Dinosaur tails redux

Getting on for ten years ago, I published a paper looking at the lengths of the tails of dinosaurs. The short version of that is that total length of tails in dinosaurs varies massively both between clades and even within groups (or within species!) which mean that a lot of the ‘total length X’ estimates for various dinosaurs are probably way out. Still, it wasn’t the biggest dataset and there’s not a lot of nuance to looking at total length vs body length, plus being restricted to only complete tails really cuts down on the number of specimens you can use.

Still, not too long after the paper was published, I set about trying to get a better dataset together as more dinosaur tails were coming out of the woodwork. That led to this appeal on here which helped reel in a few more specimens. Still, I wanted to do something more detailed and that led me to roping in my friend and colleague Steve LeComber.

At the Cheltenham Science Festival (L-R, Me, Steve Le Comber, Chris Faulkes, Jane Hallam)

Steve will be all but unknown to readers on the Musings as he never worked on dinosaurs before, though as a great science communicator he helped me out at a number of my events, especially when we went to the Cheltenham Science Festival together a few years ago.  Sadly, this will also be one of his last papers as he passed away at the end of 2019. Steve was one of my closest friends and colleagues, and was one of the most popular and friendly people I have ever met. He had an entire career as a journalist and writer before switching to science and was a superb statistician as well as a great biologist and a wonderful educator.It’s a testament to his work that papers are still coming out of his lab and his work on geographic profiling will have an important and lasting legacy in biology. He will be forever missed. (A scientific obituary was published for Steve here in the Journal of Zoology, where he was an editor for many years).

I had turned to Steve because he was tremendously creative with analyses and I had no idea how to approach the next problem I wanted to tackle – what was happening with individual vertebrae in the tails of dinosaurs? Very little has ever been written about this, and what there is implies or even states that as you go down the tail, each vertebra is shorter than the last. But you only have to look at a couple of specimens to see that this really isn’t the case. It’s true for big chunks of the tail, but the part closest to the hips often has short vertebrae but after that they tend to get longer, and in very long tails like those of sauropods you can find multiple sets of vertebrae that are lengthening. But how to capture this information? If groups of vertebrae are letting longer or shorter, and if this changes slowly or dramatically, or is just an oddity, can we capture it? Happily, he had some ideas.

As I was by now established at Queen Mary, I really lacked the time and opportunities to revisit collections and measure individual vertebrae so we then roped in Scott Persons, who has his own interest in dinosaur tails and had measurements we could use, or was able to get into some of the Canadian collections to procure more data. Various issues delayed the paper on numerous occasions but it is now out so here’s some quick take-home results (the full paper is in PeerJ and open access so you can see all the figs and data there).

First off, with more and better data, we did revisit the issue of overall dinosaur tail length and it is still very variable and unpredictable. Total length estimates without most of the tail present could easily be very wrong, and even some tails that you might think are pretty complete could easily truncate suddenly or go on much longer than you might think. There’s still a place for these of course (especially for engaging the public) but the standard ‘my theropod is longer than yours’ battles really need to stop. Total length isn’t a great indicator of size (mass is) and tail lengths, and by extension total lengths, are very hard to estimate without a near complete tail.

Obviously we do now have a much better Spinosaurus tail, but this old image by Scott Hartman demonstrates just how wildly different tail lengths (and so total lengths) of an animal could be for the same body size.

On to patterns within tails. First off we do find that individual vertebrae within tails simply don’t tend to get shorter as you go along them. There’s some interesting and cool patterns going on and I don’t want to cover all of them here (for example Coeplohysis is all over the place, and Juravenator seems really weird) but here’s a few of the more interesting ones. We use broken-stick regressions where we can have multiple different trajectories of sections of the tail lengthening, staying the same length or shortening. It’s a great tool to see what is happening and is visually nice and easy to follow, without getting mired down in the odd vertebra that’s rather out of place with the others.

First off, that means that it’s good for spotting changes in patterns of vertebrae lengths and also deal with (bits of) missing data quite well. This is also really useful for predicting the lengths of missing vertebrae and this is likely to be useful for things like working out total lengths of animals and the sizes of individual verts when reconstructing fossils. It’s also absolutely ideal for putting together skeletals and even mounted skeletons in the future.

Second, many dinosaurs have a pattern of a set of short vertebrae, then longer ones, and then the rest of the tail does indeed taper off. The second switch (were the long ones stop and it starts to taper) coincides with the ‘transition point’ in the tail, where the main leg muscles terminate, suggesting an important link between the two and from this we hypothesis that this short-longer- tapering pattern is a functional one linked to tail flexibility and muscle power. This clearly needs more work, but it’s a very interesting starting point.

Next, some exceptions. Plenty of dinosaurs don’t fit this pattern for various reasons (some it’s probably just missing data or it’s a subtlety like the tapering happens in two different phases), including some that do just generally taper. There’s some huge intraspecific variation in some but others are very consistent. All three specimens of Archaeopteryx we included show a weird humped distribution which is also very similar to Microraptor, and also different to other dromaeosaurs. That rather implies that this is flight related and that this is an important convergence, though again quite what and how is well beyond what we cover in the paper, it’s an area that hopefully others will pick up on. And aside from Microraptor, the dromaeosaurs appear to be highly variable which we attribute to their ‘sheath’ of elongated supporting rods for most of the tail length which would dominate any other functional issues and might leave the lengths of individual vertebrae to be fairly free of constraints.

I’ll leave it there since the paper is freely accessible and there’s lots that can be extracted from it, but I think this covers some of the more interesting points. The methods in particular should work well for any repeating units and while we have focused on dinosaur tails here, they should apply equally to any vertebral series or things like ribs, arthropod segments, and so on. I’d really hope that people will immediately see the use of this for describing things like sauropod or plesiosaur necks, pterosaur tails, or the lengths of neural spines or size teeth in a series. Of course I also need to say thanks to Scott and Steve and various referees and editors for helping get this published, and especially thanks to all those who contributed data to get this moving.

Hone, D.W.E., Persons, W.S.C. & LeComber, S.C. 2021. New data on tail lengths and variation along the caudal series in non-avialan dinosaurs. PeerJ. 9:e10721.

Protoceratops socio-sexual signalling again

It’s Protoceratops time again (hooray!). I have now published half a dozen papers about (or at least with a healthy dose of) Protoceratops andrewsi and a large part of that is because of the high number of really well-preserved specimens which make it one of the best dinosaurs to work on. There’s young juveniles through to large adults, large numbers of skulls and even skeletons, and all from a very narrow time and space that make it near enough (in palaeontological terms) a single population.

Among other things, I’ve used this to look at the growth and development of the frill and the implications for this major bit of dinosaurian anatomy being linked to sexual selection and dominance signals. (Here’s a recent round-up I did of all this work). The new paper out today is led by my former PhD student Andy Knapp and takes this previous simple work that only examined a few simple linear measurements and turns it up to 11.

Andy got to see almost every good Protoceratops skull going, including trips to New York, Warsaw and Mongolia to see material. Taking numerous photos of each allowed him to build 3D photogrammetry models of them and then get going with some quality morphometrics and analyses to look at how the skull changes in shape as the animals grow. It also shows that the skull can be divided up into distinct units and that these are somewhat independent of each other in their changes to size and shape and suggests evolution could act on each separately. The frill, rather inevitably, turned out to be a single unit and one that changed the most and one that is the most variable.

This fits very nicely with the prediction that the frill is used as a socio-sexual signal and if it’s true for Protoceratops then it’s probably true of other ceratopsians as well. I’d also suggest that this is good as we get very similar results from this very intensive and complicated methodology as we did from taking a few simple measurements of the skull and that gives some confidence that for future studies, the much quicker and easier methods may be more than sufficient to determine what kind of growth patterns are at play. There’s inevitably some more nuance and detail in the paper but happily this is another fully Open Access one, so click on this link here to be transported to a magical world of colourful ceratopsian skull bits in 3D.

I’ll finish here, but obviously huge congratulations to Andy on his great work on this long and complicated project and on getting an important paper out of it. I should also thank the Jurassic Foundation who gave Andy a grant to help with his travel and work on this project.

Knapp, A., Knell, & Hone, D.W.E.2021. Three-dimensional geometric morphometric analysis of the skull of Protoceratops andrewsi supports a socio-sexual signalling role for the ceratopsian frill. Proceedings of the Royal Society, Series B.