848
Journal of Paleontology 89(5):845–869
Figure 2. Marsh specimen of Rhamphorhynchus muensteri, YPM 1778. Nearly complete articulated skeleton and associated soft tissue traces of the patagia and tail vane in ventral view. Scale bar represents 3 cm.
new terms have not gained acceptance, but Wellnhofer (1987) and some subsequent authors (e.g., Bennett, 2000; Frey et al., 2003; Kellner et al., 2010) used the terms brachiopatagium, tenopatagium, and actinopatagium, though not necessarily in Schaller’s original sense. Schaller (2007) developed his ideas further and suggested that most pterosaurs had fore and hind wings supported by the fore- and hindlimbs, respectively. Wellnhofer (1987) described the Vienna specimen of
Pterodactylus antiquus (Sömmerring 1812) (NHMW 1975/ 1756/0000) from the Solnhofen Limestone, which is fully articulated and preserves traces of soft tissues of the body and wings. Wellnhofer noted fine parallel striations many of which were made visible by dendritic deposits of manganese and iron oxides in much of the patagium, interpreted the striations as part of the reinforcing system he proposed in his 1975 paper, and reconstructed the reinforcing fibers as present throughout the entire membrane spread between the forelimb, trunk, and hindlimb, which following Schaller (1985), he termed the brachiopatagium.Wellnhofer summarized Zittel’s observations of the raised longitudinal strips in the wing of Rhamphorhynchus though he referred to them as reinforcing fibers or rays (Verstärkungsfasern oder -strahlen), and repeated his earlier description of their size and spacing. He then proposed the term Aktinofibrillen (= actinofibrils), which he derived from Schaller’s term actinopatagium, for the supposed reinforcing fibers in the Vienna Pterodactylus and the Zittel wing, and presented a sche- matic reconstruction of the cross-section of the brachiopatagium with cylindrical actinofibrils dimensioned according to his
measurements from the Zittelwing (i.e., 0.05mmin diameter and spaced 0.2mm apart) and lying in the middle of the brachiopata- gium (Fig. 3.1). Pennycuick (1988) was not convinced that the raised
longitudinal strips of the Zittel wing represented solid fibers and instead interpreted them as epidermal wrinkles produced by contraction of internal elastic fibers running chordwise from the wing spar to the trailing edge. He noted that fractures across the strips showed no internal structure, and stated that:
“Although not disputing that the ridges seen in the fossils are composed of matrix material, Wellnhofer [1975] argues that such sharply defined relief could not have been produced unless the particles of matrix were compacted in between fibers of some hard material. He considers that the original fibres would eventually decay, leaving compacted ridges in thematrix.” (Pennycuick, 1988, p. 307)
Unfortunately, Wellnhofer made no such argument in his 1975 monograph nor did he make such a statement anywhere else (P. Wellnhofer, personal communication, 2012). Queries of Pennycuick and other colleagues have failed to produce an explanation or alternate source for Pennycuick’s statement. I wonder if perhaps the statement resulted from a misreading on Pennycuick’s part of the passage in which Wellnhofer (1975, p. 11–12) attributes the relative abundance of articulated Rhamphorhynchus wings to their being coarse, leathery, and tough. Pennycuick (1988, p. 308) also noted the parallel striations on the wings of the Vienna Pterodactylus as well as a “layer of
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