Functional Morphology of Animal Feeding Apparata

[theropod]

This is a place to discuss and compare the form and function of feeding apparata among vertebrates (and invertebrates if you want, but I've got the feeling there won't be many people posting on them). Not strictly just for carnivores, but I suspect that will be the main subject here.

E.g. teeth and bite forces, cranial mechanics, neck musculature etc.

Visual comparisons relevant to the subject are welcome!

Index:

Slicers (in general):

• theworldofanimals.proboards.com/post/2803/thread
• theworldofanimals.proboards.com/post/1025/thread
• theworldofanimals.proboards.com/post/1366/thread
• theworldofanimals.proboards.com/post/26775/thread

Theropoda:

•General:

• theworldofanimals.proboards.com/post/22638/thread
• theworldofanimals.proboards.com/post/23923/thread
• theworldofanimals.proboards.com/post/24933/thread
• theworldofanimals.proboards.com/post/25072/thread

•Allosaurus/Allosauridae:

• theworldofanimals.proboards.com/post/1011/thread
• theworldofanimals.proboards.com/post/1347/thread
• theworldofanimals.proboards.com/post/25071/thread

•Carnotaurus:

• theworldofanimals.proboards.com/post/17764/thread

•Ceratosaurus:

• theworldofanimals.proboards.com/post/1347/thread

•Giganotosaurus

• theworldofanimals.proboards.com/post/26769/thread

•Majungasaurus

• theworldofanimals.proboards.com/post/26005/thread

•Dromaeosauridae:

• theworldofanimals.proboards.com/post/49846/thread

•Phorusrhacidae:

• theworldofanimals.proboards.com/post/25066/thread

•Sinosaurus:

• theworldofanimals.proboards.com/post/26332/thread

•Spinosaurus/Spinosauridae:

• theworldofanimals.proboards.com/post/2978/thread

•Tyrannosaurus:

• theworldofanimals.proboards.com/post/1347/thread
• theworldofanimals.proboards.com/post/1806/thread
• theworldofanimals.proboards.com/post/7311/thread
• theworldofanimals.proboards.com/post/4764/thread
• theworldofanimals.proboards.com/post/15397/thread
• theworldofanimals.proboards.com/post/26769/thread
• theworldofanimals.proboards.com/post/34534/thread
• theworldofanimals.proboards.com/post/46580/thread

Other Dinosauria:

Triceratops

• theworldofanimals.proboards.com/post/22798/thread

Pterosauria:

•Azhdarchidae:

• theworldofanimals.proboards.com/post/15397/thread
• theworldofanimals.proboards.com/post/29494/thread

Non-mammalian Synapsida:

•Lycaenops:

• theworldofanimals.proboards.com/post/3989/thread

Carnivora + Creodonta:

•Hyaenodon:

• theworldofanimals.proboards.com/post/4633/thread

•Smilodon:

• theworldofanimals.proboards.com/post/30012/thread
• theworldofanimals.proboards.com/post/31063/thread
• theworldofanimals.proboards.com/post/46078/thread

•Felidae (general):

• theworldofanimals.proboards.com/post/43449/thread

•Jaguar:

• theworldofanimals.proboards.com/post/31241/thread

Other mammalia:

•Thylacoleo

• theworldofanimals.proboards.com/post/25072/thread

•Thylacinus

• theworldofanimals.proboards.com/post/33571/thread

Crocodylomorpha:

•Dakosaurus:

• theworldofanimals.proboards.com/post/2662/thread

•Purussaurus:

• theworldofanimals.proboards.com/post/51159/thread

Sauropterygia:

•Pliosaurus:

• theworldofanimals.proboards.com/post/25409/thread

Squamata:

•Hainosaurus:

• theworldofanimals.proboards.com/post/20730/thread

•Varanus komodoensis (Komodo dragon):

• theworldofanimals.proboards.com/post/1008/thread

Other Reptilia:

•Ocepechelon:

• theworldofanimals.proboards.com/post/4650/thread

Basal tetrapoda:

•General:

• theworldofanimals.proboards.com/post/25653/thread

Actinopterygii:

•Megapiranha:

• theworldofanimals.proboards.com/post/21775/thread

Elasmobranchii:

•Carcharodon carcharias (Great white shark):

• theworldofanimals.proboards.com/post/26769/thread

[creature386]

"We conclude that the skull and associated musculature of V. komodoensis are particularly well-adapted to exert and resist forces generated during pull-back biting. The posterior rostrum presents extra mechanical advantage to hold the prey and withstand pull-back loads imposed by retraction of its skull, and the anterior part is equally well adapted for exerting lateral pull bites.
The highly fenestrated, lightweight skull of V. komodoensis is optimized to resist a complex and finely balanced combination of adductor forces and loads generated by cervical and other postcranial muscles during killing and feeding. This combination of anatomical and behavioural features, together with its markedly ziphodont dentition, allows the animal to kill and deflesh large prey efficiently, using relatively low jaw adductor driven bite force."
onlinelibrary.wiley.com/doi/10.1111/j.1469-7580.2008.00899.x/full

[theropod]

Allosaurus/Allosauridae
Unfortunately Rayfield et al., 2001 doesn't have a nice, short conclusion. I suggest reading the whole paper, but here i quoted the most important Information.
www.academia.edu/286996/Cranial_Design_and_Function_In_a_Large_Theropod_Dinosaur

First, compared with values for extant mammalian taxa, the
muscle-driven bite force of Allosaurus fragilis appears to be relatively
weak (Table 1).

the given figure was 80 (!) to 200kg. Bates & Falkingham, 2012 gave a 4 times higher estimate for a similar sized specimen. I asked Falkingham on askabiologist and he stated this may partly be due to SMA 0005 having different morphology, but stating at least the lower range (unsurprisingly) to be probably too low.
www.askabiologist.org.uk/answers/viewtopic.php?pid=21222#p21222
The metod used to estimate the bite force was modeling the muscles in clay and using the cross-sectional area for estimating their strenght.

Second, in contrast to these weak muscular-driven bite forces, the
skull is strong enough to withstand extremely large maximum
forces at the tooth row before yielding because of tensional stress
(Table 1, modes C and D; Fig. 2a, b).

With the tension excerted by the adductor muscles and a condylar force applied, the skull of MOR 693 could still withstand about 1,8t of force. Without it, it even reached 5,5t.

Third, tensile stresses ventrally and compressive ones dorsally
re ?ect the bending moments acting on the skull during loading
(Fig. 3a, b). However, the skull of Allosaurus seems to be designed to
resist large vertically directed forces applied along the tooth row.

Studies suggest that the large Cretaceous carnivore T. rex, with
stout, conical teeth and an extremely robust skull15, could withstand
tooth±bone impacts during feeding, and was capable of generating
jaw-closure forces large enough to shatter skeletal material during
prey dismemberment12,16,17. A weak muscle-driven bite indicates
that Allosaurus was not capable of splintering bony food material in
this manner. In contrast to T. rex, Allosaurus displays recurved,
laterally compressed teeth adapted for slashing or slicing8 and a
more lightly constructed but extremely strong skull (Fig. 1a, b)5. We
believe that this `weak bite/strong skull' functional paradox may be
explained by the predatory behaviour of Allosaurus.

This seems to be analogous to V. komodoensis, hence the parallel drawn in the other paper.

If the animal was to experience large
forces such as those created by a high velocity impact of skull into
prey, as part of its regular feeding strategy, the skull could still
function within acceptable bounds of safety and the apparent
overcompensation of design can be explained.

An impact-scenario is proposed. Snively & Russel proposed something similar to this based on neck-muscle studies.
They too noted the prominent basitubera and ventrally-pointing paroccipital processes, attachment points for M. longissimus capitis, M. iliocostalis and m. rectus capitis ventralis, all head depressors.
The dorsal neck musculature (eg. M. transversospinalis capitis) is also well developed, as can be seen in the prominent nuchal crest of the parietals, however lateral musculature is less prominent than in Tyrannosaurs.


Here is what is stated in Antón et al., 2003
digital.csic.es/bitstream/10261/22490/1/102.pdf

Bakker (1998) has shown that the paroccipital
process of Allosaurs projects clearly downwards
instead of laterally as in other theropods (Fig. 2).
The paroccipital process serves for insertion of the
transversarius musc1e, which in turn transmits the
contracting force of a bundle of more posterior
musc1es inc1uding the levator scapu1ae, sternomas-
toideus and sca1enes.

The great development of a paired process called
the basitubera in the base of the cranium of
theropods indicates the presence of strong head
depressor muscles, and their enhanced vertical
action is shown by the ventral projection of those
processes (Fig. 3). Bakker (1998:152) observed the
presence of "a set of transverse thickenings in the
bone boxwork behind the basitubera", which he
interpreted as evidence showing that "un usual
stresses were incurred during head ventral-flexing".

The anterior position and hypertrophy of the
insertions for neck and head extensors such as the
spinalis capitis in the top of the braincase of
Allosaurus (Fig. 2) suggest considerable ability to
elevate the snout, fitting with Bakker's suggestion
that the animals attacked with their snout tilted
upwards, as modern crocodiles often do.

these are the neck muscles shortly outlined.

As Bakker (1998) has shown, the shape of the
cranio-mandibular articulation in Allosaurus allows
the jaw to rotate along a wider arc than is possible
for other theropods without disarticulation of the
mandible.

A wide gape is supported by the presence of an antarticular and the insertion of the muscle fibres.

This paper suggests the "Komodo dragon-analogy" and seems to generally propose a very similar biting mode. I think the Komodo dragon makes a decent analogy for many narrow-toothed theropods (ie. most if not all Carnosaurs, many coelurosaurs, many ceratosaurs...)

The newest analysis from 2013 apparently agrees with both.
palaeo-electronica.org/content/2013/389-allosaurus-feeding

These analyses indicate that ventroflexive
angular acceleration in Allosaurus was greater
than calculated for Tyrannosaurus (Snively and
Russell, 2007c), as was predicted by some previ-
ous studies (Bakker, (1998[2000]); Rayfield et al.,
2001; Snively and Russell, 2007c). Forceful ventro-
flexion supports the hypothesis that Allosaurus
augmented its bite force by ventroflexing its upper
jaws (Bakker, 1998[2000]; Rayfield et al., 2001),
and suggests that it could strike downwards
quickly.
Dorsoventral and protractive ranges of motion
mirror those observed (Snively, 2006) and exam-
ined radiographically (Samman, 2006) in volant
raptorial birds and inferred in the terrestrial phorus-
rhacid, Andagalornis (Tambussi et al., 2012).
These ranges of head position suggest the efficacy
of striking prey in the sagittal plane. In the posterior
portion of the neck, lateral range of motion appears
to be greater than observed in extant birds (Sam-
man, 2006; Snively, 2006). Anteriorly, the lateral
intervertebral range of motion is similar to that of
the bald eagle Haliaeetus leucocephalus and
snowy owl Bubo scandiacus (Samman, 2006), and
greater than in Tyrannosaurus rex (figure 6.12 in
Samman, 2006, Samman, in press; also examined
preliminarily by Snively and Russell, 2007b). Neck
and cervicocephalic range of motion in Allosaurus
will be testable more thoroughly with methods that
Stevens and Parrish applied to sauropods (Ste-
vens and Parrish, 1998; Stevens and Parrish,
2005a, b), and Samman (2006; Samman, in press)
used for tyrannosaurids.
Range of motion and accelerations may be
informative about how Allosaurus defleshed its
prey. Strong ventroflexive torque suggests a more
birdlike posterior pull on a carcass than crocodil-
ian-like shake-feeding. (The latter is more likely in
tyrannosaurids [Snively and Russell, 2007a, c; Wit-
mer and Ridgely, 2009], with their great moment-
generating capacity to decelerate and reverse
direction of the head [Snively and Russell, 2007a]).
With posterior neck dorsiflexors and anterior head
ventroflexors acting in concert for head retraction,
Allosaurus may have fed more like a large preda-
tory bird than like a crocodile (Snively, 2006;
Snively and Russell, 2007a, c). However, Allosau-
rus was probably not limited to a single strategy for
removing flesh. Its laterally compressed, ziphodont
teeth suggest forceful alternating tugs to either side
of the head, as seen in Komodo dragons (Auffen-
berg, 1981, DÂ’Amore et al., 2011).

It is also suggested the neck was very flexible, which is in accordance with its lenght, supposed ecological role of Allosaurus and possible lunging motions.
The whole thing is a mixture between Andalgalornis, extant raptors and komodo dragons. What exactly it was is still not sure, but there was certainly some sort of head depression and pull motion, and possibly, as suggested by two studies, some sort of striking/slashing-motion.

Also according to Snively et al., (the nasal paper, tough not everything in that study convinces me) 2006 The teeth of Allosaurus are broadly comparable to similar-sized Tyrannosaurs in terms of strenght (as I already noted there's certainly allometry leading to more robust structures, including teeth, in larger animals, so those cannot be directly compared to), likely result of the relatively low crown height at least in the specimen in question.
theropod nasals and skulls

N CH (mm) FABL (mm) MLBL (mm) AP str. m3 ML str. m3 Skull l. cm
Tyrannosauridae
Albertosaurus sarcophagus (As) 7 70 29.9 17 0.214 0.122 86.0
Daspletosaurus torosus (DtS) 9 37.9 16.8 11.1 0.086 0.055 57.3
Daspletosaurus torosus (DtL) 6 72.5 27.3 19.7 0.202 0.144 97.0
Gorgosaurus libratus (Gl) 13 54.8 22.90 12.90 0.12 0.213 75.0
Gorgosaurus libratus (GlL) 11 57.1 23.13 22.6 0.127 0.088 76.0
Tyrannosaurus rex (TrA) 10 101.6 37.0 28.20 0.471 0.630 119.0
Tyrannosaurus rex (TrB) 11 98.4 45.0 33.50 0.864 1.154 140.0
Tyrannosaurus rex (TrL) 9 78.6 41.4 25.70 0.659 1.042 122.0
Tyrannosaurus rex juv. (TrJ) 11 45.5 22.6 11.20 0.131 0.065 74.0
Non?tyrannosaurids
Acrocanthosaurus atokensis (Aa) 1 84.0 31.0 19.5 0.492 0.310 105.0
Allosaurus fragilis (Af) 14 44.3 18.40 12.70 0.121 0.176 76.0
Carnotaurus sastrei (Cs) 12 47.5 22.60 15.70 0.211 0.310 56.0
Ceratosaurus dentisulcatus (Cd) 10 62.9 26.10 10.60 0.079 0.193 63.0
Monolophosaurus jiangi (Mj) 13 39.8 19.80 11.00 0.103 0.187 83.0
Sinraptor dongi (Sd) 12 47.5 22.60 15.70 0.211 0.310 84.0

I think Allosaurus could have delivered a quick slashing bite, resembling the hatchetting motion but not as extreme, in ventroposterior direction, and in immediate succession bit and pulled its jaws back and down, the later stage just like V. komodoensis. Considering the narrow jaws, a shorter but more focused contact with the prey seems likely, so maybe not a long holding-onto the prey animal but a powerful but short action. That supports the striking hypothesis imo.
Pulling forces in V. komodoensis were orders of magnitude greater than the bite force (D'Amore et al., 2011), which fits the cranial strenght of Allosaurus.

An odd thing is that while other studies found a very large gape, due to the insertions of the jaw muscles and the anatomy of the mandibular articulation, Bates & Falkingham, 2012, found a very high mechanical advantage for the adductor muscles, significantly higher than in any other specimen in the study. They also stated the relatively high bite-force estimate for SMA 0005 was largely due to this.
How does this fit together?

One thing I note is that in many animals, and almost all ziphodont slicers, it is by far not only the adductor musculature that inflicts the damage, but postcranial, mainly cervical activity, inertial feeding, shaking pulling, slashing etc. of some sort. Therefore comparing adductor force to asess bite potency has lost its last bit of relevance.
In slicing animals, a traditional bite alone may do insufficient damage, due to the missing power and lenght to pierce deep. however in all taxa employing such a bite that I know of some additional activity greatlyincreases the effect, often by transforming the jaws into a sort of saw like in sharks or monitor lizards.

I outlined my thoughts here: www.paleo.keepfree.de/allobite.html

References & further reading:
Snively et al., 2007: www.ohio.edu/people/es180210/Snively%20pdfs/snively_russell_theropod_necks.pdf
Snively et al., 2013: palaeo-electronica.org/content/2013/389-allosaurus-feeding
Rayfield et al., 2001: www.academia.edu/286996/Cranial_Design_and_Function_In_a_Large_Theropod_Dinosaur
Antón et al., 2003: digital.csic.es/bitstream/10261/22490/1/102.pdf"]http://digital.csic.es/bitstream/10261/22490/1/102.pdf
D'amore et al., 2011: www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0026226
Moreno et al., 2008: www.ncbi.nlm.nih.gov/pmc/articles/PMC2423397/
Bakker, 1998: www.arca.museus.ul.pt/ArcaSite/obj/gaia/MNHNL-0000779-MG-DOC-web.PDF

PPSI: I think musculus transversarius as in Antón et al. and Bakker is the same as M. longissimus capitis profundus/superficialis as in Snively et al (both papers), isn't it?


PS: PSI=PostScriptumInterrogo

[theropod]

Reply to grey from the size-comparison's thread.

May 24, 2013 21:16:43 GMT 5 Grey said:

No, that's the entire tooth that counts in the mechanical efficiency, excluding the root part is not wise. If sharks had not evolved to replace their teeth in their own way, they would have too a very deep root and here too, the root would have to be counted in the comparison.

Anyway, both types of animals are acknowledged to have a similar, if not totally the same, killing/feeding apparatus, relative to the thape of their teeth.

But the root cannot break, it is stuck in the bone.
It is merely the crown that is exposed to the stresses of the bite, while the root will behave more or less in the same way as the surrounding jawbone. If the teeth of these animals fracture, it is not the root that breaks, but the crown, hence the part whose sturdiness you want to compare is the crown, and it provides an unbiased image because it is comparable in both of them while the root is not.
I think it is clear komodo dragons might be the best analogy for large carnosaurs that we have, shark teeth and jaws are different in morphology, however it is sufficient to show the claim of absurdely weak teeth that would shatter like glass is biased.

It does not appear any modern animal, be it one adapted for slicing flesh or for breaking bones, totally avoids bones in a fashion like the one occasionally suggested, and the tooth-crowns of carnosaurs are more robust than those of extant lamniforms, even tough probably at least in part due to allometric reasons.

[Grey]

The root does not crack but the root plays a definite role in the likelihood or not of the cracking of the tooth.
That's why we cannot properly compare the crown of a shark with the crown of a tetrapod as it produces biased results in both.

But they are used in a similar way, that's what counts at the end of the day.

[theropod]

Are there functional studies of the importance of the root for preventing a tooth from cracking? I never heard of that.

I could imagine that being the case in triangular shark teeth, were the rooth is the broadest and thickest part, but why should it be important if it is not more than just an elongation of the blade?
Still, the best way would be to compare the lever on which forces are excerted and that hence is the most exposed to stress, not the tooth including a part that could hardly be more different among the two.
Carnosaurs definitely have the more robust tooth morphology here, and their teeth are not clovis-points or flint-flakes.

[Grey]

The root first role to play is to prevent the teeth to crack.
why the root in T. rex is longer than in carnosaurs, because the works exerted by the crown is harder.
Read Bretton Kent about the mechanical efficiency of the teeth in megatooth (Hell's Teeth).

The crown of carnosaurs appear more robust because they compose a shorter part of the entire tooth and the tooth itself are far larger as the animals are larger than the white shark.

Really, the comparison here is very very limited by too many factors and using only the crown is not wise.

[theropod]

But it is better than using the whole tooth including roots that are built completely differently. Of course this comparison lacks rigour and it is not possible to draw complex conclusions from it. All we can say is that allosaurian theropods definitely had thicker tooth-crowns than lamniforms and therefore the statement of them "totally avoiding bones" is not justified, neither is the hypothesis that they could only cause a very slow death through blood loss and were unable to kill quickly, a point frequently brought up against them when their jaws are compared to those of animals with blunter, broader teeth and jaws.

T. rex has long roots because the teeth have to be prevented from breaking out. that's different from preventing the teeth from breaking. I honestly don't see what role a root that is pretty much the same depth and width as the tooth and fully embedded in the jawbone could play for the stability of the crown.
Of course the crowns are a shorter part, that's the reason why comparing the whole tooth would be biased. The functional part however has certain similar properties that make this comparison the best. It is the only part prone to breaking.

[creature386]

Theropod, it seems like Grey's opinion isn't very different from yours, because the quote of Kent says pretty much the same as you say:

The slenderness ratio is a measure of length to breadth. Needle-like teeth with high slenderness ratios are good for grasping small prey, like mackerel or flounder, but buckle under loads applied along their length, as would happen if an animal munched on the bone of a large sea mammal. Mechanical advantage relates the length of a tooth to the length of its root. Long teeth with short roots have low mechanical advantage and are more easily pulled from the jaw than teeth with comparatively larger roots.

The teeth of great whites have the high bending strength ratios of cutting teeth, high slenderness ratios, implying they are not designed for crushing, and roots that are short when compared with the overall tooth height. This means a great white's teeth have a low mechanical advantage and will likely be ripped out by struggling prey. Such a combination of parameters, says Kent, "is ideal for a shark that hunts with a slash-and-release type of attack that concentrates on the softer portions of the prey". Surveys of the corpses of marine mammals killed by great whites confirm that, when feeding on sea lions, small whales and porpoises, the sharks most frequently attack the delicate abdomens or rear flippers of their victims. Though they often tear off chunks of flesh during an attack, the sharks rarely grapple with large prey, waiting instead for the animal to die of injuries, or carrying animals in their jaws until they bleed to death, before feeding at their leisure.

Megalodon teeth, though superficially similar to those of great whites, have an entirely different set of parameters. Though they share the serrated cutting edge, megalodon teeth are comparatively thicker for their size with much lower slenderness and bending strength ratios. They also have roots that are substantially larger relative to total tooth heights, and so have a greater mechanical advantage.

Kent explains that teeth with these traits not only make good cutting tools, but are also well suited to grasping powerful prey and unlikely to crack when they bite down on bone. The differences between the teeth of great white and megalodon suggest fundamental differences in their diet, attack behaviour, or indeed both. But Kent reasons that as large mammals formed the menu of both sharks, the difference in their teeth must be related to variations in their attack behaviour.

Source: issue 2190 of New Scientist magazine, 12 June 1999, page 32
Maybe he ment being ripped out by saying "being cracked".

[theropod]

Exactly, it is about anchoring the tooth in the jawbone, not about preventing the crown from breaking. My points were merely about the sturdiness of the tooth itself, not about how well it is protected from being lost.
In this regard, sharks and theropods are not comparable, as sharks have teeth that can be replaced within hours and usually loose extreme numbers of them, while theropods have deep roots. Enlargement of the root is commonly associated with a more robust tooth, but there is no direct correlation between crown stability and the root. The crown can still break off even if the root is large, or it can be very strong while the root is to week to prevent it from being lost.

[theropod]

Does anyone have data on the cranial strenght of carcharodontosaurs? I fear the theropod-nasal paper might not be a particularly good source for the biomechanics of the whole skull, since its rather simplistic skull models possibly produced biased results against animals with more lightly constructed skulls, as can be seen in their low values for Allosaurus.

[Grey]

The degre of depth of the tooth, thus the anchoring, plays a role in the robustness of the tooth, as much as the structure of the crown itself.

In the case of the white shark, keep in mind too the actual shape of the tooth with a broad basis, this also provides robustness in the overall object.

Also, I'm not certain that the model of that tooth in the image is really accurate at all as the purpose was strictly to compare it with the thicker megatooth tooth.

I insist that these different teeth are hardly comparable, if not in their actual job when biting.

Regarding carnosaurs bites on bones, of course no one argues against the fact that they could superficially inflict marks on big bones, only they don't do what specialized bone-crushing teeth do.

[theropod]

I do not suggest bone crushing, just that smaller bones wouldn't be a hindrance for inflicting major injuries and the teeth were not in danger of fracturing on the smallest contact with robust tissue.

And of course that comparing the width when normalised for overall lenght with the root included produces a bias towards short rooted taxa. Of course the root plays an important role, but at least in Carnosaurs not for crown strenght bit for ensuring the tooth doesn't break out of its socket.

[Grey]

The main point was that forgetting the broad structure, increasing the robustness of the overall tooth, produces then a bias toward the teeth of carnosaurs. That's for showing you that the comparison can hardly be fairly done.

Of course carnosaurs can puncture small, gracile bones. But it is only said that if they bite through big bones, they just break.
Just like for the white shark, which, because of its way to replace broken teeth, is less problematic.

[theropod]

Biting into thick bones is not what these teeth are made for, in neither case. This however is not a weakness and restricting the use, it is a different purpose.
The shape differences are partly due to the jaw-orientation; in sharks, a very prominent component in damage-dealing is lateral shaking. In carnosaurs or monitor, it is posteroventral striking and pulling.
I don´t think GWS teeth are rostroposteriorly much broader than carnosaur teeth.