|Allosaurus credit Shizoform CC generic
For those who, despite these seemingly "vanilla" attributes for Allosaurus, intuitively felt that there was much more to this animal than meets the eye - your hunches might be more true than you even imagined.
In this post I aim to present a startling new theory on Allosaurus feeding technique that resolves these anatomical and ecological incongruities and offers utility in looking at other ziphodont theropod predators - to a greater or lesser extent - in terms of feeding adaptation. I will also in the process describe a unique and as yet undescribed feeding adaptation in several lineages of flesh rendering birds.
As background readers should be well versed in the paper by Bakker, 1998. Brontosaur Killers: Late Jurassic Allosaurids as Sabre-Tooth Cat Analogues. GAIA, December, 1998. Pp 145-158 available online here. and the more recent, but largely congruent study Snively, Cotton, Ridgely & Witmer 2013 Multibody Dynamics Model of Head and Neck Function in Allosaurus (Dinosauria, Theropoda). Paleontologica Electronica May 2013 online here.
In fairness it should also be mentioned that the finite element analysis of Allosaurus by Rayfield et al. supported a "slash and tear" attack (2002) while Frazzeta & Kardong come out against the "chop and slash" hatchet attack. While I lean towards the neck driven attack in Allosaurus this style is not a prerequisite for what I am going to outline. My theory will focus on the step between the initial bite/hack/purchase and the head pulling away with a bite of food. Mine is the intermediary step.
|Charles R. Knight. wiki public domain
What also should get your attention is the intricate social hierarchies between not just species of scavenging birds but between individuals of the same species. Note the lappet-faced vulture (Torgos tracheliotos) moving in and dominating the impala just before the leopard seizes it. Also note that many vultures are more concerned with fighting and establishing dominance amongst themselves rather than, you know, actually getting to the rapidly dwindling carcass. Establishing who is who in the pecking order seems to be very important to these animals. Of course looking at the intricate feeding guilds, carcass location tactics, and niche partitioning in these extant flesh rendering theropods are other fascinating avenues of discussion beyond the scope of this post. My main point being vultures are intelligent, social, and intensely competitive. Inferior and cowardly they are not.
I really love this video too. Pay special attention to the shifting eyes of the vultures as they sense the jackal moving off. And then just the mosh pit of activity as they rush in. And pay special attention to the lappet-faced vulture and how it struts it's stuff and just establishes dominion over that carcass.
And now on to the feeding adaptations of vultures (both old & new world). What I am about to introduce - and what forms the crux of a large part of my argument - is that vultures have a peculiar and, to my knowledge, as yet undescribed (anecdotally or in the scientific literature) feeding mechanism. And it is this feeding mechanism - which as I will show you is and has been in plain sight all along - that separates vultures from other birds of prey with regards to efficiency in harvesting large carcasses. And pinpoints them as more useful analogues to Allosaurus, and many other ziphodont theropods, in feeding mechanics. And, again I believe it has likely been the social stigma of "lowly scavenging carrion birds that has kept this feeding method invisible to us while it has always been in plain site.
Let's start with a youtube video - plenty more to come - that I think most succinctly captures the vulture feeding mechanism. I have seen the whole video before, it was I believe a show from the Discovery channel looking at the evolution over time of a large carcass (btw it was not a bison but I think a hippo or some other large African mammal). Be advised that the action happens very fast - you will find yourself probably watching some of these videos several time.
On the velocity of action:
"Look at the speed - it looks like it's insanely speeded up but it isn't is it? It's for real"
On the vulture feeding technique:
"Fast nibbly action rather than this sort of hooking in and pulling huge hunks out as would an eagle"
That "fast nibbly action" is what we will be looking at for a bit. And, as the vulture researcher posits, this style of feeding is different than the typical "hooking in and pulling" action of eagles, hawks, falcons and other more predatorial birds of prey. You can go check out other birds of prey and see the difference but I have a video below of some nice close ups of bald eagles feeding on discarded fish that - especially played after the above vulture video - highlights the differences.
Ok so what if all those "fast nibbly parts" are just the vulture swallowing little bits? This idea is negated by the fact that in the vulture videos they tilt their heads back to make an obvious swallowing motion when they are doing it.
It should also be stated that vultures plainly use the "hook and pull technique" like a falcon or eagle. But what should not go unnoticed is that the "fast nibbly part" almost always occurs right before the "hook and pull" part. It is almost as if the vulture is softening or chewing up the meat to better facilitate breakage when the "hook and pull" part comes. Maybe go watch the video above again "Jackal Vs. Vulture" and go to 1:40 where the Lappet-faced vulture starts feeding and you can see that it follows the pattern of "fast nibbly part" before "hook and pull".
And this feeding technique is not just found in old world vultures (Accipitridae) as depicted above but also it is characteristic of New World vultures - which represent a different lineage family (Cathartidae). Whatever is going on here is so efficient it evolved twice. You will see a convergence in feeding below in this video depicting California condors (Gymnogyps californianus) feeding on a deer carcass.
This technique is also evident in black vultures (Coragyps atratus) which starts at 5:30 although the host seems like an interesting fella too.
And here is another with loads of clear shots of the "fast nibbly parts".
And wait, if that's not enough to convince you of the utility of this feeding method in birds that render flesh off of large animals, giant petrels, both the northern/southern (Macronectes giganteus & halli family Procellariidae) use this same feeding method. You can see it to good effect in the video below of giant petrels as well as neat display and fighting behavior. So we have a remarkable case of convergence in that the most dominant large carcass rendering of the New and Old World, and southern/Antarctic oceans all have a similar style of eating.
However the giant petrels take this "fast nibbly part" a step past what vultures routinely do with dead bodies and use it on live animals. Below is the video that can give you nightmares - you have been warned. But if you sit through it (and I have several times) you will see that before the petrel literally scalps the chinstrap penguin it gives it several "fast nibbly parts" that likely helped facilitate the "hook in and pull" part pulling off a plug of skin and meat.
And, based on this video below of albatross feeding on whale blubber, it looks like other tubenosed birds utilize this feeding technique as well.
All right so what is going on here with these "fast-nibbly parts"? Let's look inside the mouth. On the roof of the mouth of birds they have something called "choanal papillae" which can take various forms but you might better know it as pseudo teeth. You can see them quite obviously in the penguin below.
|penguin with choanal papillae. via Discovery
Above is a frequently memed image of a king vulture and you can see the fine layers of choanal papillae on the roof of the mouth. But below is an even more telling picture of a turkey vulture that suggests a method as to how this whole operation works.
What you should note is that not only are there small serrated papillae lining the roof of the mouth, but the tongue too. Credit Williston Conservation Bird Trust Blog.
Now this is all common knowledge that vulture mouths have serrated/barbed tongues and papillae lined throats. But during the "fast nibbly parts" - a term I will replace from here on out as choanal grinding - I have outlined a hypothesis that best explains what is going on.
After a parcel of meat or skin is clenched in the beak the tongue rapidly retracts and grinds the food item against the choanal papillae. This action, combined with the rapid movements of the whole neck further enacting different force vectors, serves to mince and grind the meat down ultimately diminishing it's structural integrity. And then, after the tissue has been been broken down a bit, comes the classic "hook in and pull" routine which dislodges the parcel of food which is then swallowed.
Again, to the best of my knowledge this has never been suggested or explained as such. I could be wrong and there is an open comments section below.
So if we look at this "choanal grinding" it is happening almost imperceptibly fast and it provides an elegant explanation for why birds that routinely render carcasses larger than themselves all share this characteristic feeding behavior while the tug and pull "raptors" such as hawks, eagles, and falcons - which more commonly feed on items smaller than themselves - do not. Further testing could include ultra slow motion film of these birds feeding and/or in some type of controlled setting.
But while searching for images of vulture tongues - surprisingly there is an extreme paucity of such imagery on the net - I came across a bit of independent confirmation of my hypothesis. What I came across was an image, via the Vulture Conservation Foundation, of a Griffon with tongue protruding from the neck observed in France!! Evidently, probably from some anthropogenic cause, this griffon got a hole in the neck from which the tongue just kind of lopped out. But wait, it gets bettor (or worse).
|credit Francesco Panuello via vulture conservation
Vulture News 64 July 2013. An Eurasian Griffon Gyps fulvus disadvantaged for feeding. Written by Alvaro Camina and Luis Miguel Guerroro, this time seen in northern Spain. And here it gets really interesting:
"The bird had a perforation just on the throat that left the tongue outside the skin, and the tongue moved forward and backward while the bird was eating."
Now this is astounding. The tongue in this bird - which remember is not even in the oral cavity where food is being processed - still flicked forward and backward when it was feeding. This suggests that the tongue rapidly moving back and forth is involuntary and intrinsic to their feeding adaptation!! Drop the mic.
Interestingly of the picture provided in the paper of this vulture feeding it is quite clearly in "hook and pull" mode on a rather small morsel of pig.
Time to segue back into the Allosaurus question. In light of that last bit, I am going to go with birds that actually render the flesh of large tetrapods - New & Old World Vultures, Giant Petrels - as a superior model than falcons, eagles, hawks that typically on average eat animals equal to or smaller than themselves (don't light up the comments with eagles attacking deer/wolves, I mean on average). Am I suggesting that Allosaurus and other theropods had gnarly choanal papillae and serrated tongues? No, quite the contrary in fact because they had a superior set of cutlery already in place - their rows of serrated teeth.
As I mentioned at the start of the post it will prove useful to be familiar with both the Bakker, 1998 paper and Snively et al., 2013. Bakker, through comparative anatomy and Snively et al. through multibody dynamics finite element analysis converge on a similar anatomical adaptation in these animals. Allosaurus had an exceptionally powerful and flexible neck which was especially augmented for rapid dorsoventral movements. The rigorous analysis of Snively bolstered Bakker's theory of rapid neck strikes, coupled with widely gaping jaws to enact "brontophagy".
|Rayfield et al. 2001
Additionally Snively et al. suggest that this astounding musculature that facilitated such forceful hammer blows was then used when feeding to pull the head back forcefully - like falcons and other birds of prey "hook in and pull"- the flesh. And I have no problem with both the hammer saw blow from Bakker or the falcon hook in and pull technique. What I am suggesting is that a step is missing between both the initial strike and the pull back. And that step is the allosaurian equivalent to the just discussed choanal grinding in extant carcass rendering birds. But instead of barbed tongues and choanal papillae doing the cutting it was serrated teeth.
After biting into a large food item (alive or dead) then the theropod bonesaw would come into play. Rapid and forceful back and forth pumpings of the neck in the dorsoventral plane - as suggested by the work of Bakker, Snively et al., and Rayfield - then allowed a uniquely devastating and vicious sawing to commence. Now the tissue in the mouth will be - partially by it's own inertia - be sawed back and forth over both the fore and aft serrated margins of the teeth when the pumpings occur. This action - most effective with a large individual piece or a plug from a large carcass to allow more resistant inertia- will effectively negate the use of strong jaw closing muscles - which Allosaurus was relatively impoverished in - and outsource the driving musculature to the neck. A rather weak bite is actually more efficient because that will better allow the meat to slide back and forth over the teeth sawing into it. And because both sides of the serrated tooth are equally in use the cutting edge of the teeth will be better preserved.
Based on a number of factors this bonesaw shimmy as I will call it from here on out was utilized with a variety of techniques. What is probable is that the pumping bonesaw shimmy worked best on big hunks of meat and smaller bits were swallowed or fed on with the "hook in and pull technique". As in the vultures and petrels (you probably want to watch some more videos at this point), this action was occurring almost imperceptibly fast and were interspersed with various other actions, letting go to readjust, pulling back. Probably not as fast as in the birds because Allosaurus is bigger - relative muscle strength diminished with size etc etc - but still looked freakishly sped up. As in the "fast nibbly parts" the jaw is - for the briefest millisecond - opening up to allow the parcel of food to be moved in the mouth and scraped against the tongue/papillae as can be seen in this photo right here in a lappet-faced vulture. Seeing a mob of Allosaurus chowing down on a Camarasaurus carcass (or eating it alive) must have looked diabolically horrific on a disturbing scale. Take that Petrel Vs Penguin video and crank it up to 11.
|credit Kevmin fossil C. carcharias Miocene Atacma desert Chile. C.C.
Let me prime your mind a bit with some youtube clips of serrated toothed sharks feeding on whale carcasses. You will notice that as the sharks shake their heads side to side to saw off a hunk of blubber this action allows both sides of the serrated tooth to work over the tissue just as in the scenario I painted above.
Again, these rapid dorsocentral bonesaw shimmies would take advantage of both the fore and aft serrated cutting edges on the Allosaurus tooth for maximum sawing efficiency at minimal damage to tooth. Since strong bite force is not required crown breakage is curtailed and because serrations on both sides of the tooth are utilized the stress is spread out evenly across the teeth and serrations.
|Allosaurus SDNHM public domain
As I alluded to in my previous post Death Comes Ripping the recent work by Brink et al. suggests a uniquely efficient and strong tooth in ziphodont theropods which may have expanded their niche into true bone utilization. But not via crushing as I argue (with the exception of tyrannosaurids) we must disabuse ourselves of the hyena paradigm of bone consumption. Instead the bonesaw shimmy worked to effectively slice not only meat but cartilage and bone.
Here is a bit of rough sketch to give you a visual. Again, you have to imagine this happening at a very rapid pace - interspersed with the jaws opening a bit occasionally to allow movement of the bone.
Now we get back to the ecological conundrum of Allosaurus' dominance in the Morrison theropod predatory guild. It was uniquely equipped among it's contemporaries for bone slicing. Ceratosaurus had very long, narrow blade like teeth and a relatively straight neck not augmented for this type of feeding action. Although Torvosaurus was probably capable of some bone consumption it's massive bulk limited it ecologically speaking to a more specialist niche. Torvosaurus might well have been the lappet-faced vulture of the Morrison. Rare - but dominant - and it's big skull and teeth, large powerful and lithe body may have better allowed it the ability to open up and dismember exceptionally large sauropod carcasses - which allowed smaller theropods like Allosaurus better access. As I have been preaching for awhile we do not always have to look at dinosaur coexistence through the lens of competition but also through commensal/facultative relationships. So Allosaurs could exploit all parts of a large carcass efficiently. This expanded niche, with a potential mobbing behavior, and moderate size that allowed flexibility in food choice gave Allosaurus the edge.
So where is the hard evidence of Allosaurus accessing all these bones? The Morrison is so well trodden by collectors we should have bones sliced by Allosaurus lying around everywhere. Well, it turns out we already have the evidence. For whatever reasons, it never really made the news.
Now in my last post I discussed the Chure et al (1996) paper Prey bone utilization by predatory dinosaurs in the late Jurassic of North America in which the tip of an Allosaurus pubic bone was basically cleaved right off. The pubic bone - especially the foot - is not an insignificant bone to cut through. Especially if this was just incidental contact as the author's suggest. Now we don't know who was the perp - the authors suggest Ceratosaurus or Torvosaurus - but this piece of evidence should not go overlooked because we know at least some type of theropod cleaved off that piece of bone. Given the numerical dominance of Allosaurus in the Morrison chances are it was Allosaurus.
An additional layer of evidence for bone utilization by some type of theropod in the Morrison, also detailed in my last post, is provided in Karen Chin's seldom discussed paper Exploited twice: bored bone in a theropod coprolite from the Jurassic formation of Utah (2008). Chin mention not one but two likely theropod coprolites full of chunks and shards of bone (the Morrison lacked large crocodilians) one of which shows substantial amount of insect borings of the bone enmeshed within the coprolite. What is extremely interesting is that Chin can not resolve whether the insect bored into the bone first - which would suggest that the theropod in question consumed a largely defleshed bone - or if the insect bored into the bone after the theropod passed it. Chin interprets the bone as incidental bone taken in with carcass consumption, which given the generally dismissive view of bone consumption in theropods seems like the most parsimonious conclusion to make. But I would beg to differ. Allosaurus has teeth in the front of it's mouth that are D-shaped in cross section so, like tyrannosaurids, it could if it wanted to carefully pick meat off a skeleton without much incidental bone contact. If this was indeed an Allosaurus coprolite (statistically speaking that is the most likely culprit) and if Allosaurus was bone slicing using the technique as described we should expect several characteristic of the bone preserved in coprolite. These predictions are that we should see lots of small splinters/shards of bone from the sawing action together with larger chucks of bone swallowed whole. And, indeed, when we look at the coprolite - in real life view and microscopically - from Chin's study that is the pattern we see. Prediction met.
Wait there is more. Sometimes a paper - or maybe just a small bulletin - is published that falls through the cracks because it is made by people not specialized is said discipline or just is so obscure that no one more heavily invested in the issues at hand takes notice. I have found such a piece.
Although as far as I know the authors are paleontologists they are not dinosaur guys so they may have been unaware of the significance of this specimen. I tried to contact them as well but the lead author Dwayne Stone is deceased and I can't find an active email for the second author Edward Crisp although I think he is still active. It was published as an abstract at the 2000 GSA summit of Reno, Nevada. It's on the web here and I transcribed it below (screen shot did not fit here).
A LARGE MEAT-EATING DINOSAUR COPROLITE FROM THE JURASSIC MORRISON FORMATION OF UTAH
Author(s): STONE, Dwayne D., Dept. of Geology, Marietta College, Marietta, OH 45750; CRISP, Edward L., Geology Dept., West Virginia University at Parkersburg, Parkersburg, WV 26101, firstname.lastname@example.org; BISHOP, John R., Rt. 2 Box 137, Ravenswood, WV 26164
Keywords: Coprolite, Jurassic, Morrison Formation, Allosaurus
A theropod dinosaur coprolite has been excavated in Emery County, Utah from a red-brown mudstone of the Upper Jurassic Brushy Basin Member of the Morrison Formation. The coprolite is nearly complete and is divisible into two parts, the main mass and a dribbling zone. The well-indurated main mass, which tapers at both ends, is 1.52 m in length, 0.457 m in maximum width, and 10.2 cm thick. The 1.52 m long dribbling zone consists of small isolated coprolites that curve towards the end of the dribbles. The coprolite geometry is interpreted to indicate that the main mass was defecated first, then the animal walked forward to release smaller amounts of feces. The coprolite consists of dark gray bone fragments, ranging from pebble to sand size, in a red-brown matrix. Bone fragments represent about 50% of the mass and have broken, jagged ends, possibly indicating breakage by the biting action of a carnivore. XRD analysis of the bone fragments and matrix reveal that both are primarily composed of carbonate fluorapatite. The large size of the coprolite and its geometry and stratigraphic location indicate that it represents fecal droppings from a large Allosaurus. Further support for this hypothesis is the fact that a broken distal end of an Allosaurus tooth was found within the coprolite, indicating that during mastication a tooth was broken and ingested. The carnivore did not grab a portion of flesh and bone from a prey animal and then swallow it whole. Instead the eater masticated its meal and broke the bones into smaller portions. This is the largest and oldest theropod dinosaur coprolite known. Work is continuing on a second theropod coprolite higher in the section at this location and pieces of theropod coprolites from two additional localities have been identified. Morrison Formation theropod coprolites are no longer considered to be absent or scarce and future searching should reveal additional large specimens.
Ok so first things first this theropod turd blows that Tyrannosaurus turd out of the water in terms of size. This one is over 1.5 meters long and (sorry king) the turd documented in Chin's paper (1998) was a paltry .46 meters. So on the Couric scale of crap size Tyrannosaurus needs to take a seat cuz this Morrison theropod has got it beat. Not only was this poop larger - the pooper had a more fibrous diet as well - with about 50% bone mass while Chin estimates about 30-50% bone mass in the T-rex coprolite. And then it get's even better:
"The coprolite consists of dark gray bone fragments, ranging from pebble to sand size, in a red-brown matrix. Bone fragments represent about 50% of the mass and have broken, jagged ends, possibly indicating breakage by the biting action of a carnivore."
Again, as in the Chin (2008) paper, the prediction is met to have a mix of small bone chips - further embellished by the notation of "broken, jagged ends: i.e. the bone was not swallowed whole but was indeed processed.
And finally the clincher:
"The large size of the coprolite and its geometry and stratigraphic location indicate that it represents fecal droppings from a large Allosaurus. Further support for this hypothesis is the fact that a broken distal end of an Allosaurus tooth was found within the coprolite, indicating that during mastication a tooth was broken and ingested."
What is the quote, dogma can blind us to the truth? Sorry tyranno-enthusiasts they got some new competition for bone consumption in the Mesozoic. And - besides my other arguments - it just makes more sense ecologically for bone consumption to have evolved in theropoda before the tyrannosaurids. Nature abhors a vacuum.
Although this post dealt specifically with Allosaurus - there is the most data on it for starters - there is much room to investigate how alternative "bonesaw" cutting techniques evolved in different theropod clades. Allosaurus was uniquely equipped for powerful dorso-ventral neck movements but that does not preclude other relatively long necked, ziphodont theropods - certain dromaeosaurids, coelophysids off the top of my head - from engaging in a similar technique. Other lesser known Jurassic allosaurids/carnosaurs may have evolved similar mechanisms to saw. Abelisaurs and tyrannosaurs likely fell more towards the grab and pull spectrum and were doing something a little more power crunch based as opposed to slicing and dicing. There is lots to explore.
|Tsaagan Giant Petrel Style credit Markus Buhler. Bestiarius on deviantart
And now - because I think I threw a lot of stuff at you and people will need time to digest that a bit. No doubt this theory will change and evolve over time as it is refined. But for now to put you in my mind's eye, I want to go into a little speculative story telling. A Morrison tail of life, death, and cutlery in action...
Mobile Sauropod Body Processing Units of the Morrison
If the old bull Camarasaurus grandis had the emotional intelligence to develop feelings of gratitude, it would feel especially thankful to the 10 meter Torvosaurus tanneri bearing down on it's neck with jaws and claws granting it a relatively quick death. This was a far cry from the several days and nights of torturous harassment by a mob of 9 adult Allosaurus fragilis not to mention the several other dozen allosaurs arranged in assorted size classes tagging along. It was the old bull's 27th year and it suffered several broken foot bones jostling with other males. And now, during the height of the dry season migration to the floodplain seepage wetland, the ever present and ever vigilant allosaurs were quick to locate and capitalize on this hobbled sauropod. The allosaurs concentrated on biting at the cloacal region, caudemofemoralis muscle, and back of the lower legs. While the Camarasaurus had backed into dense vegetation to thwart such attacks before - exposed on the dry vernal ponds and under the pressure to get to water it had suffered debilitating blows. Attacks came almost non-stop from the hatchet head strikes followed by rapid fire bonesaw raking - the disgusting bonesaw shimmy - severing muscles and partially disembowling the 20 ton animal. But even with this trauma the Camarasaurus did not die. Sauropods in general were hard to kill having abundant air sacs throughout their skeleton able to keep the ATP machinery firing past the point where a mammal would be able to survive. But the allosaurs were persistent. They sensed the sauropods strength flagging and the foot and tail strikes no longer packed the punch they did earlier. So when the Torvosaurus arrived it was able to relatively easily dispatch the giant as it was being eaten alive.
Torvosaurus and Allosaurus have a complicated relationship. The adults of both species routinely kill and eat each other's young. But the adults are largely copesetic. Adult and near adult torvosaurs usually "adopt" their own mob of allosaurs. The allosaurs do the leg work of finding, harassing, and weakening prey. The torvosaurus usually comes in to do the coup de grace for large prey items that have been weakened by the allosaurs. But the allosaurs benefit by the torvosaurs presence at the carcass. Their absolutely larger size, stronger jaws, and bigger teeth allow more efficient dismembering of the carcass than the allosaurs are capable of. Using the torque in their large, flexible bodies and tails the torvosaurs rip femurs from hip sockets like macabre wrestlers. But torvosaurs are fiercely possessive of their mob of allosaurs and will not tolerate other torvosaurs trying to move in. So, paradoxically the adults of both species tend to tolerate each other - and often feed together on large carcasses less than a meter apart. This does not imply that they do not eat each other if wounded or deceased.
Once the feast begins it is a blur of activity. Although the torvosaur is easily twice the weight of the allosaur they both feed side by side but use different methods. The torvosaur more dependent on it's large tooth studded jaws and strong body to leverage large pieces swallowed whole. It moves slowly and deliberately. The torvosaur shows a strange penchant for swallowing tough, gristly mats of skin and ligament while the allosaurs concentrate on the flesh. But the allosaurs appear to be moving at 2x the speed. Their necks darting about at all angles followed by almost imperceptibly fast sawing motions of the head carving up meat, skin, sinew, and bone. While not challenging the torvosaur the allosaurs are constantly battling amongst themselves in a never ending quest for dominance and establishment of pecking order.
|Ceratosaurus wikicredit Richie D.
And finally after the adult allosaurs, torvosaurus, and ceratosaurus have fed move in the hordes of various age classes of allosaur. They loosely follow, but segregate themselves from the adult allosaur mob. You see, even though the allosaurs are intensely social, they are also intensely
|Juvenile Allosaurus and one juvenile Torvosaurus feast on Camarasaurus. credit Duane Nash
|Tanycolagreus topwilsoni restored skull cast credit Daderot. wiki
Finally after all the flesh, skin, fat, and marrow rich bones have been harvested and the theropods departed, a colony of specialized bone consuming termites sets up shop on the site of the remaining bones to complete the wholesale recycling of the sauropod into the ecosystem.
Brink, K.S. et. al. (2015) Developmental and evolutionary novelty in the serrated teeth of theropod dinosaurs. Scientific Reports 5, article no. 12338, July 2015
Bakker, 1998. Brontosaur Killers: Late Jurassic Allosaurids as Sabre-Tooth Cat Analogues. GAIA, December, 1998. Pp 145-158.
Camina, Alvaro & Guerroro, Luis Miguel 2013 . An Eurasian Griffon Gyps fulvus disadvantaged for feeding. Vulture News 64 July 2013
Chin, K. et. al. (1998) A king-sized theropod coprolite Nature, 393
Chin, K. et. al. (2008) Exploited twice: bored bone in a theropod coprolite from the Jurassic Morrison formation of Utah, U.S.A. Sediment-Organism Interactions: A Multifaceted Ichnology SEPM special publications No. 88
Chure, Daniel J. et. al.(1996) Prey bone utilization by predatory dinosaurs in the Late Jurassic of North America, with comments on prey bone use by dinosaurs throughout the Mesozoic. December 1996, GAIA
Hone, D.W.E. et. al. (2012) Pterosaurs as a food source for small dromaeosaurs. Paleogeography, Paleoclimatology, Paleoecology
Hone, D., & Rauhut (2009) Feeding Behavior and bone utilization by theropod dinosaurs Lethaia DOI
Stone, Dwayne D., Crisp, Edward L., Bishop, John R. (2000). A large meat-eating dinosaur coprolite from the Jurassic Morrison formation of Utah. Abs No. 50526. 2000 GSA Annual Meeting - Reno, Nevada
Snively, Cotton, Ridgely & Witmer 2013 Multibody Dynamics Model of Head and Neck Function in Allosaurus (Dinosauria, Theropoda). Paleontologica Electronica May 2013