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e and prone to cracking and breaking under strain to that end teeth emerging from oral margins tend to be more damaged and worn than those kept within a moist sealed mouth this difference can be seen with the naked eye but is particularly obvious under microscopic examination in our paper we show that the tips of alligator teeth are shorn off on their outward facing exposed surfaces with both the enamel and several layers of dentine worn to a flattened edge tooth dehydration almost certainly factors into crocodylians frequently suffering from broken and cracked dentition and they have to replace their teeth regularly something like 45 50 times in a lifetime grigg and kirchner 2015 to maintain a set of fully functional jaws these observations give us a clear hypothesis regarding theropod oral tissues if they were permanently exposed theropod teeth should show at minimum similarly obliterated enamel and dentine layers at their tips the poster children of exposed dinosaur teeth adult tyrannosaurids are especially relevant here as they replaced their teeth at a very slow sometimes even biennial rate erickson 1996 what s more they engaged in particularly violent tooth on bone feeding strategies so if any theropods are going to have knackered abraded teeth it s tyrannosaurids detailed comparisons of tyrannosaurid upper row and crocodylian lower row tooth wear note how the daspletosaurus tooth despite being over 500 days old is intact despite dinosaurs teeth having particularly thin enamel layers the alligator tooth tip by contrast has not only lost the enamel coating on its outer surface but also several layers of underlying dentine a fully intact enamel covered erupting alligator tooth is shown in panel h to show that these are features of wear not the original tooth condition from cullen et al 2023 but extracting a fully grown c 510 day old tooth from a daspletosaurus maxilla revealed a tooth tip in great condition both the inner and outer surface retained their relatively thin enamel covering and the only minor damage found was on its medial inward facing surface possibly as a result of occasional tooth on tooth contact this confirms what has generally been observed for theropod dental wear in other studies and conflicts with what we d expect from a dehydrated perpetually exposed tooth we take this as evidence of theropod dentition being maintained in a moist well hydrated setting and being located within a lipped mouth is realistically the only way this might be achieved too big to sheath we also address the possibility that some theropods had teeth that were simply too big to cover with lips e g ford 1997 our focus here is not on the widely known but still surprisingly prevalent issue of artists and researchers not accounting for tooth slippage in fossil specimens rather we focus on realistic estimates of tooth crown height when they are fully socketed in theropod mouths we calculated a ratio of tooth crown height to skull length for 37 theropod specimens and compared them with the same metric in 40 varanids the monitor lizards varanids are of course well known for possessing large theropod like teeth as well as copious amounts of lip and gum tissue this is the decay of tooth anchoring ligaments resulting in teeth sliding somewhat from their sockets preserving them at longer lengths than they held in life tooth size theropods vs varanids it turns out that both groups have similarly sized teeth relative to skull length and while this doesn t directly tell us if theropods had lips it shows that their dentition was of a size that we know can be sheathed by extra oral tissues today from cullen et al 2023 plotting these data showed that theropod and monitor teeth are about the same size for their skull lengths and even increase in proportion at the same approximate rate but the winner of the biggest tooth contest wasn t something like t rex it was the varanids some monitor species like the crocodile monitor varanus salvadorii have almost cartoonishly large dentition from this we suggest that theropods did not need unprecedentedly big lips to cover their mouths and the largest theropods wouldn t look in terms of lip proportions very different to something like a komodo dragon indeed we note that monitors are able to cover their teeth with the same basic configuration of labial and gingival tissues across a 12 fold size difference the discrepancy between the largest monitor skull and our largest theropods is only half that 6 fold so if lizard lips and gums can scale 12 times over without substantial anatomical deviation perhaps they could stretch to cover the teeth of much bigger animals without much change too whether we ve realised it or not a lot of us already evidently believe this is possible given the abundance of lippy monitor esque mosasaur reconstructions sealing the deal these are the points we cover in the main paper but this being an article in science it s essential to also check out the supplementary files for additional discussion and context therein we raise another point that resulted from our efforts at reconstructing a scientifically informed illustration of a lipless tyrannosaurus it s really really hard maybe even impossible to seal at least some theropod mouths without lips forming an oral seal even if it s just by pressing lipless jaws against one another is important to avoiding dehydration as well as maintaining basic oral health and hygiene we could not however find a way to reconstruct t rex jaws without leaving a gap behind their maxillary teeth i actually pushed our lipless reconstruction fig 1b in the paper a little beyond what i think is reasonable and we re still left with a small opening we are not the first people to ponder this issue and dinosaur literature has contrasting views on how far theropods could close their jaws some authors propose that theropod mandibles could be pulled way up into the cavity of the upper jaw and have even identified landmarks for the resting position of the lower teeth below these include depressions in the walls and roof of the upper oral chamber that seem suited to act as socket like structures for receipt of the lower dentition e g molnar 1991 ford 1997 currie 2003 hendrickx et al 2014 others most notably tyler keillor 2013 in his excellent book chapter on restoring the face of the jane tyrannosaurus have questioned this idea on grounds that theropod mandibles can t close so tightly without literally bashing into problems examples of landmarks suggested to record the resting poses of theropod lower jaws maxillary wall sockets are depressions in the internal wall of theropod oral cavities and some theropods are also preserved with round depressions in the roofs of their mouths neither are universal features of all theropods however images from osborn 1912 lü et al 2014 and cullen et al 2023 reconstructing the face of t rex for our paper saw us agreeing with tyler s conclusions at a certain point of mouth closure theropod lower jaws collide with bones under the eye socket specifically the ectopterygoid so that further adduction either requires the jaws to literally crush themselves shut or else the bones of the posterior skull act as a hinge swinging the jaw tip into the mouth but dislocating the jaw joint ford 1997 proposed that a notch in the ectopterygoid accommodated the closed lower jaw during mouth closure but we don t think this is plausible theropod ectopterygoids can be complex shapes and yes some have regions that superficially look like they could nestle the lower jaw but these were almost certainly filled by deep jaw muscles in life e g gignac and erickson 2017 there are of course theropod skulls preserved with their jaws tight shut in the fossil record but we have to be careful assuming these represent in vivo conditions given how routine processes of decay and fossilisation can pull and crush carcasses into unnatural configurations inspired by this we devote some discussion to how theropods posed their closed jaws in life beyond ruling out impossible jaw busting configurations it s difficult to know exactly how tightly theropods held their mouths but this is something for artists to consider x rays and scans of lizard carcasses show that their jaws are far from clenched shut when their mouths are closed and in some species their upper and lower dentition barely overlaps if we go full lizard with our theropod reconstructions where we apply minimal overlap of the upper and lower toothrows their lips would have been deep and their snouts much taller than we re used to we play about a little with this visually in the paper and i was struck at the blocky chunky cranial profile of our lizard like loose mouthed t rex below some of our experimentation with theropod mouth postures the crush closed pose is almost certainly impossible but it s hard to say how relaxed theropod resting gapes may have been held if modern lipped reptiles are anything to go by they may have been held far more open than we re used to modified from cullen et al 2023 conclusion theropod jaws only make sense if they had lips putting all this together our investigations of jaw structure tooth size tooth wear and jaw closing all point to the same inference theropod jaws don t make much anatomical or functional sense without lips of some kind to validate the alternative lipless model we have to engage in a lot of special pleading and scientific weaselling exposed theropod teeth would have to be unprecedentedly resistant to wear all our understanding of jaw structure and foramina distribution correlating with oral soft tissues would have to be wrong and theropods would need to be unique in not bothering to create oral seals if we re being good scientists we can t currently say that theropods definitely had lips drop the microphone and walk off stage but i think we ve made it far more challenging for anyone to legitimately object to the lipped theropod hypothesis time will tell on that front in addition to substantiating the lipped hypothesis of theropod appearance our hope is that our paper may establish some lines of inquiry for the oral tissues of other extinct animals many of the most extreme dentitions to ever evolve belong to fossil taxa after all and theropods are far from the only species with uncertain facial appearances what of nimravids gorgonopsians uintatheres or thylacosmilus and what for that matter of the superficially crocodile like spinosaurids and other weird theropods were they lipped or not if our ideas hold water they provide a relatively straightforward way of deducing whether the teeth of these animals were held within oral tissues we can t of course finish without some brief notes on the life appearance of lipped theropods we address this a little in our supplementary information and conclude that lepidosaurs the lizards and tuataras are the best modern analogue for theropod lips and gums this is admittedly a best of a bad situation recommendation because there are plenty of differences between theropod and lepidosaur jaws that preclude total confidence in their comparison but we only have so many extant reptile groups to choose from and lepidosaurs are on the whole morphologically closer to theropods in areas we think are influential on labial and gingival tissues what did lipped theropods actually look like lizard mouths have a lot more soft tissue surrounding their teeth jaw cross sections show a komodo dragon b and alligator c and this might be something we need to bring into our theropod artwork the gaping t rex shown here is outfitted not only with large lips but also a conservative 25 of its tooth height covered with gums from cullen et al 2023 supplementary data this being the case our model for theropod mouths is that they were sealed by non muscular lips covered with scales or to hedge our bets a little more whatever epidermal covering was present on the side of the snout lepidosaurs show variation in lip size with most having generous upper lips but some having thinner lower lips than others this variation continues to their gums lepdiosaur gingivae are more voluminous than those of mammals and crocodylians and generally cover at least 20 25 or so of tooth crown height this is why lizard teeth aren t always that conspicuous in their open mouths varanids take these enlarged gingivae to an extreme hiding almost all of their formidable teeth with enormous gums we currently don t have much insight into where theropods sat within this range paul 2019 independently of our study advocates for full monitor like conditions for theropods and this might be possible but we can t rule out smaller gums or indeed a unique theropodan take on oral soft tissues at this time we propose however that since extant lipped reptiles have at least 25 of their tooth crown heights covered with gums we should apply that to theropods too we ve explored this in our paper and pr art with that gummy lippy t rex shown above it s not strictly true that lizards have no muscles around their mouths some agamids have muscles that move their lips or flaps of mouth adjacent skin for communicative purposes i m no expert on these structures but i think it s fair to assume that they are specialisations of their respective lineages not the remnant of a once ubiquitous clade wide lepidosaur ability and i think we ll leave it there for now there s more to say on the tyrannosaur restorations we created for the paper with their small eyes and some of their skin details but at least some of that discussion needs to wait for developments in other research i m involved in i m so so sorry dave for now i ll thank my coauthors once again for inviting me onto such a great project and i ll leave you with this handy infographic summary of our research which you can share around the internet to instigate discussion of dinosaur life appearance wherever you may be the qr code in the corner will take you directly to the paper so if you or anyone you encounter wants to know exactly what we have to say about theropod lips you can always find it from this image enjoyed this post support this blog for 1 a month and get free stuff this blog is sponsored through patreon the site where you can help artists and authors make a living if you enjoy my content please consider donating as little as 1 a month to help fund my work and in return you ll get access to my exclusive patreon content regular updates on upcoming books papers paintings and exhibitions plus you get free stuff prints high quality images for printing books competitions as my way of thanking you for your support as always huge thanks to everyone who already sponsors my work references bakker r t 1986 the dinosaur heresies new theories unlocking the mystery of the dinosaurs and their extinction william morrow barker c t naish d newham e katsamenis o l dyke g 2017 complex neuroanatomy in the rostrum of the isle of wight theropod neovenator salerii scien...