Pterosaurs of Stuttgart and Munich

As part of my PhD, and with the help of the Geological Association of London, I've been fortunate enough to go on several research trips to some museums in Germany including Tübingen, Karlsruhe, Stuttgart, and Munich. Stuttgart and Munich in particular have excellent pterosaur collections, including many historically significant specimens.

Stuttgart

The Staatsliches Museum für Naturkunde in Stuttgart (SMNS) is a large museum that houses one of the state collections of Baden-Württemberg (the other one being in Karlsruhe). The museum has a significant collection of material from the area, as well as some excellent material on display, but I'm only going to talk about some interesting pterosaur material. 

Campylognathoides

Campylognathoides zitteli was first found in 1858, from the Early Jurassic deposits of Württemberg. In 1893, a new specimen was discovered from the Holzmaden shale, and was named as a new genus and species, "Campylognathus zitteli", based on the specimen found below. It was later discovered that the genus was already in use for an insect, and so a new genus, Campylognathoides, was named in 1928.  

Type specimen SMNS 9787 of Campylognathoides zitteli

Austriadactylus

The type specimen of Austridactylus cristatus was found in the Alps of Austria, dating back to the Late Triassic. This is one of the earliest pterosaurs known so far as Triassic pterosaurs are extremely rare. They are also very poorly preserved, and Austriadactylus is a perfect example of this. Like many pterosaurs, it is completely crushed and incomplete, but some features can be seen. It has many primitive pterosaurian features including a long flexible tail (which lacks the stiffening rods seen in later pterosaurs) and heterodont teeth. 

Type specimen of Austridactylus cristatus SMNS 56342 

Line drawing of Austriadactylus cristatus from Dalla Vecchia et al. [1]

Miscellaneous

Also in the SMNS collections is some miscellaneous pterosaur material that shows some interesting features. Specifically, there are some 3D, beautifully preserved pterodactyloid fossils that show the internal structure of pterosaur bones, the delicate trabecular structure that existed in the heads, and the trabeculae that occur in the shafts.

Close ups of the internal structure of two pterosaur bones in the shaft (above) and head (below)

Munich

The Munich palaeontology collections exist in the Bayerisches Staatssammlung für Paläontologie ind Geologie (BSP), and are quite extensive, as shown by the large number of researchers there after the Society of Vertebrate Paleontology meeting in Berlin. It is especially great for pterosaurs, including numerous historically significant specimens, including the first ever pterosaur known to sciences, Pterodactylus antiquus. Unfortunately, that specimen is so valuable that it is kept out of prying eyes and is only accessible under specific permission, so I wasn't able to see it. However, that aside, there were many other significant specimens for me to see. 

 Aerodactylus

BSP AS V 29, type specimen of A. scolopaciceps

This genus has a complicated history. In 1860, a new species, "Pterodactylus scolopaciceps" was named by Meyer after being found in the Bavarian Solnhofen limestone from the Jurassic. This was later synonomised in 1883 with "P. kochi" which was thought to be a smaller species of Pterodactylus. However, over the years it has generally been agreed that "P. kochi" is just an ontogenetic stage of P. antiquus, meaning that they are just smaller, younger individuals. The original type specimen of "P. scolopaciceps" was more recently re-described as a new genus, Aerodactylus [2]. While all specimens of A. scolopaciceps are considered to be juvenile, it has been suggested there is enough of a difference to be a different genus.  

Beautifully preserved specimen of Aerodactylus

Close up of some details of the wing of Aerodactylus. Note the long slender pteroid bone, which is unique to pterosaurs

Germanodactylus

Germanodactylus cristatus was first described as a specimen of "P. kochi" by Pleininger in 1901 after being discovered in the Solnhofen lagerstätt, another Late Jurassic German pterosaur. It was then described as a new species of Pterodactylus, "P. cristatus". In 1964, a new genus Germanodactylus was named, and BSP 1892 IV 1 was named the type specimen of G. cristatus. The precise position of Germanodactylus within the Pterosauria has been debated, but it is definitely a pterodactyloid pterosaur. 

Type specimen of Germanodactylus cristatus, BSP 1892 IV 1

The Zittel Wing
By far, the most historically interesting specimen and biggest surprise for me came when I opened up a drawer and found the "Zittel Wing", a nearly complete Rhamphorhynchus wing. This wing was described by Alfred von Zittel in 1882. It was a significant find then, and to this day still represents one of, if not the best preserved wing membrane of a pterosaur. Nearly all the wing bones are complete, and the membrane is preserved from the tip of the wing finger (the elongated 4th finger) to underneath the humerus. It showed the width of the wing, as well as the actinofibrils, which are the strengthening fibres that provided the pterosaur wing with strength when they are overlain in criss-crossing layers. This was the first sign at how pterosaur wing membranes were formed.

The Zittel Wing

 3D ornithocheird wing bones!
For me, a highlight was looking at the 3D preserved specimens from the Early Cretaceous of Brazil. These were large pterosaurs, with wingspans of 5 m or so, and were part of the pterosaur revival of the 1970s to 1990s after being described in detail by Peter Wellnhofer. These bones are all very well preserved and have mainly been prepared out of the rock, meaning that you can pick them up, move them around, and really start to understand them. These were definitely the highlight for me!

Beautifully 3D preserved Santanadactylus spixi radius, ulna, and wing carpals.

Wing metacarpal with 3 other metacarpals from Santanadactylus pricei. Note the small pneumatic foramen (see previous post on pneumaticity for details) underneath the small finger metacarpals.

Those are my highlights from the natural history museums in Stuttgart and Munich. There were many more interesting specimens, and I could go on for ages about it, but I think I'll stop here!

References:

[1] Dalla Vecchia FM et al. 2002. A crested rhamphorhynchoid pterosaur from the Late Triassic of Austria. Journal of Vertebrate Paleontology 22: 196-199.

[2] Vidovic SU and Martill DM. 2014. Pterodactylus scolopaciceps Meyer, 1860 (Pterosauria, Pterodactyloidea) from the Upper Jurassic of Bavaria, Germany: the problem of cryptic pterosaur taxa in early ontogeny. PLoS ONE 9: e110646.

Pterosaur bone mass

The last post here was on pterosaur skeletal pneumaticity, and while I said I was going to continue this discussion in the next post, I'm going to take a side-road for a bit and talk about my first research paper, which has just come out! It's still related though, and ties in with these questions nicely.

Estimating pterosaur bone mass using CT scans

Summary of Martin and Palmer 2014 [1]

At the beginning of my MSc, my supervisor (Colin Palmer) and I wanted to look at estimating pterosaur bone mass using CT scans. Total mass of pterosaurs is a controversial topic, with different methods and authors coming up with very different results, which you can read about in a previous post (and this one) if you are interested. It is essential to accurately estimate mass in pterosaurs as they were the largest animals to ever fly, an mass is extremely important in flight. The key thing to know here is that one method for estimating pterosaur body mass relied on the relationship between skeletal mass and total mass in birds [2], and applied this relationship to pterosaurs by estimating skeletal mass geometrically (i.e. a long bone is a hollow cylinder) [3]. Colin was interested in using computed tomography (CT) scans to estimate bone mass, to see how different (if any) the mass would be using this method. I agreed that it would be an interesting project, and started on my MSc at the University of Bristol.

The basic principle was that by calculating the cross-sectional area of bone in several slices throughout the bone (approximately every 5-10 mm), bone volume (as in the actual volume of bony material) could be calculated through integration. Then, mass can be estimated by applying a density and multiplying by the volume. This was done for a number of bones, but we only published on 3 first wing phalanges (the first big finger bone in the wing, herein referred to as WP1). We were then able to compare the results directly to the previous method used by Witton [3] thanks to him kindly sharing his dataset with us (thanks again Mark!).

CT scans through a pterosaur wing bone showing the shaft (A,B) and proximal head (C,D) cross-sections. Top images show unmodified CT scans, bottom images show reconstructed cortical bone and removed matrix used for area calculations. Image from Martin and Palmer [1]

What we found was quite different from what we had expected. We generally assumed that the mass would be somewhat similar to what Witton found. However, we found that all three bones were about twice as heavy using this method as previous estimates, which made us wonder what that means for the rest of the skeleton.

Table indicating measurements from 3 WP1s including mass estimates. Note the differences between mass estimates in our method and in the previous method. From Martin and Palmer [1]

While there are differences between our method and Witton's original study, he could only do what was available to him, which for many reasons, did not include CT scans. However, he did suggest in his original paper that using CT scans would be another way to do this study and likely would be more accurate, so credit to Mark for that! It was a pragmatic method at the time, and well done using the materials available to him at that point.

So why is the mass so much more using the CT method? There are several possible explanations for this. First of all, the original method did not account for trabeculae, which did add 10-15% of mass in our study. Another explanation is that the cortical thickness used by Witton (which was calculated using a regression model derived by someone else) was consistently lower than what we found in the CT scans (see the table above), which also would affect the mass. And finally, one point that is related to the last one is that the original method did not account for the variation within the cortical thickness throughout the bone.

And what does this all mean? While this information, as well as some additional new data suggests that the wings of pterosaurs were heavier than previously estimated. This isn't really a big surprise when noted that some previous estimates suggest that the pectoral muscles (the muscles around the shoulder) in pterosaurs account for 30-40% of the total body mass [4]. While these muscles are mainly used for flight, they would also be the main muscles for take off if pterosaurs did take-off using their forelimbs to launch as has been suggested [5].

This study made us wonder what the rest of the skeleton would look like if we calculated it using CT scans, which has lead to my PhD project at the University of Southampton. The amount of bone tissue in the wing bones is related to both mass and pneumaticity, which are both subjects I am interested in, as they all related to the biomechanics and flight capabilities of pterosaurs. If anyone would like to see the paper and does not have access, let me know!

Next up, I'll talk about quantifying and comparing the amount of air (pneumaticity) found within the skeletons of pterosaurs, looking at different bones, and different pterosaurs, another paper that Colin and I have published on the topic.

Acknowledgements
Just wanted to say thanks to everyone that helped me with CT scans and along with this project that I am so happy is finally out! This includes: first and foremost thanks to Colin Palmer for putting up with me the last 2 years, and to Mark Witton for sharing lots of things along the way, and also Davide Foffa, Lorna Steel, Lauren Howard, Dave Martill, the staff at Muvis, Mike Habib, Emily Rayfield, and many more! I'm so happy to finally have this paper out :)
EDIT: Also, this should seem obvious, but I'm going to add it anyways. Many thanks goes out to my wonderful partner in crime Josh Silverstone for helping through the last 7 years (and especially the last 3), and for helping me with figures of course!

References
[1] Martin, EG and Palmer, C (2014) A novel method of estimating pterosaur skeletal mass using computed tomography scans. Journal of Vertebrate Paleontology 34: 1466-1469.
[2] Prange, HD et al. (1979) Scaling of skeletal mass to body mass in birds and mammals. American Naturalist 113:103–122.
[3] Witton, MP (2008) A new approach to determining pterosaur body mass and its implications for pterosaur flight. Zitteliana, Reihe B 28:143–158.
[4] Strang, KA (2009) Efficient flapping flight of pterosaurs. Ph.D. disserta- tion, Stanford University, Stanford, California, 295 pp.
[5] Habib, MB (2008) Comparative evidence for quadrupedal launch in pterosaurs. Zitteliana, Reihe B 28:159–166.