodzoology/2009/06/19/limusaurus-is-awesome/#comment-13634 ).

This is a pretty great match for the feathering of Juravenator:

I didn't know that Juravenator's "feathering" was that sparse. Does this mean that Juravenator was scaly w/whiskers like this (as opposed to fluffy like Sinosauropteryx)?:

[Sim]

Actually, to quote Naish, "The small bumps reported on the bones of the Canjuers Compsognathus could be just about anything (they remind me of the pathological tubercles present on some Messel bird bones) and cannot be assumed to be integumentary: they're preserved on the bones, which makes it highly doubtful that they're scales" ( http://scienceblogs.com/tetrapodzoology/2009/06/19/limusaurus-is-awesome/#comment-13634 ).

Well, that's good to be aware of.  Although I'm inclined to think it is actually scales on Compsognathus.  I noticed that comment by Darren Naish is from quite some time ago - 2009.  For a long time now I've seen a number of experts in palaeontology be overly sceptical that skin patches from theropods could show scales, saying it could be bare skin instead and ultimately suggesting the patches should be ignored with regards to feathering.  It's only more recently, such as with these new Tyrannosaurus skin patches that I'm starting to see 'scaly' theropod skin patches being given more fair consideration.  As much as I like Darren Naish, his comment feels like that kind by others I referred to, where 'scaly' skin patches aren't assessed well enough.  He mentioned that since the patches are on the bones of Compsognathus, it makes it highly doubtful they're scales.  However, skin that is considered to show scales is also preserved on bone in Wyrex, and on the Psittacosaurus colour specimen.  So Darren's comment doesn't seem (entirely) up-to-date.

Additionally, Juravenator has scales on the tail and apparently lived in the same environment as Compsognathus and relatively close to it in time.  Juravenator also appears to be closely related to Compsognathus, and the patches from the tail of Compsognathus have been said to be similar to the scales of Juravenator.  With everything considered, to me it seems very likely that the patches from Compsognathus are showing scales, and that if they don't show the natural integument of the animal, Compsognathus would've probably had scales on the tail anyway like in Juravenator.  It would be interesting to see the Compsognathus patches compared with the qualities of scales in that Reptilis blog post you linked to earlier.  It seems there's a number of different papers that consider those tail patches from Compsognathus as showing scales, and none that refute this, so with everything I've said so far in mind, I don't feel Darren's comment is an adequate refutation of scaly skin from the tail of Compsognathus.  In my experience, giving too much importance to Internet comments that aren't based on strong foundation has happened a number of times in the palaeontology community, and it results in being misled and believing something unlikely.

I didn't know that Juravenator's "feathering" was that sparse. Does this mean that Juravenator was scaly w/whiskers like this (as opposed to fluffy like Sinosauropteryx)?:

I also felt what Dinoguy2 said could be interpreted as Juravenator being almost all scaly with only some short feathers on the head, just like that Tyrannosaurus by Peter Schouten.  But since Juravenator is known to have feathers above the sacrum and on dorsal parts of its tail, I interpreted what Dinoguy2 said as meaning the length and fineness of the feathers on that T. rex would be a great match for the feathers of Juravenator.

[HD-man]

It seems there's a number of different papers that consider those tail patches from Compsognathus as showing scales, and none that refute this,

What about Reisdorf & Wuttke 2012 ("in 2012 they were by Achim Reisdorf seen as plaques of adipocere, corpse wax.[8]": https://en.wikipedia.org/wiki/Compsognathus#Integument )? In retrospect, I probably should've started w/that, but the Naish quote came to mind 1st.

But since Juravenator is known to have feathers above the sacrum and on dorsal parts of its tail, I interpreted what Dinoguy2 said as meaning the length and fineness of the feathers on that T. rex would be a great match for the feathers of Juravenator.

I did too. That's why I asked, "Does this mean that Juravenator was scaly w/whiskers like this...?" B-4 then, I had always seen Juravenator depicted as being fluffy w/scales (like this: http://dustdevil.deviantart.com/art/Juravenator-38812964 ). Correct me if I'm wrong, but I'm thinking that Juravenator's sparse "feathering" & basal position would be good evidence for the "Dinosaur Whiskers" hypothesis ( http://phenomena.nationalgeographic.com/2015/03/27/dinosaur-whiskers/ ). I'm also thinking that Juravenator would've looked something like WWD's Ornitholestes:

[Sim]

It seems there's a number of different papers that consider those tail patches from Compsognathus as showing scales, and none that refute this,

What about Reisdorf & Wuttke 2012 ("in 2012 they were by Achim Reisdorf seen as plaques of adipocere, corpse wax.[8]": https://en.wikipedia.org/wiki/Compsognathus#Integument )? In retrospect, I probably should've started w/that, but the Naish quote came to mind 1st.

That appears to refer to scales that had been reported in the abdominal region of the German Compsognathus, not to the reported scales from the tail of the French Compsognathus.  This is the full quote: "Additional scales had in 1901 been reported by Von Huene, in the abdominal region of the German Compsognathus, but Ostrom subsequently disproved this interpretation;[9][17] in 2012 they were by Achim Reisdorf seen as plaques of adipocere, corpse wax.[8]"  So these reported scales from the German specimen had already been shown to not actually be scales by Ostrom.

This is the reference paper for the quote you posted, I've done a few searches and haven't found it saying anything about scales from the French Compsognathus: link

I did too. That's why I asked, "Does this mean that Juravenator was scaly w/whiskers like this...?" B-4 then, I had always seen Juravenator depicted as being fluffy w/scales (like this: http://dustdevil.deviantart.com/art/Juravenator-38812964 ). Correct me if I'm wrong, but I'm thinking that Juravenator's sparse "feathering" & basal position would be good evidence for the "Dinosaur Whiskers" hypothesis ( http://phenomena.nationalgeographic.com/2015/03/27/dinosaur-whiskers/ ). I'm also thinking that Juravenator would've looked something like WWD's Ornitholestes:

Reading that article about the "Dinosaur whiskers hypothesis" actually made me find this hypothesis unconvincing.  I can't imagine feathers starting out as facial bristles.  Also, if non-avian dinosaurs had whisker-like feathers, I would think they'd be bird-like in distribution, not mammal-like as that National Geographic article suggests both in the text and in the image.

I don't think Juravenator offers anything that supports the "Dinosaur whiskers hypothesis" more than another feather evolution hypothesis.  I don't know why Dinoguy2 said Juravenator "was 99% covered in scales", as it appears most of the Juravenator specimen has no integument of any kind preserved - no scales, no feathers, nothing.  Based on the limited integument known for Juravenator, it doesn't seem unlikely that Juravenator only had scales on some areas e.g. the underside of the tail, the feet and legs, the hands, and maybe the snout, with feathers being everywhere else.

I find theropod reconstructions like the WWD Ornitholestes, where the animal is almost entirely scaly, with the only feathers being tufts on the head and arms and sometimes on other small areas, look unpleasant and unnatural.

[HD-man]

Also, if non-avian dinosaurs had whisker-like feathers, I would think they'd be bird-like in distribution, not mammal-like as that National Geographic article suggests both in the text and in the image.

To be fair, basal coelurosaurs probably didn't have beaks like modern birds do. Also, modern birds don't grab w/their arms like basal coelurosaurs probably did.

I don't know why Dinoguy2 said Juravenator "was 99% covered in scales", as it appears most of the Juravenator specimen has no integument of any kind preserved - no scales, no feathers, nothing.

Maybe he meant that 99% of the preserved integument is scaly.

I find theropod reconstructions like the WWD Ornitholestes, where the animal is almost entirely scaly, with the only feathers being tufts on the head and arms and sometimes on other small areas, look unpleasant and unnatural.

I wouldn't call them "tufts" given how sparse the individual feathers are. In the concept art more so than the doc, it looks like WWD's Ornitholestes has a sparse covering of hair-like feathers over the entire upper body w/greater concentrations of longer feathers on certain body parts, similar to the hypothesized distribution of hair on therocephalian therapsids ( https://en.wikipedia.org/wiki/Therapsid#/media/File:Pristeroognathus_DB.jpg ). W/that in mind, I wouldn't call it "unnatural" looking. Weird looking compared to later, more fully-feathered coelurosaurs, yes, but the same can be said about therocephalian therapsids compared to later, more fully-furred therapsids (E.g. Modern mammals).

[Dinoguy2]

Yeah, I meant of preserved integument for Juravenator. The UV study being referenced is the one that revealed it had any feathers, period, isn't it? The first paper reported that feathers were totally absent from the preserved impressions. The feathers we have are from above the hip, but I don't see why a sparse covering couldn't be present across the whole body. I do think it would look more like Schoutan's and Tom's though to be honest. Just a sparse fuzz with the skin texture still visible. Think elephant.

Edit: I just checked the papers, and the wiki article is wrong. The Chiappe paper doesn't report any additional feathers, just the same 1cm wide patch on the tail reported by Tischlinger. The feathers are unbelievably fine/thin and less than 1mm in length. A baby's arm has longer, if not denser, fuzz than the tail of Juravenator. Compared to Juravenator, the tail of Sinosauropteryx looks like a yeti

No matter what was going on with the rest of the body, the Jura feathering was obviously extremely reduced compared to the Yixian "compsognathids" (hahaha! In a few years saying that is going to be like saying "thecodont". There's evidence, mostly unpublished right now, that compsognathids are an artificial group of unrelated young juvenile theropods. So I wouldn't put any stock in phylogeny to help inform their appearance).

[Sim]

Also, if non-avian dinosaurs had whisker-like feathers, I would think they'd be bird-like in distribution, not mammal-like as that National Geographic article suggests both in the text and in the image.

To be fair, basal coelurosaurs probably didn't have beaks like modern birds do. Also, modern birds don't grab w/their arms like basal coelurosaurs probably did.

Yes, I don't think a coelurosaur would have a beak if there's nothing to support it having one, e.g. the appropriate bone structure.  However, mammal faces are pretty specialised as well, and they seem to be a worse point of comparison than birds for beakless coelurosaurs.  Most mammals have a specialised fleshy nose, the rhinarium.  If my understanding is correct, mammal facial tissue can also allow it to be moved, such as to pull back lips and expose teeth, which is impossible for reptiles and birds to do as they don't have this tissue.  Maybe mammal whiskers are related to the rhinarium, or other mammalian facial features?  I get the impression the snouts of beakless theropods wouldn't tend to be almost completely covered in feathers in the way that mammal snouts tend to be almost completely covered in hair.  For a start, that integument arrangement in mammals seems to be affected by the rhinarium.  There's also specimens of Sinosauropteryx and Sinornithosaurus that show their head feather covering stopping a large distance away from the nostrils.  I get the feeling that in beakless theropods, a significant amount of the snout would be covered in either scales or hardened skin.

Even if 'basal' coelurosaurs grabbed with their arms, it should be considered these coelurosaurs are bipedal and incapable of having pronated hands (like birds), unlike mammals which mostly are quadrupedal.  So even if these coelurosaurs could use their arms to grab unlike birds, they weren't using their arms in the same way as mammals either, so I see no reason for a mammal-like whisker arrangement on the arms, especially not on the dinosaurs the National Geographic article specifically suggested this for, Velociraptor (which had large wing feathers on its arms) and Tyrannosaurus (which had tiny arms).  I think a much better point for comparison for sensory integument in the arms/legs of non-avian coelurosaurs would be found in birds - the filoplumes of owl feet.

Other things that are relevant are the lack of relation mammals have to coelurosaurs or any dinosaurs, while birds actually are coelurosaurs, and as the National Geographic article said, no signs of "whiskers" are known from non-avian dinosaurs, including specimens that have extensive feathering preserved.  Additionally, a mammal-like "whisker" distribution is not present in both extant birds and extant reptiles.  I find dinosaur reconstructions that look mammal-like look very unpleasant and unconvincing.

I find theropod reconstructions like the WWD Ornitholestes, where the animal is almost entirely scaly, with the only feathers being tufts on the head and arms and sometimes on other small areas, look unpleasant and unnatural.

I wouldn't call them "tufts" given how sparse the individual feathers are. In the concept art more so than the doc, it looks like WWD's Ornitholestes has a sparse covering of hair-like feathers over the entire upper body w/greater concentrations of longer feathers on certain body parts, similar to the hypothesized distribution of hair on therocephalian therapsids ( https://en.wikipedia.org/wiki/Therapsid#/media/File:Pristeroognathus_DB.jpg ). W/that in mind, I wouldn't call it "unnatural" looking. Weird looking compared to later, more fully-feathered coelurosaurs, yes, but the same can be said about therocephalian therapsids compared to later, more fully-furred therapsids (E.g. Modern mammals).

I'm not seeing anything on the WWD Ornitholestes that could be interpreted as a sparse covering of feathers over the entire upper body.  Even in the concept art, the distribution of the green and white colouration appears to preclude such a feather covering.  My point is that almost entirely scaly dinosaur reconstructions which only have limited feathering restricted to small areas on the animal, most often the head and/or arms, are not rare even though there is nothing to support such a way of being feathered, and I find they tend to just look like a scaly dinosaur with some feathers stuck on.  I think it's possible to make minimally feathered reconstructions that don't look unrealistic, for example the most recent Tyrannosaurus painting by Mark Witton in his recent blog post, but I almost always find minimally feathered reconstructions look like feathers have been forced onto a scaly animal in an unnatural-looking way.  Although I find these reconstructions still look better than dinosaur reconstructions with mammal-like whiskers/noses/facial integument arrangement.  Hehe!

No matter what was going on with the rest of the body, the Jura feathering was obviously extremely reduced compared to the Yixian "compsognathids" (hahaha! In a few years saying that is going to be like saying "thecodont". There's evidence, mostly unpublished right now, that compsognathids are an artificial group of unrelated young juvenile theropods. So I wouldn't put any stock in phylogeny to help inform their appearance).

Or perhaps, the Juravenator feathering could represent an early feathering state that was replaced by more extensive feathering in more 'derived' coelurosaurs?  I guess it could be a hint that feathers were not widespread among non-coelurosaur theropods.

As for compsognathids being an artificial group...  Since some ideas in palaeontology that have actually been published have been very unconvincing and/or controversial, I feel it would be premature for me to believe compsognathids are an artificial group, to say the least.  If something is published on this topic, I'll be interested in reading it.  I find it very difficult to believe the argument mentioned above that no compsognathids form a natural group.  One specimen of Sinosauropteryx is thought to be an adult that has its own unlaid eggs preserved in its abdomen.  Sinocalliopteryx preserves what appears to be the same kind of tail feathering with alternating colour bands as Sinosauropteryx, which I think makes it likely these two are closely related.

[stargatedalek]

All of this really depends on where one chooses to draw the line on what constitutes a "beak". Most expert paleontologists (which is to say the two maybe three who've ever commented on it) agree that a "beak" is "a keratinous structure covering an animals jaws and gums" instead of "a keratinous structure on the animals face or jaws" but as far as the vernacular is concerned such a distinction is only going to serve to confuse people and lead to misconceptions. When someone says "Ichthyornis didn't have a beak" (because the gums weren't covered by keratin) that's going to lead people to assume it had a lizard-like or even "stereotypical-theropod-like" (overhanging teeth, etc.) face when it actually had a structure of keratin on its face that would have looked near identical to those of modern birds.

And then you have groups like cephalopods, puffer fish, and turtles, which are typically referred to as possessing "beaks" but wouldn't fit the definition either if Ichthyornis doesn't.

A structure of hardened skin or keratin on the animals face is more or less plausible on any dinosaur. We see a very toned down version of this on crocodilians, and the logical extreme of this adaptation in birds.

[Neosodon]

I don't think the WWD Ornitholestes is supposed to have feathers. I've always thought they were quills. I don't see why people would think they are weird. I've never heard anyone complain about how they look weird on Psitacosaurus.

[stargatedalek]

I don't think the WWD Ornitholestes is supposed to have feathers. I've always thought they were quills. I don't see why people would think they are weird. I've never heard anyone complain about how they look weird on Psitacosaurus.

Ornitholestes is supposed to have feathers, not quills, so they absolutely do look weird and unnatural because they ARE unnatural/incorrect. Fairly certain in interviews it's been said they were feathers.

[Neosodon]

I don't think the WWD Ornitholestes is supposed to have feathers. I've always thought they were quills. I don't see why people would think they are weird. I've never heard anyone complain about how they look weird on Psitacosaurus.

Ornitholestes is supposed to have feathers, not quills, so they absolutely do look weird and unnatural because they ARE unnatural/incorrect. Fairly certain in interviews it's been said they were feathers.

No one really knows if it had feathers or quills. But for the movie it looks like they were trying to portray quills.