Rhamphorhynchus Wings

In 1882 Carl Zittel described a fossil pterosaur wing with the membrane preserved. This remarkable find was the highlight of that year and the Zittel wing became a very famous fossil. Casts were distributed to all of the main national museums and by 1883 most researchers and interested students of fossils had seen the wing.
In 1880, a fine fossil of Rhamphorhynchus was found in the Solenhofen Shales. The work was published in 1882 and clearly showed the wing membranes and tail fin. This specimen was originally named Rhamphorhynchus phyllurus (now Rh. muensteri) and it was sold to the Yale Peabody Museum in New Haven, USA. As with the Zittel specimen, casts were taken and distributed to National Museums.
Since the original finds of pterosaurs with wing membranes preserved, There have been many examples from the Lithographic Shales in Bavaria. The specimen above is in the Humboldt University Museum in Berlin.
Perhaps the best specimen of a Rhamphorhynchus with wing membrane preserved is the 'Dark Wing' specimen which is sometimes called the 'Tischlinger' specimen. This is a reflection of the work of Helmut Tischlinger who produced an amazing set of ultra-violet photographic images of this fossil to enable more detail of the wing to be observed.

These and other fossils enabled the wing structure to be studied in detail, from the fibres that run across the cord of the wing to the different layers of tissue within the wing membrane. As a result, there is a high level of understanding about the wing membrane structure of these and other pterosaurs.

von Zittel, K. A. 1882 Über Flugsaurier aus dem lithographischen Schiefer Bayerns. Paläontographica 29, 47–80 & pls 10–13.

Padian K & Rayner J M V; 1993, Structural fibres of the pterosaur wing: anatomy and aerodynamics. Naturwissenschaften 80: 361-364.

Martill D M and Unwin D M; 1989, Exceptionally well preserved pterosaur wing membrane from the Cretaceous of Brazil, Nature, 340:138-140

Tischlinger, H. and Frey, E. 2002. Ein Rhamphorhynchus (Pterosauria, Reptilia) mit ungewöhnlicher Flughauterhaltung aus dem Solnhofener Plattenkalk. Archaeopteryx, 20, 1-20.

Pterosaur skeletal sculptures

Bruce Mohn is a comparative anatomist, palaeontologist and Palaeo-Sculptor with a specialism in mesozoic vertebrates. He has recently completed skeletal models of pterodactylus and rhamphorhynchus for the Carnegie Museum and the quality of his work is outstanding.
The model components are crafted from observations of fossil material and investigative techniques to give a realistic life-like shape to each individual component of the skeleton. The assembly of the individual components uses wired supports to give an accurate representation of the complete skeletal structure of the animal. This technique allows the skeleton to be wired into any orientation or possition that was possible in life.
If you are able to visit the Carneige Museum, it is worth finding these skeletal models to view the 3D structure of these pterosaurs, which cannot be fully appreciated from diagrams.

Bruce is able to make bones or skeletons to order - this is fine bespoke craftsmanship of the best museum quality. His work can be seen at - Bruce Mohn sculptures at Dinoart

Bruce Mohn

Carnegie Museums

Was Rhamphorhynchus a skim feeder?

Whilst translating a passage from Peter Wellhofers' work on the Rhamphorhynchoidea, I came across this specimen from the collection of Karl Strobl.

Specimen No. 28: Fig. 24; Plate. 8, Fig. 1-3. Wintershof. Sammlung KARL STROBL, Eichstätt.
Part and counterpart of an entire skeleton with excellent preservation.

The individual is seen from the side; the arms are pushed forwards, the flight fingers in parallel with backbone extend backwards. Impressions of the flight skin and the narrow, lance shaped tail sail are present. The lower jaw extremity is relatively high, blunted in front. The upper edge line of the skull is concave. The front edge of the Infra-temporal opening is formed by the Quadrato-jugale. The cranial length measures 55.5mm.
The gastral skeleton consists of 6 curved bones still in situ; the middle pieces are displaced forwards. The Pubis is widened ventral, so that to the Ischium appears as a bay than rather a round opening. The Prepubis is strengthened centrally and hook-shaped (fig. 10 e).
Within the body cavity is a compressed fish tail and numerous single fish bone fragments indicating the stomach contents. Beside this lies 6-7mm long bananas-shaped small sausages like gastric stones of 1.5mm diameter and with zigzag-shaped textured surface (ref. fig. 44 b).

This specimen clearly shows that this individual had eaten a fish which was swallowed whole, head first. The use of gastric stones to aid mechanical digestion is an indication of a more specialised digestive system. This rhamphorhynchus may have had a gizzard, similar to that seen in birds, or a muscular and thick stomach wall to enable the churning of contents to allow the enzymes and gastric stones to break down the food both chemically and mechanically.

Very few pterosaurs give information about stomach contents, and in this case it leads to a question about fishing methods. Did rhamphorhynchus use a surface skimming technique to fish, or was it a shallow diver. It is impossible to say for certain, but the evidence is compelling. The jaws would certainly be able to scoop up a fish from near the surface of the water. and such a technique would have needed a very precise flying skill and good control of the jaws and neck.

The jaw on this specimen shows an impression of an upturned bony sheath at the end of the lower jaw. Such a structure, being free of teeth, would lend itself to surface skimming. This is a strong clue to the feeding habits of this type of Rhamphorhynchus and the group as a whole.

Wellnhofer, P. 1975 Die Rhamphorhynchoidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Teil II. Systematische beschreibung. Paläontographica A 148, 132–186.

The Rhamphorhynchoidea

In 1975, Peter Wellnhofer wrote a classic monograph on the Rhamphorhynchoidea. This set of 3 linked works has been the baseline for the study of this group of pterosaurs since then. It is a well organised work with analysis split into skeletal morphology, systematics and Palaeoecology. The texts were originally in German and as far as I am aware, they have not been translated wholesale into English. Recently, I started to translate part 2 for my own use. I would love to publish my translation, which will be a new work, but the original text is not out of copyright until 2025. I can only find paper copy, but I have been told that there is an electronic copy in German. This has eluded me, so I am transcribing and translating from the original text.

Wellnhofer, P. 1975 Die Rhamphorhynchoidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Teil I. Allgemeine Skelletmorphologie. Paläontographica A 148 , 1–33.11 plates.

Wellnhofer, P. 1975 Die Rhamphorhynchoidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Teil II. Systematische beschreibung. Paläontographica A 148, 132–186.

Wellnhofer, P. 1975 Die Rhamphorhynchoidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands. Teil III. Palökologie und Stammesgeschichte. Paläontographica A 149, 1–30. 13 plates

The problem in translating any text is the transfer of meaning. There are words in German that can have several possible meanings in English. This also works the other way around. The bulk of the meaning is transferable with a high level of confidence in the result, but there are some areas where the meaning has to be a best guess. This type of translation is often referred to as a Gist Translation. This means that in some cases, the reader may need to refer to the original text to grasp the full meaning of a sentance of passage. It is for this reason that translations should only be attempted from the original documents. Another of Peters texts that I have frequently used is the Handbook of Paleoherpetology, Part 19. This text is also only available in German, so people who are not able to read that language can only look at the wonderful line drawings and be amazed. This is of course, another text that sets the baseline for the study of pterosaurs in general. It is a little dated now, but still a valuable resource. The original text is still available to purchase on the Internet.

Wellnhofer, P. & Khun, O. 1978 Handbuch der Paläoherpetologie. Teil 19. Pterosauria. Stuttgart: Verlag Gustav Fischer.

These texts defined the study of pterosaurs for a whole generation of researchers. It would be beneficial to the subject to have them available on line both in their original German text and in English. The main difficulty is in gaining permission to freely allow access without infringing copyright law. Watch this space!

Discovering Pterosaur flight - Part 1

In the early days of research, Samuel Thomas von Soemmering made the first sensible reconstruction of a pterosaur skeleton in 1812. The reconstruction was missing several skeletal elements, but was a sound model based on the available evidence. He also proposed a wing membrane form which, although too generous in area was a good attempt at theoretical reconstruction.
It was not until 1882 that the wing structure was known from the fossil record. A specimen originally named Rhamphorhynchus phyllurus was discovered in the Lithographic Shale of Bavaria. At about the same time an isolated wing known as the Zittel wing (above) was published. Both fossils showed the outline and structure of the wing membrane of Rhamphorhynchus. This enabled much better reconstructions to be made.

In 1910 C. F. Eaton produced a paper on the osteology of Pteranodon with a skeleton reconstruction This was an inspired paper showing the skeletal form of a large pterosaur. Pteranodon was only known from crushed and distorted remains and this paper showed the skeletal form in a realistic way.

Ernst Stromer was inspired by the Rhamphorhynchus wing membranes and he produces a flying glider in 1913 which demonstrated that Rhamphorhynchus could fly with the known wing form.

By1914 the Hankin and Watson paper on the flight of Pteranodon gave us an insight into the structure of large pterosaurs that was related to the way they could fly. The framework was set for some real flying models.

Watch this space.......

von Soemmerring, S. T. 1812 Über einen Ornithocephalus oder über das unbekannten Thier der Vorwelt, dessen Fossiles Gerippe Collini im 5. Bande der Actorum Academiae Theodoro-Palatinae nebst einer Abbildung in natürlicher Grösse im Jahre 1784 beschrieb, und welches Gerippe sich gegenwärtig in der Naturalien-Sammlung der königlichen Akademie der Wissenschaften zu München befindet. Denkschr. k. bayer. Akad. Wiss. math.-phys. Kl. 3, 89–158, plates.

von Zittel, K. A., 1882, Über Flugsaurier aus dem lithographischen Schiefer Bayerns. Paläontographica 29, 47–80 & pls 10–13.

Eaton, G. F., 1910, Osteology of Pteranodon. Mem. Conn. Acad. Art. Sci. 2.pp.1-38.

Marsh, O. C., 1882, The wings of Pterodactyles. Am. J. Sci. (3)23, 251–256 & pl. 3

Stromer E., 1913, Rekonstruktionen des Flugsauriers Rhamphorhynchus gemmingi, H. v. M., Neues Jahrb. Min. Geol. Pal., II, pp. 49-68.

Hankin E. H., & Watson D. M. S., 1914, On the flight of pterodactyls. Aeronautical Journal 18, 324–335.