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J. Paleont., 80(2), 2006, pp. 400–406Copyright q 2006, The Paleontological Society0022-3360/06/0080-400$03.00

NEW INFORMATION REGARDING THE HOLOTYPE OFSPINOSAURUS AEGYPTIACUS STROMER, 1915

JOSHUA B. SMITH,1 MATTHEW C. LAMANNA,2* HELMUT MAYR,3 AND KENNETH J. LACOVARA4

1Department 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, 3Palaontologisches Museum, Bayerische Staatssammlung fur Palaontologie und Geologie,Richard-Wagner-Strasse 10/II, D-80333 Munchen, Germany, and 4Department of Biosciences and Biotechnology, Drexel University,

32nd and Chestnut Streets, Philadelphia, Pennsylvania 19104, USA

INTRODUCTION

IN THE autumn of 1912, the fossil collector Richard Markgraf,with financial support and direction from Bavarian paleontol-

ogist Ernst Freiherr Stromer von Reichenbach and the BavarianAcademy of Sciences, discovered the partial skeleton of a bizarrepredatory dinosaur in Upper Cretaceous (early Cenomanian, ;97Ma, see Ismail et al., 1989; Barakat et al., 1993; El Beialy, 1994,1995; Nabil and Hussein, 1994; Ismail and Soliman, 2001; Ibra-him, 2002; Gradstein et al., 2005) rocks of the Bahariya Forma-tion exposed in the Bahariya Oasis of western Egypt (Fig. 1, seealso Sereno et al., 1998; Nothdurft et al., 2002). This gigantictheropod, Spinosaurus aegyptiacus Stromer, 1915, possessedhighly derived cranial and vertebral features sufficiently distinctfor it to be designated as the nominal genus of the clade Spino-sauridae (Stromer, 1915, 1936). Spinosaurids, currently defini-tively known only from Europe, South America, and Africa, areimportant because of the scarcity of Cretaceous Gondwanan tet-rapod 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 controversysurrounding the postulated paleoecology of these taxa (see dis-cussions in Charig and Milner, 1997; Sereno et al., 1998; Sues etal., 2002). Questions related to spinosaurid paleoecology are par-ticularly important in the Bahariya Formation, where Spinosaurusappears to have shared its habitat (see Stromer, 1936; Smith etal., 2001) with at least two other theropods in the size range ofTyrannosaurus Osborn, 1905 (Bahariasaurus Stromer, 1934 andCarcharodontosaurus Stromer, 1931). Unfortunately, the holotypeand only known indisputable specimen of S. aegyptiacus (BSP1912 VIII 19) was lost during the night of 24/25 April 1944 in aBritish bombing raid of Munich (Nothdurft et al., 2002). Theattack severely damaged the building (dating to ca. 1583, Noh-bauer, 1987) that housed the Palaontologische StaatssammlungMunchen and destroyed most of Stromer’s Bahariya collection(see Appendix 1). Since 1944, new spinosaurid taxa have beendescribed (see Charig and Milner, 1986; Martill et al., 1996; Ser-eno et al., 1998; Sues et al., 2002) and additional material hasbeen referred to Spinosaurus (e.g., Buffetaut, 1989, 1992; Russell,1996; Taquet and Russell, 1998; Benton et al., 2000; Buffetautand Ouaja, 2002), some of it questionably. However, definitive S.aegyptiacus material, or information regarding the original spec-imen of Spinosaurus, has not been forthcoming.

We report here on two photographs of the holotype of Spino-saurus aegyptiacus as it was reposited in the PalaontologischeStaatssammlung Munchen prior to 1944, which we ‘‘rediscov-ered’’ in the archives of the Palaontologisches Museum in June2000, after they were donated to the museum by Ernst Stromer’sson, Wolfgang Stromer, in 1995 (Figs. 2, 3). These are, to our

* Current address: Section of Vertebrate Paleontology, Carnegie Museumof Natural History, 4400 Forbes Avenue, Pittsburgh, Pennsylvania 15213-4080.

knowledge, the only surviving photographs of this, the one irre-futable specimen of S. aegyptiacus, which, prior to the initial printreleases, which we authorized, of the photograph in Figure 3(Prendergast, 2001; Glut and Chiappe, 2003), has been repre-sented only by Stromer’s (1915, 1936) drawings (it has recentlybeen suggested that BSP 1912 VIII 19 may represent a chimeraof more than one dinosaurian taxon [Rauhut, 2003], but exam-ining that idea is beyond the scope of this paper). Aside fromtheir historical significance, these images are important in thatthey permit a direct comparison of several of Stromer’s (1915)illustrations with actual photographs of BSP 1912 VIII 19, there-by providing new insight into the skeleton. This is of note becauseStromer’s (1915) text and illustrations have been used as sourcesof character data for Spinosaurus and the Spinosauridae in severaltheropod phylogenetic analyses (e.g., Sereno et al., 1998; Holtzet al., 2004). It is thus beneficial to be able to determine howclosely these illustrations correspond to the actual anatomy ofBSP 1912 VIII 19 as reproduced by the photographs, an effortwhich holds implications for the use of published drawings assources of systematic information.

DESCRIPTIONS OF THE PHOTOGRAPHS

Figure 2.1 is a photograph of the right mandibular ramus ofBSP 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 pres-ervation, accurately recording the distalmost six dentary alveoliand two preserved crowns (Rd4 and Rd12, based on Fig. 2.1 anddata from Stromer, 1915; dental nomenclature after Smith andDodson, 2003). However, there are some differences between thetwo illustrations (it should be noted that, in Fig. 2.1, what couldbe interpreted as the fifth dentary crown is actually the shadowof Rd4 falling against the photographic background). The shapeof the mandibular ramus is the same in both, but some of thecracks and foramina on the surface of the dentary are only sche-matically congruent with the photograph (particularly in the cau-doventral area). More importantly, the rostral margin of the den-tary appears distinctly different in the two images. At thesymphysis, the rostral margin is more ‘squared-off’ in Figure 2.2than in Figure 2.1 (angles of ;718 vs. ;618 measured against ahorizontal line set at the ventral surface of the dentary belowRd4). We suspect that this discrepancy might partially result fromthe two images showing the mandibular ramus from slightly dif-ferent angles, but proving this is difficult. Moreover, the dorsalsurface of the dentary at the base of Rd4 is effectively the sameangle in each image (728 in Fig. 2.1 and 708 in Fig. 2.2, in anangle measured between the distalmost point on the crown baseof Rd4 and its apex). As such, we believe that the rostral marginof the lower jaw of BSP 1912 VIII 19 was incorrectly illustratedin the figure published by Stromer (1915).

As the actual orientation of the rostral margin of the right den-tary in BSP 1912 VIII 19 is seemingly more rostrodorsal-cau-doventral than previously believed, the mesialmost two dentary

401PALEONTOLOGICAL NOTES

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 oasisfrom where material in the BSP collections was found. Dashed line in 2 indicates the road to Cairo; black dot indicates the town of Bawiti (1modified 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 procumbentthan is common in theropods (see Carrano et al., 2002). Thisinterpretation is consistent with the apparent orientation of the twomesial alveoli as suggested by the occlusal view of this bone inFigure 2.2. It also accords with the orientation of Ld2 in a rostraldentary fragment referred to Spinosaurus cf. aegyptiacus from theAlbian-aged Chenini Formation of Tunisia (Buffetaut and Ouaja,2002). It is certainly possible, and perhaps probable, that the Tu-nisian 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 Cenoman-ian) and Chenini (middle Albian) formations. However, the Tu-nisian element does appear to pertain to the Spinosauridae, andthe shape of its rostral margin is extremely similar to that of BSP1912 VIII 19 (Fig. 2.1). Perhaps the morphology of the rostralend of the dentary and the orientation of the mesialmost dentaryteeth of Spinosaurus are distinct from the conditions in baryon-ychine spinosaurids (see Charig and Milner, 1986; Charig andMilner, 1997; Sereno et al., 1998). More specimens will be neededbefore this possibility can be further explored.

Figure 3 depicts BSP 1912 VIII 19 as mounted in the formerPalaontologische Staatssammlung Munchen. It shows the rightmandibular ramus with two erupted crowns and 15 additionalteeth. Stromer (1915) discussed 19 teeth in his text (four in situand 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 replacementteeth in the process of erupting, and as such did not consider themto be in situ crowns. He (p. 9) identified Ld3, Rd4, and Rd12 asin situ. At first glance, it is not clear which tooth represents thefourth in situ crown that Stromer (1915, p. 8) referred to. Basedon his (1915, p. 11) table, Ld1 and Ld4 could be interpreted asadditional in situ crowns (in Stromer’s [1915, p. 11] table, theseteeth are not listed with question marks, and consequently theirposition seems established). Therefore, one of these is probablythe aforementioned fourth in situ tooth, but Stromer (1915) didnot discuss either of them as such. The left lower jaw was ap-parently not mounted at the time the photograph was taken, andalthough Stromer (1915, pl. 1, fig. 6) did illustrate it in medialview, the dentition depicted in that illustration is reconstructedand schematic, and therefore seemingly of limited utility. The

first, third, fourth, and twelfth left dentary teeth are shown as insitu; however, based on the caption for Stromer’s plate 1, figure6 (1915, p. 32), we suspect that Ld4 and Ld12 were reconstructedfrom the in situ teeth in the right dentary. Stromer (1915. p. 9)stated that the crown of Ld1, although present, was broken andcould not be reattached to the tooth base present in the first leftalveolus (he illustrated this crown in pl. 1, fig. 7). This statement,plus the uncertainty of the measurements given for Ld4 in Strom-er’s (1915, p. 11) table, indicate that Ld1 was the fourth in situtooth 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 proximalcaudal (Stromer, 1915, 1934). Figure 3 also shows, along withtheir neural arches and spines, what appear to be the centra of thethree cranialmost dorsals of BSP 1912 VIII 19. These elementsappear identical (e.g., their lengths do not vary and they lackobvious cracks). Stromer (1915, pl. I, figs. 17–19) illustrated theunfused neural arches of these vertebrae, stating (p. 14–15, trans-lated) that ‘‘only in [vertebra] ‘f’ (pl. II, fig. 3 a, b) is the neuralarch 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 unfortunatelybeen lost.’’ Stromer (1915) lettered the vertebrae of BSP 1912VIII 19 according to their presumed anatomical position, with‘‘c’’ representing the cranialmost dorsal, ‘‘d’’ the next cranial-most, 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 ar-rangement reconstructed for museum display, and provides a viewinto the possible appearance of the dorsal margin of the sail-likestructure (but see Bailey, 1997) on the back of the animal. Thecranialmost two preserved dorsals (‘‘c’’ and ‘‘d,’’ following thelabeling scheme of Stromer [1915], see Fig. 4) were mounted asadjacent vertebrae, which appears accurate given the orientationsand relative heights of their neural spines and transverse process-es. However, we suspect, as apparently did Stromer (1934, 1936),that the more caudal dorsals were probably positioned incorrectlyin the mount.

402 JOURNAL OF PALEONTOLOGY, V. 80, NO. 2, 2006

FIGURE 2—1, Photograph of the right mandibular ramus of the holotype of Spinosaurus aegyptiacus Stromer, 1915 (BSP 1912 VIII 19), in lateralview, 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 mandibularramus. The handwritten caption in 1 translates to: ‘‘Spinosaurus aegyptiacus Stromer, from the lower Cenomanian, Bahariya, G. el Dist.’’ The twomeasurements given in the caption are ‘‘95 cm long’’ and possibly ‘‘75 cm long.’’

In the mounted skeleton (Figs. 3, 4.1), dorsal ‘‘e’’ was posi-tioned 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 probablyplaced more caudally in the series. Stromer (1934, 1936) came tothis realization, and positioned ‘‘e’’ and ‘‘h’’ caudal to ‘‘f’’ (Fig.4.2) in his 1936 reconstruction of the S. aegyptiacus skeleton. Hedid not, however, change the relative positions of the caudalmostpreserved dorsals ‘‘g’’ and ‘‘i.’’ The differing arrangements of thepreserved dorsals in the mount (Figs. 3, 4.1) and in the 1936reconstruction (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 steep-ly inclined cranially than that inferred from the mounted skeletonin Figure 3 (see Fig. 4). The photo depicted in Figure 3 does notshow any skeletal element distal to the proximal caudal vertebraillustrated by Stromer (1915, pl. I, fig. 1), so we cannot presentlydetermine if the more distal caudal vertebrae he illustrated in the1936 reconstruction were exhibited. Additionally, none of the dor-sal ribs or gastralia associated with BSP 1912 VIII 19 (see Strom-er, 1915) were apparently mounted at the time the photographwas taken.

403PALEONTOLOGICAL NOTES

FIGURE 3—A photograph of the holotype of Spinosaurus aegyptiacus (BSP 1912 VIII 19), mounted in the original Palaontologische StaatssammlungMunchen at a point prior to April 1944.

These photographs of BSP 1912 VIII 19, recently ‘‘rediscov-ered’’ in the Palaontologische Staatssammlung Munchen, provideadditional insight into the anatomy of the holotype specimen ofSpinosaurus aegyptiacus and suggest that care should be takenwhen gleaning character data from published drawings. They arealso of historical significance, in that they offer a rare view of adestroyed museum and one of its interesting exhibits. It is frus-trating that so little definitive material has been recovered of Spi-nosaurus since the loss of the Munich collection, especially con-sidering the recent paleontological attention devoted to the UpperCretaceous of North Africa. However, Stromer’s successes areheartening, and we can assume that with continued explorationNorth Africa will produce more informative specimens of thisenigmatic 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 relatedto, for assistance or discussion with, and for reviews of, this man-uscript. The Bahariya Dinosaur Project (BDP) has been supportedby funds to JBS from MPH Entertainment, Cosmos Studios, theUniversity of Pennsylvania, the Andrew W. Mellon Foundation,and the Delaware Valley Paleontological Society, and to P. Dod-son (Penn) from the late E. de Hellebranth and the University ofPennsylvania. This note is BDP Contribution 5.

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FIGURE 4—Schematic reconstructions of the dorsal series of Spinosaurusaegyptiacus, using silhouettes of the vertebrae of BSP 1912 VIII 19traced from the photograph depicted in Figure 3. Vertebral labelingscheme follows Stromer, 1915. The dorsal margin of the sail-like struc-ture implied by each reconstruction is indicated as a curve contactingthe apices of selected neural spines. 1, The dorsal vertebrae as arrangedin the mount shown in Figure 3 (baseline as in the mount). 2, Thedorsal series as reconstructed by Stromer (1936); baseline from Strom-er, 1936; gray silhouettes represent dorsals preserved in BSP 1912 VIII19; white silhouettes represent dorsal vertebrae not preserved in theholotype, but hypothesized to have been present based on the lengthsof centra and orientations of neural spines of the preserved dorsals.

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APPENDIX—Fossil material from the Bahariya Formation cataloged in the Bavarian State Collection of Paleontology (BSP) that survived the bombing ofMunich in 1944. No complete pre-World War II list of the BSP Bahariya collection is known, but these 87 specimens are what remains. Higher-leveltaxonomic assignments reflect current interpretations and, in some cases, differ from BSP specimen labels (see Fig. 1.2 for locality information).

Specimen Genus and Species Higher Taxon Element Locality

1912 VIII 46 Asteracanthus aegyptiacus Stromer, 1927 Hybodontidae dorsal spine Gebel el Dist1912 VIII 47 Asteracanthus aegyptiacus Stromer, 1927 Hybodontidae dorsal spine Gebel el Dist1912 VIII 47a Hybodus aschersoni Stromer, 1927 Hybodontidae dorsal spine Gebel el Dist1912 VIII 48c Hybodus aschersoni Stromer, 1927 Hybodontidae dorsal spine Gebel el Dist1912 VIII 50 Hybodus aschersoni Stromer, 1927 Hybodontidae dorsal spine Gebel el Dist1912 VIII 32 Cretolamna appendiculata (Agassiz, 1843) Lamniformes tooth Bahariya Oasis1912 VIII 552 Cretolamna appendiculata (Agassiz, 1843) Lamniformes tooth Gebel Maghrafa1922 X 14 Lamna planidens (Williston, 1900) Lamniformes tooth Gebel Maghrafa1911 XII 1 Scapanorhynchus subulatus (Agassiz, 1843) Lamniformes tooth crown Gebel Mandisha1912 VIII 31 Scapanorhynchus subulatus (Agassiz, 1843) Lamniformes teeth Gebel Maghrafa1912 VIII 35 Squalicorax baharijensis (Stromer, 1927) Lamniformes tooth Bahariya Oasis1912 VIII 37 Squalicorax baharijensis (Stromer, 1927) Lamniformes tooth Bahariya Oasis1912 VIII 45 cf. Trygon Batoidea indet. small teeth Gebel el Dist1912 VIII 44 cf. Rhinoptera Rhinopteridae tooth Gebel el Dist1912 VIII 537 cf. Rhinoptera Rhinopteridae tooth Gebel el Dist1912 VIII 538 cf. Rhinoptera Rhinopteridae tooth Gebel el Dist1912 VIII 539 cf. Rhinoptera Rhinopteridae tooth Gebel el Dist1912 VIII 540 cf. Rhinoptera Rhinopteridae tooth Gebel el Dist1912 VIII 541 cf. Rhinoptera Rhinopteridae tooth Gebel el Dist1912 VIII 542 cf. Rhinoptera Rhinopteridae tooth Gebel el Dist1912 VIII 543 cf. Rhinoptera Rhinopteridae tooth Gebel el Dist1912 VIII 544 cf. Rhinoptera Rhinopteridae tooth Gebel el Dist1912 VIII 545 cf. Rhinoptera Rhinopteridae tooth Gebel el Dist1912 VIII 546 cf. Rhinoptera Rhinopteridae tooth Gebel el Dist1912 VIII 551 cf. Rhinoptera Rhinopteridae tooth Gebel el Dist1912 VIII 135 Marckgrafia libyca Weiler, 1935 Sclerorhynchidae rostral tooth Bahariya Oasis1912 VIII 136 Marckgrafia libyca Weiler, 1935 Sclerorhynchidae rostral tooth Gebel Maghrafa1912 V 1 Onchopristis numidus (Haug, 1905) Sclerorhynchidae rostral tooth Gebel el Dist1912 V 2 Onchopristis numidus (Haug, 1905) Sclerorhynchidae rostral tooth Gebel el Dist1912 V 504 Onchopristis numidus (Haug, 1905) Sclerorhynchidae rostral tooth Gebel el Dist1912 V 505 Onchopristis numidus (Haug, 1905) Sclerorhynchidae tooth socket Gebel el Dist1912 V 506 Onchopristis numidus (Haug, 1905) Sclerorhynchidae rostral tooth Gebel el Dist1912 V 507 Onchopristis numidus (Haug, 1905) Sclerorhynchidae dermal denticle Gebel el Dist1912 V 508 Onchopristis numidus (Haug, 1905) Sclerorhynchidae dermal denticle Gebel el Dist1912 V 509 Onchopristis numidus (Haug, 1905) Sclerorhynchidae rostral tooth Gebel el Dist1912 V 510 Onchopristis numidus (Haug, 1905) Sclerorhynchidae rostral tooth Gebel el Dist1912 V 511 Onchopristis numidus (Haug, 1905) Sclerorhynchidae rostral tooth Gebel el Dist1912 V 512 Onchopristis numidus (Haug, 1905) Sclerorhynchidae rostral tooth Gebel el Dist1912 VIII 40 Onchopristis numidus (Haug, 1905) Sclerorhynchidae tooth Gebel el Dist1912 VIII 41 Onchopristis numidus (Haug, 1905) Sclerorhynchidae spine Gebel el Dist1912 VIII 52 Onchopristis numidus (Haug, 1905) Sclerorhynchidae vertebra Gebel Maghrafa1912 VIII 504 Onchopristis numidus (Haug, 1905) Sclerorhynchidae spine Gebel el Dist1912 VIII 505 Onchopristis numidus (Haug, 1905) Sclerorhynchidae spine Gebel el Dist1912 VIII 506 Onchopristis numidus (Haug, 1905) Sclerorhynchidae spine Gebel el Dist1912 VIII 507 Onchopristis numidus (Haug, 1905) Sclerorhynchidae spine Gebel el Dist1912 VIII 508 Onchopristis numidus (Haug, 1905) Sclerorhynchidae spine Gebel el Dist1912 VIII 513 Onchopristis numidus (Haug, 1905) Sclerorhynchidae dermal denticle Gebel el Dist

Cenoman). 3. Das Original des Theropoden Spinosaurus aegyptiacusnov. gen., nov. spec. Abhandlungen der Koniglich Bayerischen Aka-demie der Wissenschaften, Mathematisch-physikalische Classe, 28(3)Abhandlung:1–32.

STROMER, E. 1927. Ergebnisse der Forschungsreisen Prof. E. Stromers inden Wusten Agyptens. II. Wirbeltier-Reste der Baharije-Stufe (unter-estes Cenoman). 9. Die Plagiostomen, mit einem Anhang uber kano-und mesozoische Ruckenflossenstacheln von Elasmobranchiern. Abhand-lungen der Bayerischen Akademie der Wissenschaften Mathmatisch-naturwissenschaftliche Abteilung, 31(5):1–64, 63 pls.

STROMER, E. 1931. Ergebnisse der Forschungsreisen Prof. E. Stromers inden Wusten Agyptens. II. Wirbeltier-Reste der Baharije-Stufe (unterstesCenoman). 10. Ein Skelett-Rest von Carcharodontosaurus nov. gen.Abhandlungen der Bayerischen Akademie der Wissenschaften, Math-ematisch-naturwissenschaftliche Abteilung n. f., 9:1–23.

STROMER, E. 1933. Ergebnisse der Forschungsreisen Prof. E. Stromers inden Wusten Agyptens. II. Wirbeltierreste der Baharije-Stufe (unterstesCenoman), 12; die procoelen Crocodilia. Abhandlungen der Bayerisch-en Akademie der Wissenschaften Mathematisch-naturewissenshaftlicheAbteilung n. f., 15:1–55.

STROMER, E. 1934. Ergebnisse der Forschungsreisen Prof. E. Stromers inden Wusten Agyptens. II. Wirbeltier-Reste der Baharije-Stufe (unterstesCenoman). 13. Dinosauria. Abhandlungen der Bayerischen Akademie

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STROMER, E. 1936. Ergebnisse der Forschungsreisen Prof. E. Stromers inden Wusten Agyptens. VII. Baharije-Kessel und -Stufe mit deren Faunaund Flora. Eine erganzende Zusammenfassung. Abhandlungen derBayerischen Akademie der Wissenschaften, Mathematisch-naturwis-senschaftliche Abteilung n. f., 33:1–102.

SUES, H.-D., E. FREY, D. M. MARTILL, AND D. M. SCOTT. 2002. Irritatorchallengeri, a spinosaurid (Dinosauria: Theropoda) from the LowerCretaceous of Brazil. Journal of Vertebrate Paleontology, 22(3):535–547.

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WEILER, W. 1935. Ergebnisse der Forchungsreisen Prof. E. Stromers inden Wusten Agyptens, II. Wirbeltierreste der Baharije-Stufe (unterstesCenoman), 16. Neue Untersuchungen an den Fischesten. Abhandlun-gen der Bayerischen Akademie der Wissenschaften, Mathematische-naturwissenschaftliche Abteilung Neue Folge Hefte, 32:1–57.

WILLISTON, S. W. 1900. Cretaceous fishes. Selachians and pycnodonts.Kansas Geological Survey Bulletin, 6:235–256.

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406 JOURNAL OF PALEONTOLOGY, V. 80, NO. 2, 2006

APPENDIX—Continued.

Specimen Genus and Species Higher Taxon Element Locality

1912 VIII 514 Onchopristis numidus (Haug, 1905) Sclerorhynchidae dermal denticle Gebel el Dist1912 VIII 533 Onchopristis numidus (Haug, 1905) Sclerorhynchidae vertebra Gebel Maghrafa1912 VIII 534 Onchopristis numidus (Haug, 1905) Sclerorhynchidae vertebra Gebel Maghrafa1912 VIII 535 Onchopristis numidus (Haug, 1905) Sclerorhynchidae vertebra Gebel Maghrafa1912 VIII 536 Onchopristis numidus (Haug, 1905) Sclerorhynchidae vertebra Gebel Maghrafa1922 X 6 Onchopristis numidus (Haug, 1905) Sclerorhynchidae transverse process Bahariya Oasis1922 X 524 Onchopristis numidus (Haug, 1905) Sclerorhynchidae caudal vertebra Bahariya Oasis1912 VIII 144 Peyeria libyca Weiler, 1935 Sclerorhynchidae rostral tooth Ain Murun1911 XI 7 ?Ptychodus sp. Sclerorhynchidae tooth Gebel Hefhuf1911 XI 505 Schizorhiza stromeri Weiler, 1935 Sclerorhynchidae tooth crown Bahariya Oasis1911 XI 506 Schizorhiza stromeri Weiler, 1935 Sclerorhynchidae tooth crown Bahariya Oasis1911 XI 8 Schizorhiza stromeri Weiler, 1935 Sclerorhynchidae tooth crown Bahariya Oasis1912 VIII 3b Ceratodus africanus Haug, 1905 Dipnoi splenial tooth plate Ain Murun1912 VIII 5 Ceratodus africanus Haug, 1905 Dipnoi splenial tooth plate Gebel el Dist1912 VIII 6 Ceratodus africanus Haug, 1905 Dipnoi tooth Gebel el Dist1912 VIII 7 Ceratodus africanus Haug, 1905 Dipnoi tooth Bahariya Oasis1912 VIII 9a Ceratodus africanus Haug, 1905 Dipnoi splenial tooth plate Gebel el Dist1912 VIII 10 Ceratodus africanus Haug, 1905 Dipnoi splenial tooth plate Gebel el Dist1912 VIII 13 Ceratodus africanus Haug, 1905 Dipnoi splenial tooth plate Gebel el Dist1912 VIII 14 Ceratodus africanus Haug, 1905 Dipnoi palatine tooth plate Gebel el Dist1912 VIII 16k Ceratodus africanus Haug, 1905 Dipnoi splenial tooth plate Gebel el Dist1912 VIII 17o Ceratodus africanus Haug, 1905 Dipnoi splenial tooth plate Gebel el Dist1912 VIII 547 Ceratodus africanus Haug, 1905 Dipnoi splenial tooth plate Gebel el Dist1912 VIII 548 Ceratodus africanus Haug, 1905 Dipnoi palatine tooth plate Gebel el Dist1912 VIII 549 Ceratodus africanus Haug, 1905 Dipnoi splenial tooth plate Gebel el Dist1912 VIII 550 Ceratodus africanus Haug, 1905 Dipnoi splenial tooth plate Gebel el Dist1914 IV 2a Ceratodus africanus Haug, 1905 Dipnoi palatine tooth plate Gebel el Dist1914 IV 501 Ceratodus africanus Haug, 1905 Dipnoi splenial tooth plate Gebel el Dist1922 X 25 Ceratodus africanus Haug, 1905 Dipnoi palatine tooth plate Bahariya Oasis1911 XI 12 Enchodus libycus Quaas, 1902 Aulopiformes tooth? Gebel Hefhuf1911 XII 42 Enchodus sp. Aulopiformes palatine tooth Gebel Mandisha1911 VIII 502 Enchodontidae indet. Aulopiformes barbed teeth Bahariya Oasis1911 VIII 173 Enchodontidae indet. Aulopiformes barbed teeth Bahariya Oasis1912 VIII 122 ?Pleothodus libycus Tselfatiiformes toothplate Gebel el Dist1912 VIII 574 Libycosuchus brevirostris Stromer, 1914a Crocodyliformes skull and jaws Gebel el Dist1912 VIII 575 Libycosuchus brevirostris Stromer, 1914a Crocodyliformes vertebra Gebel el Dist1912 VIII 576 Libycosuchus brevirostris Stromer, 1914a Crocodyliformes vertebra Gebel el Dist1912 VIII 577 Libycosuchus brevirostris Stromer, 1914a Crocodyliformes vertebra Gebel el Dist1912 VIII 578 Libycosuchus brevirostris Stromer, 1914a Crocodyliformes vertebra Gebel el Dist1912 VIII 177 Aegyptosuchus peyeri Stromer, 1933 Crocodyliformes partial skull Bahariya Oasis