868


Journal of Paleontology 89(5):845–869


the context of the roughly planar dactylopatagium loaded in tension as in flight, actinofibrils would resist longitudinal compression and would be prevented from bending within the plane of the patagiumby the collagen fibers of the dorsal dermis. If actinofibrils consisted of collagen or elastin rather than keratin, if


they could “shorten or expand and therefore provide more flexibility to the wing membrane” as Kellner et al. (2010) suggested, then they would not have been able to resist longitudinal compression, would not redirect tension in the patagium, and would not prevent chordwise narrowing of the patagium under spanwise tension, and therewould no explanation for their radiating pattern in the dactylopatagium.


Conclusions


This paper has shown that the Zittel wing of Rhamphorhynchus, probably the most influential specimen preserving traces of the structure of the pterosaur patagium, has been misinterpreted since its first description, and actinofibrils were closely spaced


broad keratinous structures within the dorsal epidermis. The findings, when combined with evidence from other specimens, indicate that the dactylopatagium was narrow, inextensible, and somewhat stiff, which when combined with the streamlining of the pneumatic retrophalangeal wedge would produce a highly efficient airfoil more like that of sailplanes than the membranous wings of bats. That dactylopatagium was combined with a broad extensible plagiopatagium to produce a unique wing structure and planform. Various aspects of the present interpretation have been


suggested before: Wellnhofer (1975) interpreted the patagium as coarse, leathery, and tough; Padian and Rayner (1993) interpreted actinofibrils as keratinous; and Unwin and Bakhurina (1994) interpreted the dactylopatagium as stiff and relatively inelastic. However, the general acceptance of Zittel’s (1882) incorrect interpretation that the wing included widely spaced cylindrical fibers inhibited progress toward a reinterpreta- tion. It is interesting to consider thatMarsh’s (1882) interpretation of his specimen as preserving folds and wrinkles was correct, and yet for 130 years Marsh’s interpretation was largely ignored in favor ofZittel’s.Also interesting is the fact that Pennycuick (1988, p. 307) presented an essentially correct interpretation of the raised longitudinal strips of the Zittelwingwhile incorrectly attributing it toWellnhofer (1975) and rejecting it.


Acknowledgments


I thank P. Caruso, E. Frey, J. M. Hendrix, D. Kümpel, C. Palmer, C. J. Pennycuick, D. Schaller, H. Tischlinger, and P. Wellnhofer for discussion and/or other assistance. I thank H. Mayr, O. Rauhut, and R. Liebreich of the BSP, M. Kölbl-Ebert of the JM, U. Göhlich of the NHMW, E. Frey of the SMNK, M. Wilmsen of the SNSD, D. Brinkman of the YPM, and D. Kümpel for access to specimens under their care and assistance while studying them. M. Habib, D. M. Martill, K. Padian, C. Palmer, and D. M. Unwin reviewed earlier versions. This research was made possible in part by a Fort Hays State University Internationalization grant and a sabbatical semester from Fort Hays State University.


References


Ammon, L.v., 1909, Über ein schönes Flughautexemplar von Rham- phorhynchus: Geognostische Jahreshefte, v. 21, p. 227–228.


Barthel, K.W., 1978, Solnhofen: Ein Blick in die Erdgeschichte: Thun, Ott Verlag und Druckerei, 393 p.


Beckemeyer, R.J., and Hall, J.D., 2007, The entomofauna of the Lower Permian fossil insect beds of Kansas and Oklahoma, USA: African Invertebrates, v. 48, p. 23–39.


Bennett, S.C., 1995, The function of the structural fibers in pterosaur wings: Journal of Vertebrate Paleontology, v. 15(Supplement to #3), p. 19A.


Bennett, S.C., 1997a, Terrestrial locomotion of pterosaurs: a reconstruction based on Pteraichnus trackways: Journal of Vertebrate Paleontology, v. 17, p. 104–113.


Bennett, S.C., 1997b, The arboreal leaping theory and the origin of pterosaur flight: Historical Biology, v. 12, p. 265–290.


Bennett, S.C., 2000, Pterosaur flight: the role of actinofibrils in wing function: Historical Biology, v. 14, p. 255–284.


Bennett, S.C., 2001, The osteology and functional morphology of the Late Cretaceous pterosaur Pteranodon: Palaeontographica, Abteilung A, v. 260, p. 1–153.


Bennett, S.C., 2002, Soft tissue preservation of the cranial crest of the pterosaur Germanodactylus from Solnhofen: Journal of Vertebrate Paleontology, v. 22, p. 43–48.


Bennett, S.C., 2007,Asecond specimen of the pterosaur Anurognathus ammoni: Paläontologische Zeitschrift, v. 81, p. 376–398.


Bennett, S.C., 2013a, New information on body size and cranial display structures of Pterodactylus antiquus, with a revision of the genus: Paläontologishe Zeitschrift, v. 87, p. 269–289.


Bennett, S.C., 2013b, Reinterpretation of the wings of Pterodactylus antiquus based on the Vienna specimen: Journal of Vertebrate Paleontology, Program and Abstracts, v. 2013, p. 85.


Böhm, R., 1962, Beitrag zu den postnatalen Veränderungen der Lederhaut des Haushuhnes (Acta universitatis agriculturae, Brno), Sborník Vysoké školy zemědělské v Brně: Spisy Fakulty veterinární, v. 10, p. 95–100.


Bonde, N., and Christiansen, P., 2003, The detailed anatomy of Rhamphorhynchus: axial pneumaticity and its implications, in Buffetaut, E., and Mazin, J.-M., eds., Evolution and Palaeobiology of Pterosaurs: Geological Society Special Publications, v. 217, p. 217–232.


Bramwell, C.D., and Whitfield, G.R., 1974, Biomechanics of Pteranodon: Philosophical Transactions of the Royal Society London, v. B267, p. 503–581.


Brooks, A.N., MacCready, P.B., Lissaman, P.B.S., and Morgan, W.R., 1985, Development of a wing-flapping flying replica of the largest pterosaur: AIAA Paper, p. 85–1446.


Buisonjé, P.H.de., 1981, Ctenochasma porocristata nov. sp. from the Solnhofen Limestone, with some remarks on other Ctenochasmatidae: Proceedings KoninklijkeNederlandse Akademie vanWetenschappen, v. (B)84, p. 411–436.


Cowley, M., 1986, Quetzalcoatlus northropi: Model Aviation, v. 12, p. 84–90, 158, 160–162, 164.


Czerkas, S.A., and Ji, Q., 2002, A new rhamphorhynchoid with a headcrest and complex integumentary structures, in Czerkas S.J., ed., Feathered dinosaurs and the origin of flight: The Dinosaur Museum Journal, v. 1, p. 15–41.


Döderlein, L., 1929a, Über Rhamphorhychus und sein Schwanzsegel. Sitzungsberichte der Bayerischen Akademie der Wissenschaften: Mathematisch-naturwissenschaftliche Abteilung, v. 1929, p. 1–46.


Elgin, R.A., Hone, D.W.E., and Frey, E., 2011, The extent of the pterosaur flight membrane: Acta Palaeontologica Polonica, v. 56, p. 99–111.


Döderlein, L., 1929b, Ein Pterodactylus mit Kehlsack und Schwimmhaut. Sitzungsberichte der Bayerischen Akademie der Wissenschaften: Mathematisch-naturwissenschaftliche Abteilung, v. 1929, p. 65–76.


Frey, E., Tischlinger, H., Buchy, M.-C., and Martill, D.M., 2003, New specimens of Pterosauria (Reptilia) with soft parts with implications for pterosaurian anatomy and locomotion, in Buffetaut, E., and Mazin, J.-M., eds., Evolution and Palaeobiology of Pterosaurs: Geological Society Special Publications, v. 217, p. 233–266.


Frey, E., Tischlinger, H., Krüger, W., and Hone, D., 2007, Pterosaurier als Flugmaschinen - Bionische Forschung in der Paläontologie? Fossilien, v. 24, p. 79–84.


Goldfuss, A., 1831, Beiträge zur Kenntnis verschiedener Reptilien der Vorwelt: Nova acta Academiae caesareae Leopoldino-Carolinae germanicae naturae curiosorum, v. 15, p. 61–128.


Gregory, R.L., 1997, Knowledge in perception and illusion: Philosophical Transactions of the Royal Society, B, v. 352, p. 1121–1128.


Gross, W., 1937, Über einen neuen Rhamphorhynchus gemmingi. H. v. M. des Natur-Museums Senckenberg: Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, v. 437, p. 1–16.


Hildebrand, M., 1988, Analysis of vertebrate structure, 3rd ed., New York, John Wiley and Sons, Inc., 701 pp.


Page 1  |  Page 2  |  Page 3  |  Page 4  |  Page 5  |  Page 6  |  Page 7  |  Page 8  |  Page 9  |  Page 10  |  Page 11  |  Page 12  |  Page 13  |  Page 14  |  Page 15  |  Page 16  |  Page 17  |  Page 18  |  Page 19  |  Page 20  |  Page 21  |  Page 22  |  Page 23  |  Page 24  |  Page 25  |  Page 26  |  Page 27  |  Page 28  |  Page 29  |  Page 30  |  Page 31  |  Page 32  |  Page 33  |  Page 34  |  Page 35  |  Page 36  |  Page 37  |  Page 38  |  Page 39  |  Page 40  |  Page 41  |  Page 42  |  Page 43  |  Page 44  |  Page 45  |  Page 46  |  Page 47  |  Page 48  |  Page 49  |  Page 50  |  Page 51  |  Page 52  |  Page 53  |  Page 54  |  Page 55  |  Page 56  |  Page 57  |  Page 58  |  Page 59  |  Page 60  |  Page 61  |  Page 62  |  Page 63  |  Page 64  |  Page 65  |  Page 66  |  Page 67  |  Page 68  |  Page 69  |  Page 70  |  Page 71  |  Page 72  |  Page 73  |  Page 74  |  Page 75  |  Page 76  |  Page 77  |  Page 78  |  Page 79  |  Page 80  |  Page 81  |  Page 82  |  Page 83  |  Page 84  |  Page 85  |  Page 86  |  Page 87  |  Page 88  |  Page 89  |  Page 90  |  Page 91  |  Page 92  |  Page 93  |  Page 94  |  Page 95  |  Page 96  |  Page 97  |  Page 98  |  Page 99  |  Page 100  |  Page 101  |  Page 102  |  Page 103  |  Page 104  |  Page 105  |  Page 106  |  Page 107  |  Page 108  |  Page 109  |  Page 110  |  Page 111  |  Page 112  |  Page 113  |  Page 114  |  Page 115  |  Page 116  |  Page 117  |  Page 118  |  Page 119  |  Page 120  |  Page 121  |  Page 122  |  Page 123  |  Page 124  |  Page 125  |  Page 126  |  Page 127  |  Page 128  |  Page 129  |  Page 130  |  Page 131  |  Page 132  |  Page 133  |  Page 134  |  Page 135  |  Page 136  |  Page 137  |  Page 138  |  Page 139  |  Page 140  |  Page 141  |  Page 142  |  Page 143  |  Page 144  |  Page 145  |  Page 146  |  Page 147  |  Page 148  |  Page 149  |  Page 150  |  Page 151  |  Page 152  |  Page 153  |  Page 154  |  Page 155  |  Page 156  |  Page 157  |  Page 158  |  Page 159  |  Page 160  |  Page 161  |  Page 162  |  Page 163  |  Page 164  |  Page 165  |  Page 166  |  Page 167  |  Page 168  |  Page 169  |  Page 170  |  Page 171  |  Page 172  |  Page 173  |  Page 174  |  Page 175  |  Page 176  |  Page 177  |  Page 178  |  Page 179  |  Page 180  |  Page 181  |  Page 182  |  Page 183  |  Page 184  |  Page 185  |  Page 186  |  Page 187  |  Page 188  |  Page 189  |  Page 190  |  Page 191  |  Page 192  |  Page 193  |  Page 194  |  Page 195  |  Page 196  |  Page 197  |  Page 198  |  Page 199  |  Page 200  |  Page 201  |  Page 202  |  Page 203  |  Page 204  |  Page 205  |  Page 206  |  Page 207  |  Page 208  |  Page 209  |  Page 210  |  Page 211  |  Page 212