To paraphrase Dr. Robert T. Bakker in The Dinosaur Heresies: I remember the first time the thought struck me! There’s something very wrong with that toy Pteranodon. It was a Marx toy (Fig. 1) with big, floppy wings that could NOT have been folded away. The wings extended, like bat wings, to the hind limbs, and so had a very deep chord, a shape still in favor by most pterosaur workers. Even as a kid I knew birds and bats could fold their wings close to their bodies, as compact as possible. The Marx Pteranodon looked to be incapable of that. The pterosaurs in the Jurassic Park series suffered from the same deep-chord malady (Fig. 2).
Wings made pterosaurs the first vertebrates to achieve powered flight, but the shape of those wings has been argued about for decades. Most experts (Unwin and Bakhurina 1994; Elgin, Hone and Frey 2011, for instance) follow the traditional bat-wing model (Figure 3), with a deep trailing edge curving back to attach at the ankle. Such a wing is incapable of being folded away without sagging.
Another wing model emerged when Bennett (1987) reported on bicondylar and fused tail bones in Pteranodon that could move only up and down. He proposed a wing model in which trailing edge attached to the tail tip and so could be lifted and depressed like an airplane elevator (Figure 4).
At about the same time, Padian (1987) and Padian and Rayner (1993) argued against the bat-wing model in favor of a more bird-style model by attaching the trailing edge to the middle or posterior of the pelvis (Figure 4) with hind legs tucked in. Ironically, while arguing against the bat-wing paradigm, Padian and Rayner (1993) fell sway to its influence and invented more wing tissue aft of the elbow in their reconstruction than they showed in various fossils (see below).
Here’s the evidence for the traditional bat-wing model.
Elgin, Hone and Frey (2011) documented the latest evidence for pterosaur wing shape. Unfortunately, rather than firmly documenting their conclusions, they could only report, “A review of relevant pterosaur specimens … strongly suggests that the trailing edge of the wing extended down to the lower leg or ankle in all specimens where the brachiopatagium is completely preserved.” Strongly suggests???!!! I was hoping for something a little more fim. Here’s their evidence:
1. The “darkwing” Rhamphorhynchus was traced “as is” with a continuous tone deemed to represent a wing membrane that attached at mid thigh (Figure 5). Note that the left humerus was disarticulated and displaced posteriorly. Even so, the wing membrane trailing edge was directed precisely at the elbow, as in the Peters (2002) model. The strip of soft tissue extending down the thigh was on a level below the wing membrane, not continuous with it. The Zittel wing specimen of Rhamphorhynchus was not used by Elgin, Hone and Frey (2011) despite preserving a exquisite wing membrane that was extremely short at the elbow, supporting the Peters (2002) model.
2. The holotype Jeholopterus specimen (Figure 6) was traced only from the plate with no distinction made between the long pycnofibers (hairs) and wing membrane. An alternative reconstruction is offered here in which the counterplate is also employed and a more precise tracing of the soft tissues are documented separating the narrow wings from the long pycnofibers.
3. The Vienna specimen of Pterodactylus (Figure 7) was traced accurately, without much membrane posterior to the elbow. Rather than taking this example at face value (and it compares well with the Zittel wing, among others), Elgin, Hone and Frey (2011) invented the taphonomic malady “membrane shrinkage” to explain away the absence of wing membrane needed to fulfill their bat-wing paradigm. Nevermind that there was no “membrane shrinkage” elsewhere on the specimen. The Vienna specimen is animated below. It shows that if one adopts the “what you see is what you get” approach to evidence, the wing membrane extends and folds away very nicely indeed– as is!
4. The private specimen of Anurognathus (Bennett 2007a, Figure 8 ) was employed by Elgin, Hone and Frey (2011), but they did not use gray tones to delineate what they considered wing membranes. Rather three lines indicated it. They noted that no membrane attached to the ankle, but failed to note that the trailing edge they drew extended straight to the elbow, as in the Peters (2002) model. The DGS method revealed fourth wing phalanges largely buried in the matrix beneath the left foot and right tibia, revealed here.
5. Elgin, Hone and Frey (2011) employed Eosipterus (Figure 9) in which they traced patches of wing membrane not realizing that when one completes the puzzle, the results support the Peters (2002) wing shape model. Again, no material was found lateral to the ankles or shins.
6. The best example of the bat-wing model has always been Sordes pilosus (Sharov 1971, Figure 10). This is the only specimen in which wing material was preserved lateral to the ankles–but there’s more to this story! Elgin, Hone and Frey (2011) traced few details and overlooked the displaced radius (or ulna) that formed the trailing edge of their interpretation of the left wing (details here). It was indeed the left wing, but torn, displaced and preserved in such a way that the membrane continued between the ankles where it created the illusion of a uropatagium. Elgin, Hone and Frey also failed to note the continuation of the right wing trailing edge as it headed toward the metacarpophalangeal joint before turning back to the elbow and anterior femur, again as in the Peters (2002) model.
Elgin, Hone and Frey (2011) made several more mistakes when they created their reconstruction of the pterosaur wing, detailed in Figure 11. These workers, like many others before them, ignored and made excuses for clear “what you see is what you get” evidence supporting the Peters (2002) model in favor of the unsupportable, centuries-old bat-wing paradigm.
Now for the heretical view.
Peters (2002) argued for a “what you see is what you get”model and reconstructed a narrow trailing edge essentially stretched between the elbow and wing tip with a short fuselage fillet filling the gap between the elbow and mid thigh (Figures 13, 14). This wing shape is duplicated in every pterosaur in which wing material is known. Note how well it folds up to prevent damage. A deep-chord wing (at the elbow) cannot do that. Note that phalanx four either curves or is bent posteriorly, matching the curvature of the trailing edge of the tip caused by aktinofibrils separating like strips on a bamboo fan, stretching membrane between them. Note that the wing is aerodynamically taut only at full extension and that tension runs between the elbow and wing tip. The hind limb is independent.
Pterosaurs originated as flapping leapers, not as gliders. Those wings developed distally first, not as gliding membranes close to the body (contra Bennett 2008).
The bird-wing model for pterosaur wings reaches its acme in ornithocheirids like Arthurdactylus (Figure 14.) Not only does this wing shape create a nice airfoil like that of a sailplane, but it folds away to virtual invisibility!
There never has been evidence for any membrane lateral to the tibia. Please send any, if found. All available evidence supports the bird-wing model.
(There were also earlier rumblings about the correct orientation of the pteroid: forward and variable (Frey) vs. medial and fixed (Bennett 2007b, Peters 2009). Medial and fixed is correct, but we’ll take a look at that issue later.)
As always, I encourage readers to see the specimens, make observations and come to your own conclusions. Test. Test. And test again.
Evidence and support in the form of nexus, pdf and jpeg files will be sent to all who request additional data.
Bennett SC 1987. New evidence on the tail of the pterosaur Pteranodon (Archosauria: Pterosauria). Pp. 18-23 Currie, PJ and Koster EH, eds. Fourth Symposium on Mesozoic Terrestrial Ecosystems, Short Papers. Occasional Papers of the Tyrrell Museum of Paleontology, #3.
Bennett SC 2007a. A second specimen of the pterosaur Anurognathus ammoni. Paläontologische Zeitschrift 81(4):376-398.
Bennett SC 2007b. Articulation and Function of the Pteroid Bone of Pterosaurs. Journal of Vertebrate Paleontology 27(4):881–891.
Bennett SC 2008. Morphological evolution of the forelimb of pterosaurs: myology and function. Pp. 127–141 in E. Buffetaut & D.W.E. Hone (eds.), Flugsaurier: pterosaur papers in honour of Peter Wellnhofer. Zitteliana, B28.
Elgin RA, Hone DWE, and Frey E. 2011.The extent of the pterosaur flight membrane. Acta Palaeontologica Polonica 56 (1), 2011: 99-111 doi:10.4202/app.2009.0145 online pdf
Padian K 1987. The case of the bat-winged pterosaur. In: Czerkas, S.J. and Olson, E.C., eds, Dinosaurs Past and Present (Natural History Museum of Los Angeles County/ University of Washington Press, California/Seattle) Vol. II, pp. 65–81.
Padian K and RaynerJV 1993. The wings of pterosaurs. American Journal of Science 239-A, 91–166.
Peters D 2002. A New Model for the Evolution of the Pterosaur Wing—with a twist. Historical Biology 15:277-301.
Sharov AG 1971.New flying reptiles fro the Mesozoic of Kazakhstan and Kirghizia. Trudy of the Paleontological Institute, Akademia Nauk, USSR, Moscow, 130: 104–113 [in Russian].
Unwin DM and Bakhurina NN 1994. Sordes pilosus and the nature of the pterosaur flight apparatus. Nature 371: 62-64.