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There are a few fundamental biases in the fossil record and they can heavily influence what you are looking for. The older something is, the less likely you are to find it – every million years older it gets is another million years for it to be subducted, buried, eroded or destroyed. The smaller something is, the harder it is to find – 50 metre sauropods leave behind big bones that can be seen on the surface from half a mile away, rodent teeth are another matter. The more delicate something is, the rarer it is – it is more vulnerable to destruction before and after fossilisation, and even if you find it, it might be so badly crushed that all the details are lost. Finally, the rarer something was in life, the less likely you are to find it – a species with billions of individuals is more likely to enter the fossil record than one with a few hundred.
It might come as little surprise then to learn that despite intensive searches, and some very productive areas having been swept, Triassic pterosaurs are exceptionally rare. They are small, fragile, old and probably very rare (the group had jus appeared). Most of the few we have are badly crushed and have much of the animal missing.
However, (and this is a damned big ‘however’, with hobnailed boots on, and a large neon sign with ‘HOWEVER’ written on it round it’s neck), that has just changed. Raeticodactylus filisurensis is almost certainly the best preserved, most complete and generally all round excellent Triassic pterosaur known. It is in wonderful condition and tells us an enormous amount about the early days of pterosaurs. To pile glory upon glory, it is also totally bizarre and contains a number of very unusual features that make it unique in a great many ways. It would be going far too far to say it will revolutionise our understanding of the early days of pterosaurs, but it will add a massive chapter to what is currently a very short book.
That we have Raeticodactylus at all is a testament to it’s ‘father’ Rico Stecher. I have a whole blog post planned on Rico and his work, but in short he is simply a keen palaeo amateur who fond the specimen, prepared it, described it, presented his findings at a meeting (for which he had to take time off work to attend!) and then got it published. It is a fantastic achievement and one which I will write about extensively in due course. Rico will get his plaudits, but first I want to talk about the pterosaur. I met Rico while he was working on the description, he had come to Munich to look at our Triassic pterosaur material and we discussed his specimen at length, and ultimately I acted as the ‘translator’ on the description, giving the English a polish and helping him with some of the more technical aspects of the description. As a result, although I have not seen the specimen, I do know it pretty well having gone over the description several times and discussed it with Rico at length and have seen numerous photographs. As such I hope I can give some interesting insights into this bizarre little critter.
First off, as ever, I have no photos here, sorry. However there are some at Laelaps (I got beaten to the punch as Rico asked me to wait with this post until the paper was published, but Brian found it online early!) and even two seconds should be enough to convince anyone with a passing acquaintance of pterosaurs to realise how odd this guy is. Rico came to the Wellnhofer meeting in Munich and had the penultimate talk of the conference (immediately before Peter himself). I must confess to a little showmanship on my part as I was the only one who knew what was coming when he put up his first slide. I turned in my seat and quite literally saw jaws drop – yes, it is *that* weird.
Raeticodactylus is a rhamphorhyncoid pterosaur (obvious from the age and the skull shape) that can be diagnosed by any of about a dozen autapomorphies. I will not dwell on lots of interesting sublties – anyone *that* interested can go and read the paper, but there are still a large handful of very, very interesting features about Raeticodactylus that are worth discussing.
The most immediately striking thing about the specimen is the skull. Pterosaurs are well known for sporting some bizarre headgear, but even so this is extreme – especially in what must be a very basal taxon. Bony crests appeared *very* early in pterosaur evolution. The lower jaw is almost as strange, it has a deep anterior ‘keel’, the whole thing is very robust, the dentary is about half the size one might expect of a pterosaur and the articular is simply colossal (though quite possibly tied to the fact that the jaw is so damned big in the first place – it will need a big lever to shift it!).
The teeth are also interesting. Heterodont dentition is not that rare in pterosaurs (think Dimorphodon!) but there are unusual regardless. They show some nice fangs at the front and behind some nicely cusped teeth. Now previously cusps have only been seen in eudimorphodontids in pterosaurs, but several features suggest that this is *not* an eudimorphodontid. In short, cusped teeth evolved twice, and furthermore, there are a bunch of Triassic pterosaur fragments assigned to Eudimorphodon purely on the basis of cusped teeth – we might need to look at these again! Even this is not quite the end, and the teeth in the upper and lower jaws are arranged differently – tightly packed and overlapping in the mandible, but well spaced in the upper jaw. It gets harder with each passing sentence not to add an exclamation mark to the end of each line here, this thing is nuts.
The humerus is interesting as it is extremely long and slender. This does not make sense. If the damn thing was trying to fly it wants a short and robust humerus. Short to reduce the cycle time of a wingbeat, robust to support the heavy and powerful flight muscles required for powered flight (especially in such a basal taxon that one has to assume is not a great flier compared to some of the later clades). What was it doing? It certainly has the classic complex humeral head of a pterosaur indicating that there were extensive flight muscles, it is certainly not a glider or anything like that, but it is strange.
The femur is however, to those in the know, perhaps the crowing glory / weirdness of Raetico. Basically it looks like that of a theropod, or for that matter any of the derived archosaurs / ornithodirans that pterosaurs are derived from (in all probability – see number 15). It is long, slender, shows a nice sinusoidal shape and comes with a proper 90 degree angled head with a distinct caput. One of the ongoing issues over pterosaur origins is the relative lack of characters to tie them to possible ancestor clades and while the balance of power is with an ornithodiran origin, the evidence is still scant. This is therefore an enormous piece of the puzzle, which certainly *implies* a very close tie between Raetico (and by extension other pterosaurs) to the ornithodians.
Clearly there is still much to do. The paper out now is ‘simply’ an involved description and diagnosis and at the very least it’s systematic relationship to other pterosaurs needs to be determined, and further to that the possible implications for pterosaur origins. Other work is in preparation already on this taxon and Rico has kindly invited me to be involved with this work, so there is more to come. On the face of it, it does have characters that tie it to several other basal pterosaur clades and yet it is still very unusual. Rather like the anurognathids, it has some *very* basal features which are apparently combined with some highly derived features, and thus it could be a very basal taxon that is simply highly modified.
I am speculating (with an experienced eye) but I suspect it might just be deserving of its own family. It certainly has more autapomorphies than pretty much any other pterosaur I can think of off-hand given that it is perhaps only 50% complete. It’s systematic position is likely to be complex too because of the combination of features – it *could* just, possibly, turn out to be the most basal pterosaur known. It could also nest with the scaphognathines, or even collapse the rhamphorhyncoid tree to a bush. In short there is much to do here, and while it will be quite sometime before something is published on it’s systematic and phylogenetic affinities I suspect that pterosaur researchers are already plugging it into their matrices to see what happens.
For now though, the conclusions are simple. We have a great new specimen of a Triassic pterosaur that is well preserved, fairly complete and shows off a hatful of unusual features, both basal and derived. So, welcome Raeticodactylus to the literature, there are many people waiting to see what secrets you can reveal.
Edit: Without my knowledge, I have discovered that this post has been cross-posted to another site. For any visitors coming from there, please do explore the rest of the site. If nothign else, I have written two more pieces on Raeticodactylus!
Last edited by David Hone (2008-04-22 09:30:51)
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This is one amazing discovery. My first impression was that the skull looked suspiciously like a theropod rather than a pterosaur! That was just on first glance. Then, there's that crest and ventral keel...how bizare! I'm guessing as cranial ornamentation appear early and persist in pterosaur evolution, it must have been subjected to extensive sexual selection - or is it function?.
I haven't yet read the paper but from what you describe here, I am interested to know what the significance of a slender humerus is...
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I love this bugger, and I've already read the paper. I also wonder how it flew, not necessarily because of the long humerus, but because of how the patagia surfaces connect together. Without a sprawling hindlimb, one would think that the cheiropatagia would not have attached to the leg. Thus, my wonderance on my blog whether it caught fish with its feet.
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Hey David, what is your opinion regarding possibly synonymy with Caviramus? Reading the paper leaves me with the strong feelings that the few differences are very minor and can be accounted for by the ontogenetic age and preservational condition of the two specimens.
The keel looks structural to me. If so, we have something similar, but we call it a chin.
First off, Zach, sorry but I can see absolutely nothing in your fishing idea. Extant raptors have robust leg and foot bones (more so that terrestrial or just heavy flying birds) and of course the femur here is rather slender. Furthermore, you might expect the feet to be well spread out (either as a kind of reversed hallux kind of way, or just with a wide spread of the toes as seen in fishing bats), and we do not see this. The pedal claws also apprea to be small, blunt and not curved, exactly the opposite to what you mgith expect. Finally the mouth is full of odd teeth, and if it was catching and killing fish with its feet, why would it need a large heavy jaw and complex heterodont dentition if all it has to do is swallow? Why not have no teeth at all, rahter than big fangs and then these unusual unevenly spaced cusped teeth. And another finally, surely the most parsimonious explanation to all of this is jsut that is has an archosaurian femur, becuase its, well, and archosaur. Sorry to be soooo negative, but well, nothing matches up at all really.
I also don't see why a parasagital femur would prevent an leg or ankle attachment for the brachiopatagium. The patagium sweeps in close to the hindlimb and is highly elastic, it should not really affect the locotmotiuon of the animal on the ground. Look at flying squirrels and dermopterans.
Christopher: as for Caviramus, while I have not seen it Rico has. The drawins in the paper suggest that it has a different tooth set (look at the position of the formaina and sockets) and a different tooth count. I also doubt that it represents a juvenile and the shape of the articular is rather different. Finally, pterosaurs grow really isometrically, even in the head (excet of course for head crests) and so I would be surprised if the chnages were so dramatic late in development. It is possible they are the same, but I would be very surprised and there is nothing wrong with giving this it's own name.
Manabu: yes, it is an odd humerus. I might havre to try and persuade Ross to look into it at some point (lucky him). As I say it implies a weak wingstroke, but the rest of the humerus and the wing does not, perhaps it is less pneumatic that other ones and thus stronger? And while Raetico. is weird with the crests, we do see other Tirassic critters with them, notably Austriadactylus so bony crested pterosaurs are known to be very basal.
Nathan: I am not sure what you mean by 'it look structural'? It could be a genuine crest (i.e. a disaply feature) or of course a way of strengthing the lower jaw (though this does seem an odd thing to do really in the manner in which it has occured) though as I said, it does match the large articular.
Right, that's it for now. Expect the interview with Rico to go up in a day or two.....
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Thanks for your insights into the fishing idea, David. I knew it was a long shot, but it seems like Raetico must be doing something different with its legs!
Now, my patagium problem is how the animal would have flapped through the air if its main patagia attached to the thigh or outer ankle of a parasagittal wing. I wish I could post a picture to show you the problem!
Okay, wings flap on the lateral plane, but the legs are held beneath the body, folded up (like a bird) or hanging down (like...raptors). Because the legs are not sprawling as the arms are, how would they move WITH the motion of the arms? See? It's very hard to explain without being able to post a picture. You know what? I'll just sketch something and post it to my blog later today, so you can see my conundrum.
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Well I think we can be fairly certain that the brachiopatagium attached to the ankle in all pterosaurs (or at least in all that we have a good preservation of the wing membrane), so this would be a problem for all pterosaurs. It hardly affects bats through as they hold the legs out behind, and this is the consensus for pterosaurs. Just becuyase they don't have a parasagital femur, doesn't mean they can't raise the legs to a horizontal position. Pterosaurs have incredibly loose hip joints, and this should not be an especially difficult position for them to get their legs into.
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Well, I think bats have more of a naturally sprawling hindlimb posture, but I see your point. So the pterosaurs just held their legs back...now that I think about that, I think that's how Skrepnick restores his pterosaurs and it works just fine.
Crap! Should I be retrofit the patagium onto my Nyctosaurus' ankle? ARGH! It currently stops at his knee!
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Yeah, it doesn't really stop them, and dlook at vampires and mysticinus, they can *gallop*!
And actually don't worry about the Nyctosaurus. While there is no direct evidence for it, and based on parsimony one should keep with the ankle attachement, I think the nyctosaurids / pteranodontids might be the few taxa that reduced their attachement based on how extreme they were with their wings. It is unscientific speculation (naughty me) but I think reasonable enough, certainly when it coems to doing a reconstruction rather than a scientific paper, so keep it short if you want.
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