AN ANNOTATED AND ILLUSTRATED CATALOGUE OF SOLNHOFEN (UPPER JURASSIC, GERMANY) PTEROSAUR SPECIMENS AT CARNEGIE MUSEUM OF NATURAL HISTORY

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1 ANNALS OF CARNEGIE MUSEUM vol. 82, number 2, PP DEcEMBEr 2013 AN ANNOTATED AND ILLUSTRATED CATALOGUE OF SOLNHOFEN (UPPER JURASSIC, GERMANY) PTEROSAUR SPECIMENS AT CARNEGIE MUSEUM OF NATURAL HISTORY DaviD W. E. HonE School of Biological and Chemical Sciences, Queen Mary University of London, Mile End Road, London, United Kingdom MicHaEl B. HaBiB [Research Associate, Section of Vertebrate Paleontology, Carnegie Museum of Natural History] Keck School of Medicine, University of Southern California, Bishop Research Building, Room 403, 1333 San Pablo Street, Los Angeles, California MattHEW c. lamanna Assistant Curator, Section of Vertebrate Paleontology, Carnegie Museum of Natural History 4400 Forbes Avenue, Pittsburgh, Pennsylvania ABSTRACT We present an annotated and illustrated catalogue of all original fossils, casts, and sculpted replicas of pterosaur specimens from the Upper Jurassic Solnhofen limestones of southern Germany that are housed at Carnegie Museum of Natural History (Pittsburgh, Pennsylvania, U.S.A.). The museum obtained its substantial Solnhofen pterosaur fossil collection almost certainly the largest currently held outside of Europe as part of its purchase of the vast private collection of the Belgian Baron Ernest de Bayet in Original fossils include six partial to nearly complete skeletons and a beautifully preserved skull of the rhamphorhynchine Rhamphorhynchus muensteri (Goldfuss, 1831), two skeletons of the ctenochasmatid pterodactyloid Aurorazhdarcho micronyx (von Meyer, 1856), and one large pterodactyloid partial appendicular skeleton of indeterminate affinity. Three of the fossils preserve significant soft tissues, and one of these is also among the very few specimens of Rhamphorhynchus von Meyer, 1847 known to include probable gut contents (in this case, fragmentary fish remains). Two other Rhamphorhynchus fossils have been prepared nearly free of sediment in three dimensions. Despite the significance of the Carnegie Museum s Solnhofen pterosaur collection, it has, in general, been underutilized by the paleontological community. The primary purpose of this catalogue is therefore to increase awareness, and ideally study, of this scientifically and historically important collection of Late Jurassic flying reptiles. KEy WorDs: Archosauria, Aurorazhdarcho, Ctenochasmatidae, paleoecology, Pterodactyloidea, Rhamphorhynchidae, Rhamphorhynchus, soft tissue Pterosaurs are an extinct group of Mesozoic reptiles that were the first vertebrates to evolve powered flight. Although they have a rich fossil record that extends across all seven continents, their highly pneumatized skeletons mean that a great many of the best preserved and most complete specimens come from a few limited Lagerstätte deposits (Kellner 2003). Of these, the most famous and probably the most important are the Upper Jurassic Solnhofen and surrounding beds of southern Germany. The first pterosaurs to be recognized by science in the late 18 th century came from the Solnhofen (Wellnhofer 2008) and hundreds of pterosaur specimens have now been recovered from these limestone beds. Important new material continues to be discovered and described (e.g., Frey et al. 2003, 2011; Tischlinger 2010; Frey and Tischlinger 2012; Bennett 2013). The Solnhofen remains a critical part of our understanding of pterosaur evolution for three key reasons. First, it represents part of the major evolutionary turnover in pterosaurian biotas as the more derived pterodactyloids begin to supersede more basal forms both are present in the Solnhofen, but the basal forms INTRODUCTION had vanished by the end of the Early Cretaceous at the latest. Second, material is available in large numbers, allowing assessments of ontogenetic and intraspecific variation. Finally, since the Solnhofen beds are historically the earliest pterosaur-bearing sediments, and remain some of the most productive, a considerable quantity of research has been based on material from these deposits, and many of the taxa that they have yielded are among the best-studied pterosaurs. Solnhofen pterosaurs were long prized for their often exceptional quality of completeness and preservation, with some specimens even including soft tissues. As such, many of these were sold by dealers to nations outside of Germany, and consequently, specimens came to reside in collections as far from Bavaria as Budapest, Copenhagen, London, Dublin, Uppsala, New York, Pittsburgh, and Tokyo. This historic material continues to enter the literature as it is rediscovered in collections and its scientific significance is appreciated (e.g., Ősi and Prondvai 2009; Ősi et al. 2010; Hone 2012; Vullo et al. 2012). Here we briefly describe a series of original fossils,

2 166 annals of carnegie MusEuM vol. 82 Table 1. Taxonomic identity and selected measurements (mm) of CM Solnhofen pterosaur fossils. CM was deemed too fragile to measure. Listed genus and species follow the CM specimen database ( See Supplementary Table (online) for additional measurements. Abbreviations: MC, metacarpal; n/a, not applicable (element not preserved); Ph, phalanx. Specimen Listed genus Listed species Revised genus Revised species Skull length Skull mid-orbit CM Pterodactylus elegans Aurorazhdarcho micronyx CM Pterodactylus sp. Aurorazhdarcho micronyx CM Rhamphorhynchus muensteri Rhamphorhynchus muensteri 95.6 unmeasurable CM Rhamphorhynchus longiceps Rhamphorhynchus muensteri CM Rhamphorhynchus muensteri Rhamphorhynchus muensteri CM Rhamphorhynchus? sp. unidentified unidentified n/a n/a CM Rhamphorhynchus carnegiei Rhamphorhynchus muensteri 63.3 unmeasurable CM Rhamphorhynchus cf. longicaudus Rhamphorhynchus muensteri 42.3 unmeasurable CM Rhamphorhynchus gemmingi Rhamphorhynchus muensteri (distorted) Table 1. continued Specimen Humerus length Humerus midshaft Ulna length MC IV length Ph IV-1 length Femur length Femur midshaft CM unmeasurable (left) 19.7 unmeasurable CM (right) ~ (left) 48.7 (left) CM (right) 5.4 (right) (right) 29.1 (left) 2.4 (left) CM (left) (left) (left) 28.6 (left) 4.2 (left) CM broken 85.3 n/a n/a CM CM (left) (left) 21.2 (left) CM (left) broken broken CM n/a n/a n/a n/a n/a n/a n/a casts, and models of Solnhofen pterosaurs in the collection of Carnegie Museum of Natural History (Pittsburgh, Pennsylvania, U.S.A.). Although almost all of these were obtained more than a century ago, collectively they have been only rarely mentioned in the literature, and consequently little information has been published about them. With ten original specimens (many of which constitute nearly complete, articulated skeletons), five casts (not including casts of one of the Carnegie s own specimens), and three modeled skeletons, this represents perhaps the most important collection of Solnhofen pterosaurs outside of Germany. Indeed, a century ago, the Carnegie Museum s Solnhofen pterosaur collection would have been one of the most extensive in the world (including Germany); that it remains important attests to the quality and quantity of specimens contained therein. Recent reassessments of the taxonomy of Solnhofen pterosaurs (e.g., Bennett 1995, 1996, 2013; Jouve 2004) require that the identity of specimens be reexamined carefully. By providing descriptions, photographs, and measurements of this material herein, we hope to facilitate future studies of this important collection. HISTORICAL CONTEXT The original pterosaur fossils detailed below were acquired by Carnegie Museum of Natural History (then known as the Carnegie Museum) in 1903 as part of the museum s purchase of the enormous private fossil collection of the Baron Ernest de Bayet of Brussels, Belgium (Kay 1951, 1955; Dawson and McGinnis 1972; Wellnhofer 1973; Mc- Ginnis 1983). The Baron put his collection up for sale for US $25,000, reportedly to generate funds needed to support his purchase of a chalet on Lake Como, Italy, which had been requested by his recently-wedded bride (Dawson and McGinnis 1972; McGinnis 1983; Rea 2004). Carnegie Museum paleontologist John Bell Hatcher learned of the availability of the collection in October 1902, and notified the museum s director, William Jacob Holland, who in turn convinced Pittsburgh industrialist and Carnegie Museum founder Andrew Carnegie to underwrite the purchase. The sale was finalized in the early summer of 1903 for the reduced price of $20,500 (worth approximately $540,000 in modern currency), with the museum also agreeing to pay for packing and shipping (Dawson and McGinnis 1972). Accompanied by his son, Holland

3 2013 HonE Et al. solnhofen PtErosaurs at carnegie MusEuM 167 travelled to Brussels to personally supervise the packing efforts (Rea 2004). Numbering into the tens of thousands, the fossils, contained within more than 250 cases, arrived in Pittsburgh in September 1903, and were initially stored in a rented warehouse due to a shortage of space in the museum itself (Dawson and McGinnis 1972; Carter 1973; Rea 2004). Among them were European plant, invertebrate, and vertebrate fossils that collectively spanned most of the Phanerozoic, including especially rich assemblages from the Jurassic of Holzmaden (Germany) and Cerin (France), the Eocene of Monte Bolca (Italy), and the Solnhofen beds. Many of these specimens come from localities that are no longer accessible (McGinnis 1983). Within the vast Solnhofen collection were the ten pterosaur fossils detailed below. Although the Carnegie Museum s Solnhofen pterosaur collection was studied by German paleontologist Peter Wellnhofer during a six-month residency at the museum in 1972 (Wellnhofer 1973), and all specimens have been at least briefly mentioned in previous works (e.g., Koh 1937; Wellnhofer 1970, 1975; Padian and Rayner 1993; Witmer et al. 2003; Chatterjee and Templin 2004; Padian et al. 2004; Bennett 2013), the assemblage has never been comprehensively reviewed in the literature. THE CATALOGUE Our descriptions of the material herein are deliberately brief. Solnhofen pterosaur taxa have been described extensively in the past (e.g., Wellnhofer 1970, 1975; Bennett 1995, 1996, 2007a, 2007b, 2013; Frey et al. 2003, 2011) and therefore detailing or revising the anatomy of these forms is not necessary. Our intent is instead to inform researchers of the type and quality of the material curated at Carnegie Museum of Natural History, and by extension, its suitability for various avenues of pterosaur research. Specimens are ordered by taxon rather than by catalogue number. Fundamental measurements of all specimens are provided in Table 1, with additional details presented in the Supplementary Table ( Supplementary_and_additional_files_of_Carnegie_pterosaurs_/780794). Institutional abbreviations. BSPG, Bayerische Staatssammlung für Paläontologie und Geologie, Munich, Germany; CM, Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, U.S.A.; GPIB, Geologisch- Paläontologisches Institut der Universität Bonn, Bonn, Germany; NHMW, Naturhistorische Museum, Vienna, Austria; NMB, Museum für Naturkunde der Humboldt- Universität Berlin, Berlin, Germany; RGM, Nationaal Natuurhistorisch Museum, Leiden, The Netherlands; SMNK, Staatliches Museum für Naturkunde Karlsruhe, Karlsruhe, Germany; YPM, Yale Peabody Museum, New Haven, Connecticut, U.S.A. CM Original Fossils Taxonomic identity. Rhamphorhynchus muensteri (Goldfuss, 1831). Previous mentions in the literature. Koh (1937); Wellnhofer (1973, 1975 [his example 59 ]); McGinnis (1983:52 53); Padian and Rayner (1993); Faux and Padian (2007). Description. Large (wingspan approximately 1040 mm), articulated and nearly complete non-pterodactyloid specimen exposed mostly in ventral view (Fig. 1). The large size of the animal and its well-ossified sternum, mature humeral shape (sensu Bennett 1995), and fused pelvis all indicate that it represents an adult individual. The surrounding matrix is very pale gray with yellow patches, with the bone mostly being light brown. Some bones are in good condition and even preserved in three dimensions, but most have a damaged cortex. Other elements are badly broken or missing entirely, existing only as impressions in the matrix. Low-angle lighting reveals that a colorless preservative has been applied to the bones and matrix. The skull is visible in left lateral view and poorly preserved. The anterior part is damaged and the posterior even more so, with much of the bone missing or the cortex lost. The mandible is articulated with the skull in a natural position. Some teeth are present in the jaws and project anteriorly. A partial sclerotic ring appears to be preserved in the orbit, but this is also in poor condition. The cervical series appears to be complete and articulated but is poorly preserved. In addition, repair work on a break across the line of the cervicals has likely concealed additional parts of the series. The dorsal sequence is articulated and apparently complete but also in poor condition. The posterior dorsals are partially covered by sediment. Given the good preservation of the sacrals, the most posterior dorsals may be more intact than are the more anterior parts of the axial series. The dorsal ribs are mostly articulated with their corresponding vertebrae, though some have been dislocated. The gastralia have been slightly displaced from their natural positions and lie in series between the right elbow and knee. The sacrals are articulated and complete as a series; they are much better preserved than the preceding vertebrae. Their lateral processes are fused into broad sheets of bone that in turn are co-ossified to the ilia. The tail is incomplete as the slab is broken but at least the anteriormost ten caudal vertebrae are articulated to each other and to the sacrals. Posteriorly the caudals are well preserved but anteriorly they are present only as impressions. The elongate chevrons have partly separated from the caudal vertebrae and some lie at an angle from the main series. The left scapula and the proximal part of the left coracoid are present as impressions in the matrix. The pectoral elements on the right side cannot be seen. The left forelimb

4 168 annals of carnegie MusEuM vol. 82 Fig. 1. CM 11427, Rhamphorhynchus muensteri in ventral view. Scale equals 10 cm with 1 cm divisions. Abbreviations: L, left side of specimen; R, right side of specimen. is in poor condition and the humerus is damaged although the outline of the bone is clear. The radius and ulna are barely preserved proximally and their more distal parts are missing. Of the distal left forelimb, only parts of wing phalanges 1 and 2 remain, with the wing metacarpal, wing phalanges 3 and 4, and digits I III being absent. Wing phalanx 1 is broken and the distal part of phalanx 2 is cut off at the edge of the slab. The proximal part of wing phalanx 1 lies over the skull, suggesting that the wing metacarpal and other parts of the manus may originally have been present but were lost in an effort to expose the skull. This interpretation is supported by evidence of repair around the proximal part of wing phalanx 1 where it abuts the skull, suggesting that the wing elements were removed and replaced later. The right forelimb is more complete than the left, though many of its component elements are damaged. The humerus is fragmentary and the head is missing, and the radius and ulna are preserved primarily as impressions in the matrix. The carpus is an indistinct mass of bone and perhaps calcite, but the pteroid is clearly preserved. There are traces of at least two of the first three metacarpals and a shallow impression of the wing metacarpal. An indistinct patch of matrix around the distal part of the wing metacarpal may correspond to impressions of digits I III but this is not clear. All four wing phalanges are present and articulated, though the first is badly damaged. Both ilia are present, though the left is incomplete, and these appear co-ossified with the sacrals. Part of the right ischium is fused onto the ilium and projects from the slab as it is preserved in three dimensions, but is also broken. Two spar-like fragments of bone anterior to the ilia probably represent parts of the prepubes, but no details can be resolved. Both hind limbs are well preserved although the distal elements of the left pes are absent. Otherwise both of these limbs are complete, articulated, and preserved largely in three dimensions. Padian and Rayner (1993: table 1) intimated that the wing membrane of CM is discernable via grooves or impressions in the matrix associated with the specimen. This is almost certainly correct. Immediately posterior to the right wing finger, the slab is marked by a series of faint, posterodistally-oriented linear features that strongly resemble the wrinkles or folds in the wing membrane observed in better-preserved fossils of Rhamphorhynchus von Meyer, 1847 (e.g., Marsh 1882; Zittel 1882; Padian and Rayner 1993; Frey et al. 2003). Furthermore, the posterior margin of this wrinkled area is aligned precisely with the distal tip of wing phalanx 4, as would be expected for an impression of the wing membrane.

5 2013 HonE Et al. solnhofen PtErosaurs at carnegie MusEuM 169 Fig. 2 CM 11428, Rhamphorhynchus muensteri. Scale equals 10 cm with 1 cm divisions. A pair of very small, dark brown elements close to the gastralia appear to be osteichthyan scales or teeth. A mass that may correspond to additional fish parts lies between the gastralia and dorsal ribs; these include a putative series of vertebrae, a fin ray, and a series of small discs that may be scales. Comments. No original taxonomic identification was given on the specimen, but in the Carnegie Museum of Natural History collections database it is listed as Rhamphorhynchus muensteri, an assignment that was also provided by Wellnhofer (1975). We follow that identification here, because the presence of edentulous jaw tips, anteriorly directed anterior teeth, a femur that is shorter than the humerus, and a first wing phalanx that is approximately the same length as the skull (though not the longest phalanx in the series) clearly identify this as a specimen of Rhamphorhynchus (Bennett 1995; though see also Hone et al. 2012). Moreover, here and below, we follow Bennett (1995) in considering all known specimens of Rhamphorhynchus to be referable to R. muensteri, with smaller supposed species representing earlier ontogenetic stages of this taxon. An associated label identifies the specimen as having a Dpt. No and notes that it came from the Bayet collection. Two old labels on the reverse of the specimen are nearly unreadable except that one contains only the number 4. The most interesting aspect of this specimen is the putative fish remains associated with the ribcage. Although these are difficult to identify with certainty, there is a mass that is unlike anything else on the slab and that appears to be composed of fossilized organic remains that resemble fish scales and bones (see Supplementary Figure at This lies between the gastralia and ribs and is clearly not derived from the pterosaur. As such, it most likely represents fish remains in the gut of this Rhamphorhynchus individual. Several Rhamphorhynchus specimens with fish as gut contents are already known (Wellnhofer 1975: fig. 44; Unwin 2005; Frey and Tischlinger 2012), but gut contents have not previously been reported in CM Examination by a paleoichthyologist may reveal the identity of the fish in this Carnegie Museum pterosaur. CM Taxonomic identity. Rhamphorhynchus muensteri. Previous mention in the literature. Wellnhofer (1975 [his example 83 ]). Description. The specimen is of a large (wingspan approximately 960 mm) non-pterodactyloid pterosaur (Fig. 2). The size of the specimen and the shape of its skull both suggest that it corresponds to an adult individual (Bennett 1995). The specimen is preserved on a large, heavy slab of light gray limestone with yellow streaks, with the bones being a pale brown color. The skeleton is partly disarticulated and largely preserved in three dimensions but the condition of individual bones varies. Some are preserved only as impressions in the matrix, while others are split such that the interior of the bone is exposed but its external surface is not. The elements preserved are the skull and

6 170 annals of carnegie MusEuM vol. 82 Fig. 3. Interpretive line drawing of CM 11428, Rhamphorhynchus muensteri, to show identities and positions of various elements. Abbreviations: cdv, caudal vertebrae; cv, cervical vertebrae; dr, dorsal ribs; dv, dorsal vertebrae; fe, femur; ps, pes; sk, skull; ti, tibia; wpx, wing phalanx. articulated mandible, a partially articulated cervical series, the articulated posterior cervical and dorsal vertebrae, a nearly complete caudal series, both wings (one complete and fully articulated, the other almost complete and partly articulated), both femora and tibiae, and scattered ribs, gastralia, and pedal elements (Fig. 3). The skull is exposed in right lateral view and is articulated with the mandible, with the jaws wide open. The skull is in excellent condition and is prepared in relief, such that its three dimensional preservation is evident. Further preparation is probably possible on the skull, and would likely reveal additional details, especially of the palate, which appears to be present. The teeth are mostly missing, with only one large premaxillary tooth, the remnants of two smaller maxillary teeth, and one curved tooth in the anterior part of the left dentary in situ. The presence of matrix in and around the alveoli suggests that most teeth were lost prior to burial. At least two of these dissociated teeth are preserved as impressions in the vicinity of the cervical and dorsal vertebrae. Four articulated but poorly preserved cervical vertebrae are present in the middle of the slab and visible in right ventrolateral view. Nearby is an extensive axial series; exposed in ventral view, it consists of two cervical vertebrae, one indeterminate element that is most likely the first dorsal, and then 11 dorsals. A piece of bone representing a further possible cervical is preserved close to the distal tail. At least three ribs are articulated with their corresponding dorsal vertebrae, and approximately 15 other ribs and gastralia are scattered on the slab, though most are represented only by impressions. The tail is poorly preserved, with some parts (including the tip) missing and others covered with calcite. At least 25, and possibly up to 30, caudal vertebrae are present, suggesting that the majority of the tail is preserved (Wellnhofer [1975] states that Rhamphorhynchus possesses up to 40 caudals). Most limb elements are jumbled, with a number of bones lying over each other. One wing (it is not clear if this is the left or right) is complete and although slightly disjointed is in articulation. The proximal end of the humerus is broken off, but the bone is otherwise largely complete. The articulated radius and ulna have lost much of their cortices and calcite is visible in their interiors. The wing metacarpal is complete and well preserved. None of the other metacarpals of this limb can be seen. Wing phalanx 1 is likely complete but largely hidden below other elements; its proximal and distal ends are visible and in good condition. Wing phalanges 2 4 are present but in poor condition and the distal part of phalanx 4 is missing. Almost all of the elements of the other wing are also present. Its humerus is more complete than that of the wing described above, but also lacks much of the proximal end and is crushed and covered by calcite. The radius and ulna are complete and articulated but in poor condition. The wing metacarpal is largely hidden under other elements and matrix but its distal end is visible and in good condition. None of the other metacarpals or digits I III can be seen. Wing phalanx 1 has suffered extensive damage and has been repaired with glue. This glue has stained the bone a dark color and has also made the surrounding matrix and parts of adjacent elements darker. Wing phalanges 2 4 are disjointed but articulated and lie over the tail. All of phalanx 2, the distal

7 2013 HonE Et al. solnhofen PtErosaurs at carnegie MusEuM 171 Fig. 4. CM 11429, Rhamphorhynchus muensteri in primarily right lateral view. Scale equals 10 cm with 1 cm divisions. Abbreviations: cdv, caudal vertebrae; cv, cervical vertebrae; dv, dorsal vertebrae; hu, humerus; L, left side of specimen; ps, pes; sk, skull. part of phalanx 3, and all that can be seen of phalanx 4 (the distal part of which is missing) are presented in cross section along the long axes of the elements, as the bones have been split and filled with matrix. An articulated femur, tibia, and possible astragalus lie together and are also visible as longitudinal cross sections. An impression of what may be a metatarsal lies across the femoral head. A second femur and tibia lie close to the jumble of wing elements, with the proximal end of the femur partially hidden under the distal end of phalanx 2 of the fully articulated wing. Close to this second hind limb are impressions of three elements that likely represent pedal digit V, given the short proximal element and the very long and straight middle element. Comments. A Carnegie Museum label with the specimen identifies it as Rhamphorhychus [sic] longiceps Smith Woodward, 1902, an identification that was also given by Wellnhofer (1975). Here we refer the specimen to R. muensteri, due to its possession of edentulous jaw tips, anteriorly directed anterior teeth, an antorbital fenestra that is larger than the external naris but considerably smaller than the orbit, a humerus that is longer than the femur, and a first wing phalanx that is the longest in the series and that is subequal in length to the skull (Bennett 1995). The museum label indicates that the skeleton was purchased as part of the Bayet collection in 1903, though no locality information is given apart from Solnhofen, W. Germany. An aged label affixed to the specimen states Acc (the Carnegie Museum s accession number for the Bayet collection) and Dept. 2477, though the latter has been struck through by hand and replaced with 11428, suggesting that the fossil was re-catalogued at some point. The origin of the Dept number is unknown: it may have been a notation of Bayet s collection, but we consider this unlikely as the label is clearly from the Carnegie Museum. Instead we suggest that it may have been a number used by Holland or other Carnegie personnel in cataloging the material before it was given a formal CM specimen number. Also included with the specimen is a cryptic piece of paper that states only #16. This is clearly very old but there is nothing associated with the specimen to indicate what this might refer to or why it has been kept. We suspect that it may be an original number applied by the Baron de Bayet. On the reverse of the specimen there is evidence of a blue-edged label having been removed. The position of the mandible in a gaping position (with the long axis of the mandible forming close to, or even exceeding, a right angle with that of the maxilla) is not uncommon in specimens of Rhamphorhynchus. It is also seen, for example, in CM and CM (see below), as well as in some specimens in other collections such as BSP 1884 XI 1 and SMNK PAL The cause of this condition is not known. Although the opisthotonic posture hypothesis (see, for example, Faux and Padian 2007) attempts to explain the contorted posture and position of the joints in some fossil vertebrate specimens, opisthotony is unlikely to be the case here. In addition to the controversy regarding this hypothesis (see Reisdorf

8 172 annals of carnegie MusEuM vol. 82 and Wuttke 2012), specimens such as BSP 1884 XI 1 are otherwise in a normal posture. Furthermore, some Rhamphorhynchus specimens that Faux and Padian (2007) considered to have adopted an opisthotonic posture (e.g., CM 11427) have closed jaws. CM Taxonomic identity. Rhamphorhynchus muensteri. Previous mentions in the literature. Koh (1937); Wellnhofer (1973, 1975 [his example 53 ]); Padian and Rayner (1993). Description. This is a large (wingspan approximately 800 mm) non-pterodactyloid pterosaur that is preserved largely in three dimensions and that has significant soft tissues present (Fig. 4). The size of the specimen and especially the shape of the preserved tail vane suggest that this is a subadult animal that was close to maturity at the time of death (Bennett 1995). Much of the specimen is preserved as impressions in the matrix, or with heavy calcitic buildup on the elements. A partial counterslab is present and has been affixed to the main slab, and this contains parts of the left wing and probable left hind limb. The matrix is a pale fawn color with the bones being light brown and the soft tissues orange. The skull is poorly preserved and primarily visible in right lateral view. However, the bones appear to have been split between the slab and a missing part of the counterslab, such that the bones of the left orbital region are visible in medial view. The rostrum has also been twisted such that it is presented in ventral view, and teeth protrude from both sides of the upper jaw. A number of these are present, but most are represented only by impressions in the matrix. The mandible is articulated with the skull such that the jaws are widely agape (as seen in a number of other Rhamphorhynchus specimens, including CM and CM 11431; see above), but only its posterior part is preserved as bone. The remainder of the mandible is represented by impressions in the matrix, though some teeth are present anteriorly. The cervical series is almost entirely obscured by the mandible, and what is visible is poorly preserved and little detail can be discerned. The dorsal series is preserved in left lateral view but can be seen only as impressions in the matrix. A number of ribs are present. The anterior ribs are articulated with the dorsal vertebrae while closer to the posterior part of the series the ribs are disarticulated and lie in a variety of orientations alongside several gastralia. A single large excavation between the dorsals and the base of the tail is interpreted as remnants of the sacrum. Based on the extent of the tail vane, the caudal sequence is probably complete, but its posteriormost part is obscured under a large calcite crystal. Part of the posterior tail is preserved only as an impression in the matrix, and the majority of the centra are obscured by calcite, although the chevrons and at least some of the elongate zygapophyses are visible. The left forelimb is largely present but generally in poor condition. The articulated radius and ulna are present on a large block of matrix that lies ventral to the dorsal series. These are in turn articulated with a poorly preserved mass of bone and calcite that corresponds to the articulated carpal block and wing metacarpal. Close to the distal end of the left wing metacarpal are impressions of manual digits II and III. Along with manual digit I, these two digits are well preserved in the counterslab. All four left wing phalanges are present and articulated, although the bones are split between the main slab and the counterslab, and the distal tip of phalanx 4 may be missing. A twisted shaft of bone that lies proximal to the right humerus is identified as the middle part of a scapular blade. The right forelimb is represented by a nearly complete but obscured and broken humerus, the radius and ulna, and part of the wing. The radius and ulna are complete and well preserved, though the carpus and metacarpus are hidden below matrix and parts of the left wing. Only parts of right wing phalanx 1 are preserved, and this bone courses alongside the left wing (as seen in the dark wing specimen of R. muensteri; Frey et al. 2003), suggesting that the wings are folded together. However, the rest of the wing finger is missing from the slab where this is broken and these parts are lost. One hind limb is partially preserved as the distal end of the femur, the tibia, and the pes. The distal femur and proximal tibia lie on top of the corresponding regions of the right humerus and radius/ulna, respectively. The position of this knee atop the right wing strongly suggests that this hind limb is from the left side of the animal, but this is difficult to definitively determine given that some parts of the skeleton are disarticulated or rotated. As preserved in the slab, the tibial midshaft is represented only by an impression, while its distal part is bone. The pes is damaged but largely complete. The pedal elements are mostly present as impressions in the slab; as might be expected, however, most parts of the tibia and pes that are missing from the slab are preserved as bone in the counterslab. A large piece of indeterminate bone lies just below the tip of the mandible. This does not appear to be any part of the pterosaur, and may therefore represent a chance association of some part of another animal. The whole specimen is surrounded by a halo of orange stains (Fig. 4) that is remarkably similar to that seen in another Solnhofen Rhamphorhynchus specimen and that was interpreted as partial remnants of the soft tissues of the animal (Hone 2012). We favor the same interpretation for these orange stains in CM Two observations support this: first, the tail vane is clearly preserved and is of a similarly orange tone; and second, what appears to be the posterior edge of the left wing membrane coursing between the distal end of the wing and the ankle is also preserved as an orange stain. This color is not closely associated with any other part of the matrix (though some pale orange areas are present elsewhere on the slab) and the vast majority is limited to areas adjacent to bones and

9 2013 HonE Et al. solnhofen PtErosaurs at carnegie MusEuM 173 Fig. 5. CM 11431, Rhamphorhynchus muensteri in left lateral and dorsal views. Scale equals 10 cm with 1 cm divisions. Abbreviations: dv, dorsal vertebrae; hu, humerus; L, left wing; mc, metacarpus; mn, manus; R, right wing and hind limb; r, radius; sc, scapula; ti, tibia; ul, ulna. other places where soft tissues would be expected. The tail vane is preserved as a broad, diamond- or kitelike shape, as seen in other large specimens of Rhamphorhynchus (Bennett 1995), and measures 58 mm long with a maximum height of 43 mm. The edges of the vane are clear, though no structural fibers or similar anatomical features can be seen in the tail as in some other specimens (Wellnhofer 1975). Comments. The associated Carnegie Museum label identifies this specimen as Rhamphorhynchus munsteri [sic]. We follow this identification (and that of Wellnhofer 1975) as R. muensteri as a result of the presence of the following diagnostic morphologies: ten teeth per side in the upper jaws, where the anterior teeth are anteriorly directed; orbit considerably larger than antorbital fenestra; and first wing phalanx longest in series, with its length subequal to that of the skull (Bennett 1995). The museum label also identifies the specimen as having been obtained as part of the Bayet collection in The most significant parts of this specimen are the preserved soft tissues, and in particular, the traces that are considered probable remnants of the left wing membrane. The trailing edge of the wing is nearly straight. Although its proximal and distal ends are rather indistinct in the main slab, these areas are clear on the counterslab. Following the line of tissue as it is preserved, the distal part would appear to connect with the tip of the wing, as would be expected, whereas the proximal part would be in line to connect with the ankle as seen in a number of other pterosaurs, including other Rhamphorhynchus specimens (Frey et al. 2003; Elgin et al. 2011). As preserved, the chord of the wing is relatively broad along its entire length, and the orange stain is likely not an exact representation of the extent of the wing, given, for example, the patch around the pes on the counterslab. Nevertheless, at least part of the inferred posterior edge is too straight and too well positioned (compared to what is known of Rhamphorhynchus wing anatomy [e.g., Marsh 1882; Zittel 1882; Padian and Rayner 1993]) to be a coincidence, and therefore it must at least partly represent the genuine contour of the wing. CM Taxonomic identity. Rhamphorhynchus muensteri. Previous mentions in the literature. Koh (1937); Wellnhofer (1975 [his example 56 ]); Chatterjee and Templin (2004). Description. CM is an articulated skeleton of a

10 174 annals of carnegie MusEuM vol. 82 Fig. 6 CM 11431, Rhamphorhynchus muensteri prior to acid preparation and mounted in a wooden frame. The pictured ruler is 15 cm. large (wingspan approximately 950 mm) non-pterodactyloid pterosaur that was originally exposed primarily in left lateral and dorsal views (see Wellnhofer 1975: pl. 14, figs. 1 2). It is superbly preserved although somewhat compressed. In 1992, the specimen was loaned to Sankar Chatterjee (Texas Tech University, Lubbock, Texas, U.S.A.), who had it acid-prepared nearly free of matrix; it is therefore now visible in three dimensions, although extraordinarily fragile (Figs. 5 6). Some sediment remains, however, especially on the left side of the skull and the dorsal part of much of the remainder of the skeleton; this is colored green, presumably as a result of staining from glue or other preservatives employed during the preparation process. The specimen is nearly complete, lacking only the tail, the distal ends of the wings, much of the pelvis, and the left hind limb and the distal end of the right. The bones are medium brown in color. The fusion of the elements comprising the skull and mandible, the neurocentral sutures, the extensor tendon process to the proximal end of the wing metacarpal, and the sacrum to the ilium all suggest that the animal was an adult at death. The skull is complete and articulated with the mandible. Most of the left sclerotic ring is preserved, although damaged, and lies within the orbit. Nearly all of the teeth are apparently present, but two from the left maxilla are displaced from their sockets and directed dorsally. Little can be said of the cervical series; however, it appears to be both complete and articulated. The posterior cervical and anterior dorsal vertebrae are concealed on one side of the specimen by matrix and on the other by a sheet of bone that represents the sternum and probably the right coracoid. There is a very sharp bend in the presacral vertebral column, such that the anterior cervicals and the dorsals are antiparallel to one another, suggesting that two or more vertebrae are dislocated somewhere along the cervicodorsal transition. The dorsal series also appears to be complete and articulated but only two possible ribs or gastralia can be seen. At least two, and probably three, sacral vertebrae are in sequence with the dorsal series and can be identified by their large lateral processes that articulate with the partial right ilium. Both scapulae and coracoids are present and articulated. The forelimbs are also well preserved and in nearly natural articulation. They are almost complete, though both wing fingers lack their distal elements. The humeri are well preserved, although the right has suffered some damage to the deltopectoral crest. Both pairs of radii and ulnae are well preserved and articulated. The two carpal blocks appear to be fused. The pteroids have apparently been lost, although the most proximal part of the right may be present on the corresponding carpus. The two wing metacarpals are well preserved and articulated, although the left metacarpals I III have rotated out of position. Except for the wing fingers, the phalanges of digits I III appear complete and articulated in both manus. The right wing finger has phalanges 1, 2, and the most proximal part of 3 preserved. Phalanx 2 has split along its long axis and its two parts are somewhat separated. On the left side, only the first wing phalanx is completely preserved, though most of phalanx 2 is also present. The latter is somewhat crushed and appears to be close to splitting in a fashion comparable to its counterpart on the right side. Of the pelvis, only the anterior ramus of the right ilium is preserved. Part of the right hind limb, consisting of the complete femur and tibia, is articulated with the ilium. No other hind limb elements are preserved. Comments. CM was still on loan to S. Chatterjee when DWEH visited Carnegie Museum of Natural History in November 2011 and was therefore not seen by the first author during the writing of this manuscript. As such, our description is based partly on photographs. The specimen has since been returned to the museum and was directly examined by MCL and MBH in February The museum s specimen database identifies CM as R. muensteri, a taxonomic assignment also given by Wellnhofer (1975). Here we follow this assignment owing to the presence of the following characters: edentulous jaw tips; large and anteriorly directed anterior teeth and smaller, subvertical posterior teeth; antorbital fenestra larger than naris but both considerably smaller than orbit; and humerus longer than femur (Bennett 1995). The fragility of the specimen makes it difficult to measure; however, we estimate its wingspan at approximately 950 mm based on the material and comparison to similarly sized specimens described by Wellnhofer (1975). This is somewhat higher than the 860 mm wingspan reported for this specimen by Chatterjee and Templin (2004). For a Solnhofen pterosaur, CM is unusual in its preparation, and perhaps unique in the manner in which the skeleton has been almost entirely freed of matrix. Although traces of rock remain to help maintain the structural integrity of the specimen, it is accessible in both major dimensions, allowing individual bones to be viewed from multiple angles.

11 2013 HonE Et al. solnhofen PtErosaurs at carnegie MusEuM 175 Fig. 7. CM 11432, Rhamphorhynchus muensteri. Scale equals 10 cm with 1 cm divisions. Abbreviations: cdv, caudal vertebrae; cv, cervical vertebrae; fe, femur; hu, humerus; L, left hind limb; ps, pes; R, right hind limb; r, radius; sk, skull; st, sternum; ti, tibia; ul, ulna; wmc, wing metacarpal; wpx, wing phalanx. CM Taxonomic identity. Rhamphorhynchus muensteri. Previous mentions in the literature. Koh (1937); Wellnhofer (1975 [his example 54 ]). Description. A large (wingspan approximately 890 mm), nearly complete and articulated specimen exposed in multiple views (Fig. 7). The large body size, fusion of the scapula and coracoid, and ossification of the sternum suggests that this animal had reached adult size, or nearly so, at the time of death. The matrix is pale yellow with the bones very dark brown. A halo of orange stains is preserved around most skeletal elements. The slab is mounted in a plaster block that sits in a wooden frame and that has undergone significant repair, with large breaks having been filled with dark-colored plaster. Plaster possibly covers some parts of the skeleton (e.g., the base of the tail). Overall the bones are in poor condition and in many cases badly damaged. The skull is well preserved and exposed in ventral view, although the ventral end of the braincase is broken off and part of the posterior palate is missing. The teeth are mostly missing or barely exposed, but dendrites on the matrix extend from the alveoli, giving the impression that teeth are

12 176 annals of carnegie MusEuM vol. 82 present. The mandible is not preserved. The cervical column is articulated with the skull but not with the dorsal vertebrae. It is seen in right lateral view and appears to be in reasonably good condition. However, the anterior parts are partially covered by matrix and could therefore be exposed with further preparation. Cervical ribs are present and articulated with the vertebrae. The dorsal series is complete and articulated but also not fully prepared. Several anterior dorsal ribs are articulated with their respective vertebrae, but others are not and lie with the gastralia in the vicinity of the posterior dorsal region. These ribs and gastralia are mostly preserved as impressions in the matrix. The sacrum is indistinct and largely present only as an impression, though the orientations of the ilia and the fact that they overlie the proximal ends of the femora suggest that it is in dorsal view. The base of the tail is missing or covered by repair to the slab. The position of the pieces of matrix that comprise the specimen and the orientation of the posterior half of the tail suggest that it is in a natural position relative to the remainder of the skeleton and that therefore its apparent length is genuine. The sternum is present and lies below the dorsal vertebrae and ribs. Unfortunately, however, it is in poor condition and little detail can be made out. The?right scapulocoracoid is present in?anterior view. Both forelimbs are preserved, although it is not clear which is the left and which is the right. Both have suffered extensive damage and the cortices of their bones are often broken or missing; furthermore, some parts are represented only as impressions in the matrix. The proximal end of one forelimb lies under the body but then overlaps the other. In the former, the head of the humerus is missing or buried in sediment. The radius and ulna are apposed but have been slightly disarticulated and lie somewhat apart on the slab. The wing metacarpal is nearly in articulation with the distal ends of the radius and ulna. Two fragments of bone and impressions likely represent parts of the phalanges of manual digits I III. Another blocky element that lies near the distal end of the wing metacarpal likely represents part of the carpus. The wing phalanges of this limb are missing. The other wing is nearly complete and mostly articulated. The humerus is in fair condition although its proximal end has suffered some damage. The radius and ulna are articulated and also damaged proximally. These elements have not separated as in the other wing, but they do have matrix in the space between them, suggesting they are not as tightly appressed as is usual for pterosaurs. A block of bone at the distal end of the radius and ulna represents the proximal carpus. The disarticulated distal carpal lies close to the distal end of the wing metacarpal. The wing metacarpal and distal wing are somewhat displaced, such that the proximal end of the metacarpal lies close to that of the ulna. The proximal parts of digits I III are adjacent to the wing metacarpal and the distal parts including the unguals are present as impressions in the matrix. All four wing phalanges are at least partially present, though damaged. The distal parts of phalanges 1 and 2 and nearly all of phalanx 3 are missing, represented only as impressions. The proximal part of phalanx 4 is present as an impression but the remainder is missing as the slab is broken. Both anterior rami of the ilia are present and in a natural position with respect to the vertebral column, suggesting that they may be fused to the sacrum. However, no other parts of the pelvis can be seen. Both hind limbs are present and in articulation, though damaged. The right femur and tibia are in poor condition and mostly preserved only as impressions in the matrix. The right pes is disarticulated and indistinct, being covered in part by the dorsal vertebrae and ribs, and only a few elements can be identified. The left hind limb is more complete and in much better condition than the right. Both the femur and tibia are present and lie in line with one another. The tarsus is preserved but indistinct. Three metatarsals are present and articulated, but it cannot be determined if they correspond to metatarsals I III or II IV, and some have impressions of phalanges distal to them. Comments. The label accompanying the specimen identifies it as simply Rhamphorhynchus while the Carnegie Museum s database lists it as R. carnegiei Koh, This latter name was considered by Wellnhofer (1978) to be a junior synonym of R. muensteri. Here we follow the latter identification because the anterior teeth are anteriorly directed, the humerus is longer than the femur, and the first wing phalanx is the longest in the series (Bennett 1995). The museum label notes that the specimen was part of the Bayet collection. No date is listed for its acquisition, but given that other Bayet specimens with both higher and lower Carnegie Museum catalogue numbers were acquired in 1903, this skeleton presumably was as well. The separation of the radius and ulna in one forelimb is unusual, especially given the otherwise reasonably wellarticulated nature of the specimen. Although the radius and ulna of pterosaurs did sometimes separate during decomposition, this seems to have been an extremely rare occurrence (DWEH pers. obs.). This suggests that, in life, the two bones were typically very well bound together by soft tissues, and that the proximal and/or distal ligaments that connected these elements might have been especially strong, perhaps even supplemented by interstitial tissue along the lengths of the bones. This tight coupling of the antebrachial bones may have been an adaptation to resisting flight forces. The center of lift on the pterosaur wing would have acted posterior to the radius and ulna (near one-quarter the length of the chord), creating a torque that would have tended to rotate the ulna relative to the radius if these bones had not been tightly bound together. CM Taxonomic identity. Rhamphorhynchus muensteri. Previous mentions in the literature. Wellnhofer (1975 [his example 21 ]); Padian et al. (2004).

13 2013 HonE Et al. solnhofen PtErosaurs at carnegie MusEuM 177 Fig. 8. CM 11433, Rhamphorhynchus muensteri in right dorsolateral view. Scale equals 10 cm with 1 cm divisions. Abbreviations: cdv, caudal vertebrae; cv, cervical vertebrae; hu, humerus; L, left wing; ps, pes; R, right wing; r, radius; sk, skull; st, sternum; ul, ulna; wmc, wing metacarpal; wpx, wing phalanx. Description. A small (wingspan approximately 390 mm), nearly complete and articulated specimen of a nonpterodactyloid pterosaur exposed primarily in right dorsolateral view (Fig. 8). The skeleton is poorly preserved with little bone present, mostly consisting of impressions in the matrix. Based on its small size, proportionally large skull, and apparently unfused pelvis, this would appear to represent an immature individual. It is preserved on a small, thin slab of yellowish-brown matrix. Glue or a preservative has been applied to much of the torso and proximal wings, and as a result these areas are darker than the rest of the specimen. This substance is highly reflective and as such it hinders examination of the specimen. Numerous calcite crystals are present in the matrix in association with radiating patterns that may well represent organic remains (given their common association with cartilage; Bennett 2007a), but not of the pterosaur. The skull is seen in dorsolateral view and crushed at an oblique angle. As such, parts of the orbits are visible, the right more so than the left, but other details of the skull are difficult to discern. Fragments or impressions in the matrix show that at least 11 teeth are present in the rostrum. The right sclerotic ring appears to be at least partially preserved. The cervical series is complete and articulated; however, it is difficult to distinguish between individual elements owing to the poor preservation of the bones and the buildup of calcite upon them. Even so, the cervical ribs appear to be present and in natural articulation with the vertebrae. The dorsal series is similarly preserved, and although it is complete and articulated, few details can be made out. The dorsal and sternal ribs are present and in near natural articulation, though some gastralia have been dislocated and lie next to the pedes. The pelvic elements appear to be missing, which suggests that they were not fused to each other, or to the sacrum. The sacrum is present and links the dorsal and caudal vertebrae, but this part of the specimen is in very poor condition. The tail is the best-preserved part of the axial series and nearly every individual vertebra therein can be identified, with only the very anterior part poorly preserved and the tip missing and left as an impression. At least 31 centra can be identified, and, given the distal tapering of the tail, the terminal caudal is likely present. An impression of the sternum is present anterior to the dorsal ribs. This bears a striking similarity to sterna described for small specimens of Rhamphorhynchus (cf. longicaudus [Münster, 1839]) (Lü et al. 2011), with a long, slender cristospine and only a small sternal plate. Both pectoral girdles and forelimbs are nearly complete and in close to natural articulation. The left scapula is partially preserved, but its blade is only an impression, while the left coracoid is an indistinct impression alongside the dorsal series, with bone preserved around the margins. The

14 178 annals of carnegie MusEuM vol. 82 Fig. 9 CM 11434, Rhamphorhynchus muensteri. Skull in following views: A, left lateral; B, right lateral; C, dorsal; D, ventral. Scale bar is 20 mm with one mm divisions. left humerus is poorly preserved and there is extensive calcite buildup along its shaft. The articulated left radius and ulna are incomplete, with only their most proxima parts preserved and their more distal sections being impressions in the matrix that overlie the skull roof. Two wing phalanges (?2 and?3) lie close to the skull, and an isolated wing

15 2013 HonE Et al. solnhofen PtErosaurs at carnegie MusEuM 179 phalanx 4 is close to the sternum. The left metacarpals and other manual phalanges cannot be seen. The right wing, by contrast, is articulated and complete, if mostly preserved as elements that have split along their long axes, revealing their internal surfaces. The right scapula and coracoid are present but almost completely concealed behind other elements. The right humerus lies alongside the dorsal edge of the dorsal vertebral column. The articulated right radius and ulna are perhaps the best-preserved elements in the specimen, having the most amount of well-preserved bone. Although the wing metacarpal is indistinct, metacarpals I III and manual digits I III are preserved as clear impressions in the matrix. All four right wing phalanges are complete, with the fourth showing a limited degree of posteriorly-directed curvature. The pelvic elements (ilia, prepubes, pubes, and ischia) all appear to be absent. Both hind limbs are complete and articulated, with the exception of the left femur, which cannot be seen and may therefore be obscured behind the dorsal series. Most hind limb elements are preserved as high-fidelity impressions in the matrix, although some of the distal phalanges are less clear. The right femur and tibia overlie the posterior dorsal vertebrae and the left tibia is partially hidden by the axial column. Comments. The Carnegie Museum label on this specimen identifies it as Rhamphorhynchus cf. longicaudus, an identification shared by Wellnhofer (1975). Here we interpret it as R. muensteri, as it has large, procumbent anterior teeth and a first wing phalanx that is both the longest in the phalangeal series and close to the length of the skull (Bennett 1995). An old, red-bordered label on the specimen bears the number 2049, but this has been struck through and replaced with 11433, suggesting a replacement catalogue number. An identical red-bordered label appears on the reverse of the specimen, except that, on this label, 2049 has not been crossed out. An apparently even older label with a blue border is affixed to another corner of the specimen, and gives the number 649. Yet another label reads 9 and Jurassique [word illegible] Solnhofen but there is no obvious indication as to what these may refer to. Perhaps, since its text is in French, and French is spoken widely in Belgium (especially in Brussels and nearby areas), this last label may date to prior to 1903, when the specimen was still owned by the Baron de Bayet. Unsurprisingly, the Carnegie Museum label gives the source of the specimen as the Bayet collection and its acquisition date as The provenance of the specimen is given simply as Solnhofen (i.e., Carnegie Museum locality 494). Accompanying documents prepared by personnel at the Museum of the Rockies (Bozeman, Montana, U.S.A.) state that a coracoid, radius, ulna, and wing phalanx of CM were sampled for histology and subsequently repaired. However, based on the included sketch of the specimen, the element that these persons identified as a Fig. 10 CM 11434, Rhamphorhynchus muensteri in posterior view. Scale bar is 10 mm with one mm divisions. coracoid we interpret as a humerus. The results of this histological analysis were detailed by Padian et al. (2004). In that paper, specimens CM (described below) and CM are identified as Pterodactylus Cuvier, 1809, and Rhamphorhynchus, respectively. Oddly, although the main text of the paper lists the numbers of these specimens correctly, the only figure in which they are shown (Padian et al. 2004: fig. 6) misidentifies them as CM and CM 11400, respectively. CM Taxonomic identity. Rhamphorhynchus muensteri. Previous mentions in the literature. Huene (1914); Wellnhofer (1973, 1975 [his example 89 ]); Padian (1984); Witmer et al. (2003); Chatterjee and Templin (2004). Description. The specimen is an isolated skull that is preserved and acid-prepared in three dimensions (Figs. 9 10). It is generally in excellent condition, though it lacks all teeth and the tip of the rostrum. The bone is a pale brown color with yellow parts and small orange highlights, the latter especially around the edges of fenestrae. As preserved, the skull is 92.4 mm in length, which corresponds to some of the larger specimens of Rhamphorhynchus. The skulls of CM and CM are only slightly longer (see Table 1), suggesting that the wingspan of the individual represented by CM approached 1000 mm. Wellnhofer (1975: pl. 16.3) published a photograph of the palate of CM 11434, in which considerable amounts of matrix are clearly visible around the skull. In 1979, the specimen was loaned to John Ostrom and Kevin Padian of Yale University (New Haven, Connecticut, U.S.A.); subsequently, in 1992, this loan was transferred to S. Chatterjee of Texas Tech University, who had technician Zhong Zheng extensively and superbly prepare CM such that it is now essentially free of sediment (Witmer et al. 2003). Only a few cranial sutures can be discerned, with

16 180 annals of carnegie MusEuM vol. 82 Fig. 11. CM 11425, Aurorazhdarcho micronyx in right lateral view. Scale equals 10 cm with 1 cm divisions. Abbreviations: L, left wing; R, right wing. others being obliterated, suggesting that CM belonged to an adult or nearly adult animal, a conclusion that is further supported by the large size of the specimen. As a whole, the skull is somewhat dorsoventrally flattened, and the skull roof is depressed, which has at least affected the shape of the orbits and the orientation of the narial openings. A raised ridge courses along the midline of the skull roof, which nonetheless seems to have been crushed downward, especially in the area of the nares. The palate, by contrast, is in good condition and appears complete. There is some buildup of calcite in the foramen magnum and this may well continue further inside the skull. Comments. This specimen was on loan during DWEH s visit to Pittsburgh, and as such it was not examined by the first author firsthand during the writing of this manuscript. The description is therefore based on photographs provided by Lawrence Witmer (Ohio University, Athens, Ohio, U.S.A.) and personal observations by MCL and MBH. The specimen is listed in the Carnegie Museum database as R. gemmingi von Meyer, 1846, and was identified by Wellnhofer (1975) as Rhamphorhynchus sp. Here it is identified as R. muensteri as it possesses a narrow lower temporal fenestra and a large and rounded upper temporal fenestra (Bennett 1995). CM was subjected to a high-resolution computed tomographic scan by Witmer et al. (2003) as part of their investigations of pterosaur neuroanatomy. This revealed that the osseous labyrinth is largely preserved internally. Witmer et al. (2003) interpreted the morphology of this structure to indicate that, in life, the head of Rhamphorhynchus was habitually held subhorizontally relative to the axial column. CM Taxonomic identity. Aurorazhdarcho micronyx (von Meyer, 1856). Previous mentions in the literature. Wellnhofer (1970 [his example 36 ], 1973); McGinnis (1983:52 53); Bennett (2013).

17 2013 HonE Et al. solnhofen PtErosaurs at carnegie MusEuM 181 Description. A small (wingspan approximately 325 mm), articulated, and nearly complete pterodactyloid specimen visible predominantly in right lateral view and compressed into two dimensions (Fig. 11). The matrix is a pale fawn color with yellow streaks across the surface, and the bones are pale brown. Almost the entire skeleton is represented as impressions in the matrix, although these are often quite detailed. Bone fragments remain in situ in the anterior ends of the jaws, the skull roof dorsal to the orbit, some ribs, parts of the distal wings, and the midshaft of the right femur. Owing to the nature of the preservation, it is not clear what ontogenetic age the animal may have been at the time of death, since no sutures can be seen between elements and the lack of disarticulation gives no indication as to what bones may remain unfused. However, the small size of the specimen implies that it belonged to a young individual. The skull is intact and articulated with the mandible, and shows that the tips of the jaws contained numerous small teeth. These are larger at the anterior ends of the jaws and rapidly become smaller posteriorly. Traces of bone lie below the posterior end of the skull and seem to be part of the mandible as they appear to be larger than would be expected for the hyoids. As such, it appears that the posterior part of the mandible is disarticulated, though this is difficult to determine. The cervical vertebral series is complete and articulated, though the morphology of its constituent elements is not clear. These are mostly represented as deep impressions in the matrix as a result of the laterally broad nature of many pterodactyloid cervical centra. The dorsal series is also intact and articulated, and is better preserved than the cervical series, providing clear outlines of individual vertebrae. The dorsal ribs are articulated with their respective vertebrae; the gastralia are also present and remain close to their in vivo positions. The sacral vertebrae are partially concealed behind the right ilium but the dorsal ends of their neural spines are visible. A number of caudal elements are also present but appear to be at least partially disarticulated. An impression of the right scapula overlies the anterior dorsal vertebrae. Below this, a deep cavity likely represents the part of the coracoid that contributes to the glenoid. Both forelimbs are complete and articulated. Diminutive structures including the thin metacarpals and phalanges of digits I III and the pteroid are all visible, as are some details of the left carpus. The right ilium is complete but it is not clear if other pelvic elements are also preserved. Indistinct marks ventral to the ilium may represent other parts of the pelvis but this is uncertain. As with the forelimbs, both hind limbs are complete and articulated. Although they are preserved primarily as impressions, details of delicate elements including the fibulae and tarsals are evident. Comments. No original taxonomic identity was given on labels with this fossil, though the museum s electronic database identifies it as Pterodactylus elegans Wagner, Wellnhofer (1970) identified it as a specimen of Pterodactylus micronyx von Meyer, Here it is interpreted as a juvenile of Aurorazhdarcho micronyx following Bennett (2013). Bennett (1996) considered specimens of P. micronyx to be juveniles of Gnathosaurus subulatus von Meyer, 1833, but more recently has assigned the species micronyx to the genus Aurorazhdarcho Frey et al., 2011 (Bennett 2013). We follow that identification here based on the long and slender rostrum and the concave dorsal margin of the skull, although the nasoantorbital fenestra is slightly longer than is typical for A. micronyx (approximately 33% of skull length as opposed to 20 25%; Bennett 2013). Wellnhofer (1970:45) stated that, relative to other P. micronyx skeletons, the humerus of CM was unusually short, but its proportional length differs little from that of other, similarly-sized specimens of this taxon (Wellnhofer 1970: Übersichtstabelle). On the reverse side of the specimen is a red-bordered label bearing the number 2047 ; this has been struck through with written in its place. As with several other specimens (e.g., CM 11428; see above), the number 2047 may relate to the cataloguing of the material prior to its formal accession into the CM collections. An old handwritten label is also on the reverse side of the specimen but is unreadable. The slab on which the specimen is preserved has two holes drilled in it (now repaired with plaster), presumably for mounting the specimen on a wall. This is similar to (but not exactly the same as) the way in which some other Solnhofen specimens were drilled through to secure them to frames by early fossil dealers (Hone 2010). CM Taxonomic identity. Aurorazhdarcho micronyx. Previous mentions in the literature. Wellnhofer (1970 [his example 44 ]); McGinnis (1983:52 53); Bennett (2013). Description. A small (wingspan approximately 510 mm) pterodactyloid pterosaur that is complete, articulated, and exposed primarily in left lateral view (Fig. 12). Although all bones are apparently present, many are in poor condition or preserved in only two dimensions. Calcite crystals have formed at some of the joints, as is common in Solnhofen pterosaurs (Bennett 2007a). Furthermore, preparation is incomplete and some elements are only partially exposed. The ontogenetic age of the specimen is difficult to determine, but parts of the skull and pelvis appear very slightly disarticulated, suggesting that these areas had not fused and thus that the animal was not an adult at death. The small size of the material also implies juvenile or subadult status. The matrix is a pale gray color with the bones being medium and occasionally dark brown. The skull is nearly complete and undistorted, and the dorsal edge of the rostrum is noticeably concave. The skull

18 182 annals of carnegie MusEuM vol. 82 Fig. 12. CM 11426, Aurorazhdarcho micronyx in left lateral view. Scale equals 10 cm with 1 cm divisions. Abbreviations: L, left wing; R, right wing. roof along the rostrum is slightly damaged, and it appears that splints of bone have been lost. The posterior end of the skull has extensive calcite growth. The mandible is complete and articulated with the skull. Both the upper and lower jaws have numerous small teeth in their tips. A partial sclerotic ring is preserved in the orbit; the scleral ossicles are darker in color than the other bones, and the ring is narrow in diameter. The hyoids are present as a pair of thin bones positioned ventral to the mandible and in the same anteroposterior plane as the orbit. The cervical vertebral series is complete, articulated, and exhibits low neural spines. The dorsal series is mostly absent, but it is not clear if the elements are missing or simply obscured by overlying matrix and calcite. The posterior dorsal vertebrae are preserved but in poor condition. The dorsal ribs are articulated with their corresponding vertebrae, or, more accurately, where these vertebrae are likely situated, and the gastralia also appear to be close to their natural positions. The sacrum is apparently in situ but hidden medial to the left ilium. The caudal series is articulated and probably complete (the posteriormost preserved caudal is indistinct but very small, and most likely represents the terminal caudal). The caudal series is strongly curved, such that its tip points dorsally. A mass of bone ventral to the anterior end of the dorsal series appears to be at least one scapula and coracoid and part of the sternum. However, the poor preservation of this area and incomplete preparation make this difficult to determine. The left humerus is incomplete, and its proximal part and midshaft are preserved only as impressions. The left radius and ulna are complete but in poor condition. A long, thin element lies adjacent to these bones. Near its midpoint, this bone is oddly kinked around a calcite crystal. It may be a pteroid, but it appears to be too long to be this element, and so instead it may be a dislocated gastralium or even a displaced metacarpal I, II, or III. Metacarpal IV appears unusually thin, but it has rotated out of position, as can be seen by the appearance of the lateral face of the distal articular end. As such, metacarpals I III are not visible (unless the elongate, kinked element mentioned above corresponds to one of these bones). Manual digits I III are poorly preserved but complete and articulated. The wing finger is also complete and articulated, although phalanx 2 has suffered some damage to its proximal part. Only part of the right humerus is visible, and only the distal ends of the radius and ulna are exposed (with the rest of these bones being covered by other elements or damaged). All four metacarpals are present and articulated, and digits I III are preserved as in the left arm. The wing finger is not evident, but as with other elements from the right side, it is probably simply buried in matrix given the otherwise complete and articulated nature of the specimen (and especially the presence of the tail). One (the left?) prepubis is preserved in lateral view. Part of the left ilium is also preserved, though poorly, and it is partly concealed by the left femur. The left hind limb is complete, articulated, and well preserved, largely

19 2013 HonE Et al. solnhofen PtErosaurs at carnegie MusEuM 183 in three dimensions. Notably, the proximal part of the left fibula is present as a long, thin splint of bone. A poorly preserved piece of bone below the prepubis is most likely the underlying right femur. The remainder of the right hind limb is not seen but, as with the right forelimb, it is probably buried in matrix. As noted by Wellnhofer (1970), CM preserves what appear to be impressions of soft tissues adjacent to the ventral parts of the mandible and cervical series, the dorsal part of the cervical series, and the left forelimb. Impressions associated with the forelimb are vestiges of the folded wing membrane, while those near the lower jaw and the ventral part of the neck may correspond to a throat sac, as has been reported in selected specimens of Aurorazhdarcho (Frey et al. 2011), Pterodactylus (Wellnhofer 1991; Frey and Martill 1998; Bennett 2013), and Rhamphorhynchus (Unwin 2005: fig. 5.3), among other genera. Comments. No original taxonomic identity is provided on labels associated with the specimen, though the museum s electronic database identifies it as Pterodactylus sp. Wellnhofer (1970) identified it as a specimen of Pterodactylus micronyx. Here it is interpreted as Aurorazhdarcho micronyx following Bennett (2013). As with CM 11425, the skull is very long and low with a dorsally concave rostrum and numerous, slender, closely packed anterior teeth. As is also the case for CM 11425, at some point in its history, the matrix of the specimen was drilled through to be mounted. Old labels on the reverse side are also similar to that on CM A number on a red-edged label reads 2047, which has been struck through and replaced with A second, blue-edged label reads simply 3. A large, handwritten label cannot be read as it is faded and damaged, but it appears to be written in the same hand as that of CM The degree of curvature seen in the tail is unusual, perhaps even unique for a pterodactyloid. The lack of disarticulation and the presence of all caudal elements suggest that this curvature is genuine and not the result of taphonomic distortion, although it could be linked to opisthotony. Although, as is also the case for CM and CM 11427, CM is currently on public display in Carnegie Museum of Natural History s Mesozoic fossil gallery, an unlabeled, unpainted white plaster cast of CM is stored in the museum s Vertebrate Paleontology collections. This cast is housed in a drawer with CM and a similar, unpainted plaster cast of this latter specimen (see below). CM Taxonomic identity. Aurorazhdarcho micronyx. Previous mention in the literature. None (see below). Description. A small pterodactyloid pterosaur exposed in right lateral view (Fig. 13); this is the counterplate of CM The material is primarily preserved as fine impressions in the matrix, with fragments of bone representing some parts of the maxilla, orbital region, cervical vertebrae, ribs, pectoral girdle, humerus, proximal and distal radius and ulna, and the proximal part of metacarpal IV. The matrix is a thick slab of Solnhofen limestone that has several large cracks across it but that has been adequately repaired with plaster. The matrix is fawn in color with the few fragments of bone being light brown. The skull is preserved mostly as an incomplete, shallow impression on the surface of the slab. A small fragment of bone makes up part of the posteroventral portion of the maxilla, and another fragment is situated at the anterodorsal corner of the orbit. The mandible is present and there is a suggestion of teeth at the tips of the jaws. The cervical series is preserved almost entirely as impressions, with just a few bone fragments being preserved. There are indistinct traces that likely represent part of the anterior dorsal series, as well as fragments of anterior rib shafts. There are also some bony parts and clear impressions of three posterior dorsal ribs that are positioned such that they would likely articulate with their respective vertebrae. There is no trace of the sacral or caudal series. The left forelimb is partially preserved. There are traces of the scapula and the humeral shaft is largely represented by bone. Most of the radius, ulna, and wing metacarpal remain only as impressions. Two faint impressions likely represent the first two wing phalanges. There are traces of the left hind limb, with partial but indistinct impressions of what are likely the femur, tibia, and part of the pes. Comments. A Carnegie Museum label identifies CM only as Pterodactylus, while the museum s specimen database lists it as Pterodactylus sp. Here we consider it to be a juvenile individual of Aurorazhdarcho micronyx (Bennett 2013), and more specifically, the counterplate of CM Specimens CM and CM are strikingly similar in size, shape, and posture, though as mirror images of one another. However, this is not uncommon for Solnhofen pterosaur specimens: for example, the pose of CM is very similar to that of CM 11425, and the Vienna and Munich specimens of Pterodactylus (NHMW 1975/1756 and BSPG 1937 I 18, respectively) are also remarkably similar to one another. Indeed, at some point in the past, the similarity of the former two (CM) specimens appears to have caused confusion that may have ultimately led to the dissociation of CM from CM Specimen CM has CM (note that in this and the following paragraphs we have emphasized the terminal numbers of specimens CM and CM for clarity) written at least once, and possibly as many as three times, on the slab. A note, handwritten in 1978 on the reverse of the CM specimen label by former CM fossil preparator Robert Cardillo, reads This was formerly but was not the counterpart of on display. Therefore it has a new no. 8/78 R.F.C. As Cardillo noted, there is no reason to consider CM

20 184 annals of carnegie MusEuM vol. 82 Fig. 13 CM 35002, Aurorazhdarcho micronyx in right lateral view. This is the counterplate to CM (see Figure 12). Scale equals 10 cm with 1 cm divisions as the counterpart of CM The matrices of the two specimens are different colors, the sizes of various elements and their exact positions are quite different, and CM is unusually high in relief, with a large bulge in the middle of the specimen that would not fit well with the very flat CM In contrast, a compelling case can be made that CM is the counterpart of CM The convex surface of CM fits well with the strong concavity of CM 11426; this has been tested by MCL with casts of the two specimens held in the CM collections. Furthermore, many anatomical measurements of the specimens correspond closely (if admittedly not quite perfectly, a fact that we attribute to the incompleteness and generally poor preservation of CM 35002). For example, the lengths of the skulls and ulnae differ by less than 1 mm, and the wing metacarpals by just 1.2 mm (see Supplementary Table). The preservation of the middle parts of the shafts of a radius and ulna in CM also corresponds well with the missing parts of these elements in CM Further confirmation comes from the similarity in the color of the matrices of the two specimens. Lastly, and most convincingly, digitally reflecting and overlaying photographs of the two specimens illustrates their remarkable correspondence, to the point that we are very confident that CM is the counterpart of CM Why, then, was it formerly considered the counterpart of CM 11425? Although we cannot be certain, we suspect that this error resulted from confusion over the closely similar specimen numbers of CM and CM We believe that the slab that is now CM was originally and correctly identified as the counterpart to CM 11426; regrettably, however, the number CM (instead of CM ) was written on the CM slab by mistake. In the late 1970s, Carnegie Museum staff (Cardillo and possibly others) correctly recognized that CM could not be the counterpart of CM 11425, so the former was given the separate catalogue number that it now bears. Incidentally, the fact that casts of the critical pair of specimens CM and CM share a single drawer in the CM collections suggests that, in the past, another researcher also came to the conclusion reached here (i.e., that these two specimens are part and counterpart of the same pterosaurian individual). This has, however, remained uncorrected in the museum s catalogue. CM Taxonomic identity. Pterodactyloidea sp. indet. Previous mention in the literature. Padian et al. (2004). Description. A large (wingspan approximately 1000

21 2013 HonE Et al. solnhofen PtErosaurs at carnegie MusEuM 185 Fig. 14. CM 11430, Pterodactyloidea indet. in ventral and left lateral views. Scale equals 10 cm with 1 cm divisions. Abbreviations: cp, carpus; fe, femur; hu, humerus; mn, manus; ptd, pteroid; ti, tibia; ul, ulna; wmc, wing metacarpal; wpx, wing phalanx. mm), incomplete pterodactyloid appendicular skeleton seen largely in ventral and left lateral views and consisting of an articulated but incomplete right wing and left hind limb (Fig. 14). The presence of an unfused humeral epiphysis suggests that this was not an adult animal. The majority of the material is preserved in a slab, with impressions and some additional bone in an associated counterslab. The two parts are mounted together in a painted wooden frame. Gaps between the frame and specimen and a number of large cracks on the slabs have been repaired with plaster. The frame appears to have been added relatively recently (as the wood does not look old) to help support the material and to hold the slab and counterslab together. The bones are generally in good condition and preserved in three dimensions, but the joints between them are heavily obscured by calcite crystals. The bones are a pale graybrown color on a generally yellow matrix and there are extensive dendrites on the matrix around the bones; these are more clearly visible on the counterslab. Only the broken distal part of the humerus is visible; the proximal part is preserved only as an impression in the matrix. In the counterslab, a piece of bone adjacent to the proximal end of the humerus is interpreted as an unfused epiphysis. Of the forearm bones, only the ulna is visible, and it is not clear if the radius is buried in the matrix or lost. The ulna is well preserved but its cortex is slightly damaged at midshaft, a circumstance that is due, at least in part, to destructive sampling for a recent histological analysis (Padian et al. 2004; see below). The carpus is an indistinct mass of calcite and no details can be made out. Articulated with the carpus and directed medially is the pteroid; this is unusually long (60% the length of the ulna) and thin. Metacarpals I III cannot be seen, but the wing metacarpal is present, although both its proximal and distal ends are covered in calcite and the latter is damaged. Manual digit I is complete and articulated; digit II is also present but its penultimate phalanx and ungual are slightly disarticulated from the proximal phalanx. The wing finger is incomplete, with only a small part of the proximal end of phalanx 3 being present in the slab, most of the shaft of

22 186 annals of carnegie MusEuM vol. 82 this bone preserved in the counterslab, and phalanx 4 missing (or possibly buried in matrix). The wing phalanges are in good condition but their articular ends are partly covered in calcite. The femur is well preserved and noticeably bowed anteriorly. The head is buried in matrix and cannot be seen. The tibia is articulated with the femur, but its proximal and distal ends are damaged. The counterslab appears to be a generally good representation of the slab. Parts of the carpus, wing phalanx 3, and the tibia are all in the counterslab, as are one or more pieces that represent the very distal end of the wing metacarpal and/or the most proximal part of wing phalanx 1. A large, subquadrangular element lies distal to the tibia, but is too large to be a tarsal. It is not well preserved, and could be an isolated wing carpal or a large middle cervical vertebra. Comments. The specimen is listed in the Carnegie Museum of Natural History specimen database as Rhamphorhynchus sp but it is clearly that of a pterodactyloid. An accompanying handwritten note from Alexander Kellner dated 30 Oct 95 notes the long metacarpal IV relative to the humerus and gives this as Pterodactyloidea indet. Pterodactylus? The accompanying museum label indicates that the specimen is from Carnegie Museum locality 494 (i.e., Solnhofen), and confirms that it was brought to the museum as part of the Bayet collection in Documentation accompanying the specimen shows that the ulna, wing phalanges 1 and 2, the femur, and the tibia were sampled for histological analysis by John Horner and K. Padian in March 1999, and that the removed parts were later replaced with painted casts by Ellen-Thérèse Lamm of the Museum of the Rockies. The specimen is listed as Pterodactylus on these documents and identified as such in the resulting publication (Padian et al. 2004). As noted by Kellner, the material is obviously that of a pterodactyloid and not Rhamphorhynchus, given the proportionally short humerus, long, slender pteroid, and long ulna, wing metacarpal, and femur. However, its assignment to Pterodactylus is highly questionable (see below). The putative unfused humeral epiphysis implies that this individual had not reached skeletal maturity at the time of death; indeed, the animal may be a juvenile that died at considerably less than adult size. This is supported by the histological analysis of Padian et al. (2004), who concluded that, in CM 11430, there was no evidence of the slowing or cessation of growth in the external cortex of the bones. CM is comparable in size to some of the larger pterodactyloid specimens from the Solnhofen beds. Nevertheless, its probable immaturity suggests that it attained a still larger size at adulthood, and further complicates its generic identification, given that pterosaurs exhibit ontogenetic shifts in limb proportions that can confound taxonomic work (see, for example, Bennett 1995, 1996). With only limb elements available, determining the identity of the specimen is therefore difficult. Absolute size is a poor diagnostic character within Pterodactyloidea, given, for example, the size discrepancy exhibited by adult males and females of at least some species (Bennett 2001). Nevertheless, CM is almost certainly not referable to Pterodactylus, in that it is likely immature but also nearly 50% larger than the largest known specimen of this genus (RGM St , that has a wingspan of 700 mm). This hypothesis is further supported by the relative proportions of wing phalanges 1 and 2: in CM 11430, wing phalanx 2 is slightly longer than wing phalanx 1, but in larger specimens of Pterodactylus, it is only 91% the length of wing phalanx 1 (Bennett 2013). It is conceivable that CM could pertain to Ctenochasma von Meyer, 1852, given that the proportions of its proximal forelimb elements are nearly identical to those in the largest known specimens of this genus. In CM 11430, the humerus comprises 30.5% the length of the proximal wing, whereas the ulna is 42.2% and the wing metacarpal is 27.4%. In the largest Ctenochasma specimen (BSP 1935 I 24, Elgin and Hone, unpublished data), these proportions are 30.6%, 41.7%, and 27.8%, respectively. However, even though, as above, CM is probably not an adult, it is already much larger than the largest known Ctenochasma, and other limb proportions are also inconsistent with this taxonomic identity. In the largest Ctenochasma skeleton with both the humerus and femur preserved (the Schöpfl specimen ; Bennett 2007b), the humerus (47 mm) is slightly longer than the femur (44 mm) whereas in CM the femur (71 mm) is longer than the humerus (68 mm; Table 1). It therefore seems unlikely that the CM specimen belongs to Ctenochasma. No definitive comparisons can be made to Gnathosaurus von Meyer, 1833, given the lack of postcranial material for that genus. However, the longest known Gnathosaurus skulls are similar in length to those of the largest Ctenochasma individuals (Wellnhofer 1970), which would seem to rule out this generic identification as well. CM is similar in size to the largest known Aurorazhdarcho skeleton (NMB Sh 110), but the proportions of these specimens are very different, suggesting that they do not belong to the same taxon. For example, Aurorazhdarcho has a wing metacarpal that is longer than the ulna, but in CM 11430, the wing metacarpal is only two-thirds the length of the ulna. Specimens of Ardeadactylus (Bennett, 2013) are also comparable in size to CM (Bennett 2013), but as is the case for Aurorazhdarcho, the wing metacarpal is considerably longer than the other forelimb elements in this genus. Two species of Germanodactylus Young, 1964, are currently recognized in the Solnhofen, though they may not be congeneric and consequently are considered separately here. Both species are comparable in size to CM 11430, with the only known specimen of Germanodactylus cristatus (Wiman, 1925) (BSP 1892 IV 1) having a humeral length of 56 mm, and the largest G. rhamphastinus (Wagner, 1851) (BSP AS I 745) of 60 mm. The proportions of the proximal forelimb are also similar between the three forms (differing by less than 4% for each value of the humerus, ulna, and

23 2013 HonE Et al. solnhofen PtErosaurs at carnegie MusEuM 187 wing metacarpal). Again, however, CM also differs in some areas: for example, its femur is slightly longer than the humerus; while similar to the condition in G. cristatus, in G. rhamphastinus the humerus is much longer (data from Wellnhofer 1970). Similarly, in G. cristatus, the first wing phalanx is slightly longer than the second, but the reverse is true in CM Therefore, although both species of Germanodactylus are plausible candidates for the identity of CM 11430, neither is especially convincing. With a humeral length of 68 mm and a wingspan of approximately 1 m, CM is roughly half the size of the largest pterodactyloid specimens in the Solnhofen (Elgin and Hone, unpublished data), that have humeral lengths in excess of 115 mm and a wingspan of over 2 m. The wing proportions of CM are also different from those of these large specimens; for example, in the Carnegie specimen, the wing metacarpal is much shorter than the humerus, rather than subequal in length as in the latter form. However, given the size and apparent immaturity of CM 11430, it is not impossible that this specimen might represent a juvenile individual of the large and as-yet unidentified form. Despite the highly incomplete nature of CM 11430, a number of systematically informative characters can be observed and compared to the conditions in other pterodactyloids (though these should be treated with caution given the probable immaturity of the individual). The length of the humerus relative to that of the wing metacarpal (Kellner 2003, character 54) is greater than that seen in Nyctosauridae, and the ratio of the length of wing phalanges 1 and 2 (Kellner 2003, character 68) rules out Azhdarchoidea as well. The lack of characters of these derived pterodactyloid clades suggests that CM represents a relatively basal form, but beyond this little more can be said. At least for the moment, the specimen must be assigned to Pterodactyloidea indet. It is intriguing, however, in that its placement remains uncertain despite renewed attention to Solnhofen pterodactyloid taxonomy (e.g., Bennett 2013). CM 55 Casts Specimen number of original. GPIB Taxonomic identity. Scaphognathus crassirostris (Goldfuss, 1831). Comments. This is a low-fidelity cast of the holotype of the rare non-pterodactyloid pterosaur Scaphognathus Wagner, It is painted to have orange bones and yellow-gray matrix that is similar to, but somewhat brighter than, the original fossil. An attached museum label on the reverse reads Pterodactylus crassirostris, which was the original name given to this specimen (Goldfuss 1831) before it was removed to its own genus (Wagner 1861). The label also notes that the cast was obtained from Ward s Natural Science Establishment (Rochester, New York, U.S.A.). An additional label in the box containing the cast reads Ichthyosaurus quadricissus, Ob. Lias Holzmaden. This appears to be an older label, probably German in origin, that relates to another specimen (presumably one of the Carnegie Museum s several fossils of the ichthyosaur Stenopterygius quadricissus (Quenstedt, 1856), from Holzmaden, Germany, that were obtained as part of the Bayet collection). CM 54 Specimen number of original. YPM Taxonomic identity. Rhamphorhynchus muensteri. Comments. This is a cast of the Yale specimen of Rhamphorhynchus, a specimen that preserves a complete tail vane and partial wing membranes (Marsh 1882). The cast is of poor quality and is painted in colors that differ greatly from those of the original specimen, with dark brown bones, medium brown wing membranes and tail vane, and yellow-gray matrix. The original specimen lacks the distal wings and these are molded into the surrounding mount to give the impression that the skeleton is complete. In this cast, however, no distinction is made between what is represented in the original fossil versus what parts are restored. The number 1286 is written on the reverse, but it is not clear what this might relate to. The accompanying label identifies the taxon as Rhamphorhynchus phylurus [sic] Marsh, 1882, another name that has fallen from use and presumably is linked to the antiquity of the cast as indicated by its very low number in the Carnegie Museum catalogue. CM 4027 Specimen number of original. BSPG 1883 XVI 1. Taxonomic identity. Pterodactylus antiquus (von Sömmerring, 1812). Comments. A low quality cast, painted much darker than the original fossil and depicted with orange-black bones, an orange halo around the specimen, and gray matrix. A label on the front gives the number 3614 but this is not the current specimen number, and it may therefore refer to an old catalogue number. A red-bordered label on the reverse has the current specimen number (4027) written twice. CM Specimen number of original. BSPG 1937 I 18. Taxonomic identity. Pterodactylus antiquus. Comments. This is an excellent, high-fidelity plaster cast of the beautifully-preserved and complete Pterodactylus skeleton BSPG 1937 I 18 (Broili 1938; Wellnhofer 1970 [his example 21 ]). The plaster is white but has been

24 188 annals of carnegie MusEuM vol. 82 point, as a repaired diagonal crack courses through it. On the reverse of the cast, this crack is covered and reinforced by a fabric bandage impregnated with epoxy and/or glue. Also on the reverse is the handwritten text Pterodactylus scolopaciceps H. v. MEYER; Ob. Malm, Eichstätt; Orig. zu BROILI 1938; Bayer. Staatsslg. München No I 18; CM The species name P. scolopaciceps von Meyer, 1850 has not been in use for decades (Wellnhofer 1970). The most recent works that explicitly mention BSPG 1937 I 18 Prondvai and Ősi (2011) and Bennett (2013) refer the specimen to P. kochi (Wagner, 1837) and P. antiquus, respectively. Here we follow Bennett (2013) in considering P. kochi a junior synonym of P. antiquus, and consequently regard BSPG 1937 I 18 (and, therefore, CM 19646) as a specimen of the latter. CM Specimen number of original. BSPG AS I 739. Taxonomic identity. Pterodactylus antiquus. Comments. This is a cast of the holotypic specimen of P. antiquus, the first pterosaur specimen ever described (Wellnhofer 2008). The cast is of exceptional quality and very high fidelity, and is colored in a manner that closely reflects the original specimen (DWEH pers obs). Sculpted Skeletons Another uncommon aspect of Carnegie Museum of Natural History s Solnhofen pterosaur collection is the inclusion therein of three fully reconstructed, three-dimensional, articulated skeletal models representing two species. As one of these sculptures was made by a renowned expert in pterosaur anatomy, and the other two by an acclaimed natural history artist under the supervision of professional paleontologists, their degree of accuracy is high. They are briefly mentioned here given the global rarity of such sculptures. CM Fig. 15. Notable aspects of selected Solnhofen pterosaur specimens curated at Carnegie Museum of Natural History. A, CM 11427, Rhamphorhynchus muensteri, right thoracic region showing probable fish remains in gut. The black arrow points to a putative fish tooth (see also Supplementary Figure); B, CM 11431, Rhamphorhynchus muensteri, showing three-dimensionally preserved and prepared pectoral and proximal wing elements; C, CM 11426, Aurorazhdarcho micronyx, skull and cervical series showing soft tissues and diminutive osteological structures such as sclerotic ring and hyoid elements. The black arrow points to the ventral margin of the striated throat pouch. Scale for each equals 2 cm. Abbreviations: co, coracoid; hu, humerus; r, radius; sc, scapula; ti, tibia; ul, ulna. coated in a yellow wash, presumably to more closely resemble the color of the matrix in the original specimen. Unfortunately, the cast was clearly broken in two at some Species represented. Rhamphorhynchus muensteri. Comments. This sculpture was made by German pterosaur specialist P. Wellnhofer assisted by Carnegie Museum taxidermists Christine and Otto Epping during Wellnhofer s six-month residency at the museum in 1972 (Wellnhofer 1973). Based on the R. muensteri skeleton CM 11427, it is mounted in a flying position and has a wingspan of approximately 900 mm. It consists of 50 individual pieces that were originally constructed from wax, wire, lead, plastic, and wood, and subsequently cast in epoxy. A photo of the sculpture was published by Wellnhofer (1973). According to Wellnhofer (1973), this model was the first of its kind ever made.

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