J. Paleont., 80(2), 2006, pp. 400–406 Copyright q 2006, The Paleontological Society 0022-3360/06/0080-400$03.00
NEW INFORMATION REGARDING THE HOLOTYPE OF SPINOSAURUS AEGYPTIACUS STROMER, 1915 JOSHUA B. SMITH,1 MATTHEW C. LAMANNA,2* HELMUT MAYR,3
AND
KENNETH J. LACOVARA4
Department of Earth and Planetary Sciences, Washington University, 1 Brookings Drive, Campus Box 1169, St. Louis, Missouri 63130-4899, USA, ,smithjb@wustl.edu., 2Department of Earth and Environmental Science, University of Pennsylvania, 240 South 33rd Street, Philadelphia 19104-6316, USA, 3Pala¨ontologisches Museum, Bayerische Staatssammlung fu¨r Pala¨ontologie und Geologie, Richard-Wagner-Strasse 10/II, D-80333 Mu¨nchen, Germany, and 4Department of Biosciences and Biotechnology, Drexel University, 32nd and Chestnut Streets, Philadelphia, Pennsylvania 19104, USA 1
INTRODUCTION
autumn of 1912, the fossil collector Richard Markgraf, with financial support and direction from Bavarian paleontologist Ernst Freiherr Stromer von Reichenbach and the Bavarian Academy of Sciences, discovered the partial skeleton of a bizarre predatory dinosaur in Upper Cretaceous (early Cenomanian, ;97 Ma, see Ismail et al., 1989; Barakat et al., 1993; El Beialy, 1994, 1995; Nabil and Hussein, 1994; Ismail and Soliman, 2001; Ibrahim, 2002; Gradstein et al., 2005) rocks of the Bahariya Formation exposed in the Bahariya Oasis of western Egypt (Fig. 1, see also Sereno et al., 1998; Nothdurft et al., 2002). This gigantic theropod, Spinosaurus aegyptiacus Stromer, 1915, possessed highly derived cranial and vertebral features sufficiently distinct for it to be designated as the nominal genus of the clade Spinosauridae (Stromer, 1915, 1936). Spinosaurids, currently definitively known only from Europe, South America, and Africa, are important because of the scarcity of Cretaceous Gondwanan tetrapod fossils (see Krause et al., 1999, 2003; Carrano et al., 2002; Lamanna et al., 2002). Moreover, fossils of Spinosaurus Stromer, 1915 and other spinosaurids are significant because of controversy surrounding the postulated paleoecology of these taxa (see discussions in Charig and Milner, 1997; Sereno et al., 1998; Sues et al., 2002). Questions related to spinosaurid paleoecology are particularly important in the Bahariya Formation, where Spinosaurus appears to have shared its habitat (see Stromer, 1936; Smith et al., 2001) with at least two other theropods in the size range of Tyrannosaurus Osborn, 1905 (Bahariasaurus Stromer, 1934 and Carcharodontosaurus Stromer, 1931). Unfortunately, the holotype and only known indisputable specimen of S. aegyptiacus (BSP 1912 VIII 19) was lost during the night of 24/25 April 1944 in a British bombing raid of Munich (Nothdurft et al., 2002). The attack severely damaged the building (dating to ca. 1583, No¨hbauer, 1987) that housed the Pala¨ontologische Staatssammlung Mu¨nchen and destroyed most of Stromer’s Bahariya collection (see Appendix 1). Since 1944, new spinosaurid taxa have been described (see Charig and Milner, 1986; Martill et al., 1996; Sereno et al., 1998; Sues et al., 2002) and additional material has been referred to Spinosaurus (e.g., Buffetaut, 1989, 1992; Russell, 1996; Taquet and Russell, 1998; Benton et al., 2000; Buffetaut and Ouaja, 2002), some of it questionably. However, definitive S. aegyptiacus material, or information regarding the original specimen of Spinosaurus, has not been forthcoming. We report here on two photographs of the holotype of Spinosaurus aegyptiacus as it was reposited in the Pala¨ontologische Staatssammlung Mu¨nchen prior to 1944, which we ‘‘rediscovered’’ in the archives of the Pala¨ontologisches Museum in June 2000, after they were donated to the museum by Ernst Stromer’s son, Wolfgang Stromer, in 1995 (Figs. 2, 3). These are, to our
I
N THE
* Current address: Section of Vertebrate Paleontology, Carnegie Museum of Natural History, 4400 Forbes Avenue, Pittsburgh, Pennsylvania 152134080.
knowledge, the only surviving photographs of this, the one irrefutable specimen of S. aegyptiacus, which, prior to the initial print releases, which we authorized, of the photograph in Figure 3 (Prendergast, 2001; Glut and Chiappe, 2003), has been represented only by Stromer’s (1915, 1936) drawings (it has recently been suggested that BSP 1912 VIII 19 may represent a chimera of more than one dinosaurian taxon [Rauhut, 2003], but examining that idea is beyond the scope of this paper). Aside from their historical significance, these images are important in that they permit a direct comparison of several of Stromer’s (1915) illustrations with actual photographs of BSP 1912 VIII 19, thereby providing new insight into the skeleton. This is of note because Stromer’s (1915) text and illustrations have been used as sources of character data for Spinosaurus and the Spinosauridae in several theropod phylogenetic analyses (e.g., Sereno et al., 1998; Holtz et al., 2004). It is thus beneficial to be able to determine how closely these illustrations correspond to the actual anatomy of BSP 1912 VIII 19 as reproduced by the photographs, an effort which holds implications for the use of published drawings as sources of systematic information. DESCRIPTIONS OF THE PHOTOGRAPHS
Figure 2.1 is a photograph of the right mandibular ramus of BSP 1912 VIII 19 in lateral view. Judging from this photograph, it appears that, in general, Stromer’s (1915) illustration (Fig. 2.2) of this element faithfully reproduces its anatomy and state of preservation, accurately recording the distalmost six dentary alveoli and two preserved crowns (Rd4 and Rd12, based on Fig. 2.1 and data from Stromer, 1915; dental nomenclature after Smith and Dodson, 2003). However, there are some differences between the two illustrations (it should be noted that, in Fig. 2.1, what could be interpreted as the fifth dentary crown is actually the shadow of Rd4 falling against the photographic background). The shape of the mandibular ramus is the same in both, but some of the cracks and foramina on the surface of the dentary are only schematically congruent with the photograph (particularly in the caudoventral area). More importantly, the rostral margin of the dentary appears distinctly different in the two images. At the symphysis, the rostral margin is more ‘squared-off’ in Figure 2.2 than in Figure 2.1 (angles of ;718 vs. ;618 measured against a horizontal line set at the ventral surface of the dentary below Rd4). We suspect that this discrepancy might partially result from the two images showing the mandibular ramus from slightly different angles, but proving this is difficult. Moreover, the dorsal surface of the dentary at the base of Rd4 is effectively the same angle in each image (728 in Fig. 2.1 and 708 in Fig. 2.2, in an angle measured between the distalmost point on the crown base of Rd4 and its apex). As such, we believe that the rostral margin of the lower jaw of BSP 1912 VIII 19 was incorrectly illustrated in the figure published by Stromer (1915). As the actual orientation of the rostral margin of the right dentary in BSP 1912 VIII 19 is seemingly more rostrodorsal-caudoventral than previously believed, the mesialmost two dentary
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FIGURE 1—1, Location of the Bahariya Oasis in Egypt, ;320 km southwest of Cairo. 2, Locations of landforms in the northeastern area of the oasis from where material in the BSP collections was found. Dashed line in 2 indicates the road to Cairo; black dot indicates the town of Bawiti (1 modified from Smith et al., 2001; 2 modified from Stromer, 1914b and El-Akkad and Issawi, 1963).
teeth in S. aegyptiacus might have been slightly more procumbent than is common in theropods (see Carrano et al., 2002). This interpretation is consistent with the apparent orientation of the two mesial alveoli as suggested by the occlusal view of this bone in Figure 2.2. It also accords with the orientation of Ld2 in a rostral dentary fragment referred to Spinosaurus cf. aegyptiacus from the Albian-aged Chenini Formation of Tunisia (Buffetaut and Ouaja, 2002). It is certainly possible, and perhaps probable, that the Tunisian specimen does not pertain to the species S. aegyptiacus, given the approximately nine million years (see Gradstein et al., 2005) separating the deposition of the Bahariya (lower Cenomanian) and Chenini (middle Albian) formations. However, the Tunisian element does appear to pertain to the Spinosauridae, and the shape of its rostral margin is extremely similar to that of BSP 1912 VIII 19 (Fig. 2.1). Perhaps the morphology of the rostral end of the dentary and the orientation of the mesialmost dentary teeth of Spinosaurus are distinct from the conditions in baryonychine spinosaurids (see Charig and Milner, 1986; Charig and Milner, 1997; Sereno et al., 1998). More specimens will be needed before this possibility can be further explored. Figure 3 depicts BSP 1912 VIII 19 as mounted in the former Pala¨ontologische Staatssammlung Mu¨nchen. It shows the right mandibular ramus with two erupted crowns and 15 additional teeth. Stromer (1915) discussed 19 teeth in his text (four in situ and 15 isolated, the latter of which are all illustrated in Fig. 3), but his table (Stromer, 1915: 11) includes 20 teeth. Stromer (1915, p. 8) recognized Rd13 and Ld6, 8, 12, 13, and 14 as replacement teeth in the process of erupting, and as such did not consider them to be in situ crowns. He (p. 9) identified Ld3, Rd4, and Rd12 as in situ. At first glance, it is not clear which tooth represents the fourth in situ crown that Stromer (1915, p. 8) referred to. Based on his (1915, p. 11) table, Ld1 and Ld4 could be interpreted as additional in situ crowns (in Stromer’s [1915, p. 11] table, these teeth are not listed with question marks, and consequently their position seems established). Therefore, one of these is probably the aforementioned fourth in situ tooth, but Stromer (1915) did not discuss either of them as such. The left lower jaw was apparently not mounted at the time the photograph was taken, and although Stromer (1915, pl. 1, fig. 6) did illustrate it in medial view, the dentition depicted in that illustration is reconstructed and schematic, and therefore seemingly of limited utility. The
first, third, fourth, and twelfth left dentary teeth are shown as in situ; however, based on the caption for Stromer’s plate 1, figure 6 (1915, p. 32), we suspect that Ld4 and Ld12 were reconstructed from the in situ teeth in the right dentary. Stromer (1915. p. 9) stated that the crown of Ld1, although present, was broken and could not be reattached to the tooth base present in the first left alveolus (he illustrated this crown in pl. 1, fig. 7). This statement, plus the uncertainty of the measurements given for Ld4 in Stromer’s (1915, p. 11) table, indicate that Ld1 was the fourth in situ tooth mentioned by Stromer. The high-spined dorsals illustrated by Stromer (1915, pls. I, II) were mounted, as were the ?axis and ?middle cervical (pl. II, figs. 1, 2), fragmentary sacrals, and a questionably referable proximal caudal (Stromer, 1915, 1934). Figure 3 also shows, along with their neural arches and spines, what appear to be the centra of the three cranialmost dorsals of BSP 1912 VIII 19. These elements appear identical (e.g., their lengths do not vary and they lack obvious cracks). Stromer (1915, pl. I, figs. 17–19) illustrated the unfused neural arches of these vertebrae, stating (p. 14–15, translated) that ‘‘only in [vertebra] ‘f’ (pl. II, fig. 3 a, b) is the neural arch still in natural articulation, in ‘g’ less certainly, in ‘h’ and ‘i’ (pl. II, figs. 5 a, b, and 6) very probably associated, while in ‘c,’ ‘d,’ and ‘e’ (pl. I, figs. 17–19) the centrum has unfortunately been lost.’’ Stromer (1915) lettered the vertebrae of BSP 1912 VIII 19 according to their presumed anatomical position, with ‘‘c’’ representing the cranialmost dorsal, ‘‘d’’ the next cranialmost, and so on. As such, the cranialmost three dorsal ‘centra’ shown in Figure 3 were almost certainly reconstructions. DISCUSSION
Figure 3 depicts the dorsal vertebrae of Spinosaurus in an arrangement reconstructed for museum display, and provides a view into the possible appearance of the dorsal margin of the sail-like structure (but see Bailey, 1997) on the back of the animal. The cranialmost two preserved dorsals (‘‘c’’ and ‘‘d,’’ following the labeling scheme of Stromer [1915], see Fig. 4) were mounted as adjacent vertebrae, which appears accurate given the orientations and relative heights of their neural spines and transverse processes. However, we suspect, as apparently did Stromer (1934, 1936), that the more caudal dorsals were probably positioned incorrectly in the mount.
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FIGURE 2—1, Photograph of the right mandibular ramus of the holotype of Spinosaurus aegyptiacus Stromer, 1915 (BSP 1912 VIII 19), in lateral view, taken at a point prior to its destruction in April 1944. 2, Reproduction of Stromer’s (1915, pl. I, fig. 12a) illustration of the right mandibular ramus. The handwritten caption in 1 translates to: ‘‘Spinosaurus aegyptiacus Stromer, from the lower Cenomanian, Bahariya, G. el Dist.’’ The two measurements given in the caption are ‘‘95 cm long’’ and possibly ‘‘75 cm long.’’
In the mounted skeleton (Figs. 3, 4.1), dorsal ‘‘e’’ was positioned immediately caudal to ‘‘d,’’ followed by ‘‘h,’’ ‘‘f,’’ ‘‘g,’’ and ‘‘i.’’ However, as the neural spine of ‘‘e’’ is broken dorsally, and consequently was longer in life, this vertebra was probably placed more caudally in the series. Stromer (1934, 1936) came to this realization, and positioned ‘‘e’’ and ‘‘h’’ caudal to ‘‘f’’ (Fig. 4.2) in his 1936 reconstruction of the S. aegyptiacus skeleton. He did not, however, change the relative positions of the caudalmost preserved dorsals ‘‘g’’ and ‘‘i.’’ The differing arrangements of the preserved dorsals in the mount (Figs. 3, 4.1) and in the 1936 reconstruction (Fig. 4.2) illustrate the evolution of Stromer’s ideas
regarding the anatomy of this theropod. Stromer’s (1936, p. 65) reconstruction results in a ‘‘sail’’ of Spinosaurus that is less steeply inclined cranially than that inferred from the mounted skeleton in Figure 3 (see Fig. 4). The photo depicted in Figure 3 does not show any skeletal element distal to the proximal caudal vertebra illustrated by Stromer (1915, pl. I, fig. 1), so we cannot presently determine if the more distal caudal vertebrae he illustrated in the 1936 reconstruction were exhibited. Additionally, none of the dorsal ribs or gastralia associated with BSP 1912 VIII 19 (see Stromer, 1915) were apparently mounted at the time the photograph was taken.
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FIGURE 3—A photograph of the holotype of Spinosaurus aegyptiacus (BSP 1912 VIII 19), mounted in the original Pala¨ontologische Staatssammlung Mu¨nchen at a point prior to April 1944.
These photographs of BSP 1912 VIII 19, recently ‘‘rediscovered’’ in the Pala¨ontologische Staatssammlung Mu¨nchen, provide additional insight into the anatomy of the holotype specimen of Spinosaurus aegyptiacus and suggest that care should be taken when gleaning character data from published drawings. They are also of historical significance, in that they offer a rare view of a destroyed museum and one of its interesting exhibits. It is frustrating that so little definitive material has been recovered of Spinosaurus since the loss of the Munich collection, especially considering the recent paleontological attention devoted to the Upper Cretaceous of North Africa. However, Stromer’s successes are heartening, and we can assume that with continued exploration North Africa will produce more informative specimens of this enigmatic and important theropod. ACKNOWLEDGMENTS
We thank J. Milio, M. J. Peltier, M. Hufnail, W. Nothdurft, H.D. Sues, P. M. Barrett, S. E. Novak, J. D. Harris, A. Milner, D. Weishampel, S. Sachs, B. Grandstaff, A. Tumarkin-Deratzian, J. Maisey, and K. Carpenter, respectively, for providing data related to, for assistance or discussion with, and for reviews of, this manuscript. The Bahariya Dinosaur Project (BDP) has been supported by funds to JBS from MPH Entertainment, Cosmos Studios, the University of Pennsylvania, the Andrew W. Mellon Foundation, and the Delaware Valley Paleontological Society, and to P. Dodson (Penn) from the late E. de Hellebranth and the University of Pennsylvania. This note is BDP Contribution 5.
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APPENDIX—Fossil material from the Bahariya Formation cataloged in the Bavarian State Collection of Paleontology (BSP) that survived the bombing of Munich in 1944. No complete pre-World War II list of the BSP Bahariya collection is known, but these 87 specimens are what remains. Higher-level taxonomic assignments reflect current interpretations and, in some cases, differ from BSP specimen labels (see Fig. 1.2 for locality information). Specimen 1912 1912 1912 1912 1912 1912 1912 1922 1911 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912
VIII 46 VIII 47 VIII 47a VIII 48c VIII 50 VIII 32 VIII 552 X 14 XII 1 VIII 31 VIII 35 VIII 37 VIII 45 VIII 44 VIII 537 VIII 538 VIII 539 VIII 540 VIII 541 VIII 542 VIII 543 VIII 544 VIII 545 VIII 546 VIII 551 VIII 135 VIII 136 V1 V2 V 504 V 505 V 506 V 507 V 508 V 509 V 510 V 511 V 512 VIII 40 VIII 41 VIII 52 VIII 504 VIII 505 VIII 506 VIII 507 VIII 508 VIII 513
Genus and Species
Higher Taxon
Asteracanthus aegyptiacus Stromer, 1927 Asteracanthus aegyptiacus Stromer, 1927 Hybodus aschersoni Stromer, 1927 Hybodus aschersoni Stromer, 1927 Hybodus aschersoni Stromer, 1927 Cretolamna appendiculata (Agassiz, 1843) Cretolamna appendiculata (Agassiz, 1843) Lamna planidens (Williston, 1900) Scapanorhynchus subulatus (Agassiz, 1843) Scapanorhynchus subulatus (Agassiz, 1843) Squalicorax baharijensis (Stromer, 1927) Squalicorax baharijensis (Stromer, 1927) cf. Trygon cf. Rhinoptera cf. Rhinoptera cf. Rhinoptera cf. Rhinoptera cf. Rhinoptera cf. Rhinoptera cf. Rhinoptera cf. Rhinoptera cf. Rhinoptera cf. Rhinoptera cf. Rhinoptera cf. Rhinoptera Marckgrafia libyca Weiler, 1935 Marckgrafia libyca Weiler, 1935 Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905)
Hybodontidae Hybodontidae Hybodontidae Hybodontidae Hybodontidae Lamniformes Lamniformes Lamniformes Lamniformes Lamniformes Lamniformes Lamniformes Batoidea indet. Rhinopteridae Rhinopteridae Rhinopteridae Rhinopteridae Rhinopteridae Rhinopteridae Rhinopteridae Rhinopteridae Rhinopteridae Rhinopteridae Rhinopteridae Rhinopteridae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae
Element dorsal spine dorsal spine dorsal spine dorsal spine dorsal spine tooth tooth tooth tooth crown teeth tooth tooth small teeth tooth tooth tooth tooth tooth tooth tooth tooth tooth tooth tooth tooth rostral tooth rostral tooth rostral tooth rostral tooth rostral tooth tooth socket rostral tooth dermal denticle dermal denticle rostral tooth rostral tooth rostral tooth rostral tooth tooth spine vertebra spine spine spine spine spine dermal denticle
Locality Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Bahariya Oasis Gebel Maghrafa Gebel Maghrafa Gebel Mandisha Gebel Maghrafa Bahariya Oasis Bahariya Oasis Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Bahariya Oasis Gebel Maghrafa Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel Maghrafa Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist
406
JOURNAL OF PALEONTOLOGY, V. 80, NO. 2, 2006
APPENDIX—Continued. Specimen 1912 1912 1912 1912 1912 1922 1922 1912 1911 1911 1911 1911 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1912 1914 1914 1922 1911 1911 1911 1911 1912 1912 1912 1912 1912 1912 1912
VIII 514 VIII 533 VIII 534 VIII 535 VIII 536 X6 X 524 VIII 144 XI 7 XI 505 XI 506 XI 8 VIII 3b VIII 5 VIII 6 VIII 7 VIII 9a VIII 10 VIII 13 VIII 14 VIII 16k VIII 17o VIII 547 VIII 548 VIII 549 VIII 550 IV 2a IV 501 X 25 XI 12 XII 42 VIII 502 VIII 173 VIII 122 VIII 574 VIII 575 VIII 576 VIII 577 VIII 578 VIII 177
Genus and Species Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Onchopristis numidus (Haug, 1905) Peyeria libyca Weiler, 1935 ?Ptychodus sp. Schizorhiza stromeri Weiler, 1935 Schizorhiza stromeri Weiler, 1935 Schizorhiza stromeri Weiler, 1935 Ceratodus africanus Haug, 1905 Ceratodus africanus Haug, 1905 Ceratodus africanus Haug, 1905 Ceratodus africanus Haug, 1905 Ceratodus africanus Haug, 1905 Ceratodus africanus Haug, 1905 Ceratodus africanus Haug, 1905 Ceratodus africanus Haug, 1905 Ceratodus africanus Haug, 1905 Ceratodus africanus Haug, 1905 Ceratodus africanus Haug, 1905 Ceratodus africanus Haug, 1905 Ceratodus africanus Haug, 1905 Ceratodus africanus Haug, 1905 Ceratodus africanus Haug, 1905 Ceratodus africanus Haug, 1905 Ceratodus africanus Haug, 1905 Enchodus libycus Quaas, 1902 Enchodus sp. Enchodontidae indet. Enchodontidae indet. ?Pleothodus libycus Libycosuchus brevirostris Stromer, 1914a Libycosuchus brevirostris Stromer, 1914a Libycosuchus brevirostris Stromer, 1914a Libycosuchus brevirostris Stromer, 1914a Libycosuchus brevirostris Stromer, 1914a Aegyptosuchus peyeri Stromer, 1933
Higher Taxon
Element
Locality
Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Sclerorhynchidae Dipnoi Dipnoi Dipnoi Dipnoi Dipnoi Dipnoi Dipnoi Dipnoi Dipnoi Dipnoi Dipnoi Dipnoi Dipnoi Dipnoi Dipnoi Dipnoi Dipnoi Aulopiformes Aulopiformes Aulopiformes Aulopiformes Tselfatiiformes Crocodyliformes Crocodyliformes Crocodyliformes Crocodyliformes Crocodyliformes Crocodyliformes
dermal denticle vertebra vertebra vertebra vertebra transverse process caudal vertebra rostral tooth tooth tooth crown tooth crown tooth crown splenial tooth plate splenial tooth plate tooth tooth splenial tooth plate splenial tooth plate splenial tooth plate palatine tooth plate splenial tooth plate splenial tooth plate splenial tooth plate palatine tooth plate splenial tooth plate splenial tooth plate palatine tooth plate splenial tooth plate palatine tooth plate tooth? palatine tooth barbed teeth barbed teeth toothplate skull and jaws vertebra vertebra vertebra vertebra partial skull
Gebel el Dist Gebel Maghrafa Gebel Maghrafa Gebel Maghrafa Gebel Maghrafa Bahariya Oasis Bahariya Oasis Ain Murun Gebel Hefhuf Bahariya Oasis Bahariya Oasis Bahariya Oasis Ain Murun Gebel el Dist Gebel el Dist Bahariya Oasis Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Bahariya Oasis Gebel Hefhuf Gebel Mandisha Bahariya Oasis Bahariya Oasis Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Gebel el Dist Bahariya Oasis