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On the Traces of Cranial Veins in Saurischians and Ornithischians, As Well As Several Other Fossil and Recent Reptiles1

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On the Traces of Cranial Veins in Saurischians and Ornithischians, As Well As Several Other Fossil and Recent Reptiles1 On the traces of cranial veins in Saurischians and Ornithischians, as well as several other fossil and recent reptiles1 By WERNER JANENSCH, Berlin with 5 Figures 1 Original citation: Janensch, W. 1936. Über bahnen von Hirnvenen bei Saurischiern und Ornithischiern, sowie einigen anderen fossilen und rezenten Reptilien. Paleont. Zeitschrift 18(3-4):181-198. Translated 2009 by J.A. Whitlock, University of Michigan. There are not many observations on the traces of veins on the braincases of fossil reptiles to be found in the literature. They cannot be noticable, since in the adult head the intracranial veinous system sends only a very few branches by the skull wall. If, however, foramina are interpretable as insertion places of veins, then their experiences over the ontogenetic development of the intracranial venous system must be considered in our interpretation. C. VAN GELDEREN (1924/25) stated very clearly how this occurs with the different sauropsids and thereby how important elements are backformed to a large extent. Since such evolutionary investigations—understandably—treats only the process of the container, but not or only in small measure the question of whether traces impress themselves into the fully developed bony braincase2, then the interpretation of questionable venous foramina is difficult for the paleontologist 2 [Ed.]—Hirnkapsel-lit. cranium, here considered as ʻbraincaseʼ 182 Werner Janensch, and always afflicted with a fairly large measure of uncertainty. During the investigation in the collections of the Berlin Geological and Paleontological Institute and Museum of skull caps recovered from the sauropods from the Tendaguru layers of German East Africa it resulted that that certain morphologic details can probably be interpreted best as the traces of cranial nerves. I could use the results of the following investigations in the detailed representation of the skulls of the sauropods of the Tendaguru layers (1935). Fig.1. Brachiosaurus brancai JANENSCH. Medial view of the right wall of the brain cavity (Cat. Nr. Y 1). 2/3 nat. size. d.e Foramen for the Ductus endolymphaticus, fe.ep Fenestra epioptica, fs.sa Fossa sub-arcuta, v.c.m Foramina and groove for the Vena cerebralis media, v.c.p Foramen for the Vena cerebralis posterior, lV-XII IV.-XII. Cranial nerves. I begin with the the description of the sometimes very distinct vascular traces, which I believe to be attributable to the Vena cerebralis media. The preparation of the cranium of three genera of East African sauropods exposed a sulcus above the foramen trigemini which travels, in a more or less pronounced arc, upward from the rear wall. In the endocast3, the sulcus presents itself as accordingly formed, raised border or line. Such a sulcus was in the genera Brachiosaurus, Dicraeosaurus and Barosaurus. 3 [Ed.]—Hirnschädelausguß, here interpreted as ʻendocastʼ On the traces of cranial veins in Saurischians and Ornithischians, etc. 183 The traces are notably developed in three braincases of Brachiosaurus brancai JANENSCH. The sulcus has a length of approximately 3 cm, is several millimeters broad and ends both above and below in an approximately 3 mm broad foramen (Fig. 1). On two skulls the lower foramen perforates the wall of the braincase so near to the Foramen trigemini that only a plate of bone a few mm wide remains between them, and opens laterally; on the third skull the lower foramen is also closely placed above the Foramen trigemini in the wall of the brain cavity, but it does not reach the external wall but instead opens already into the upper wall of the Foramen trigemini. Fig. 2. Barosaurus africanus (E. FRAAS). Medial view of the left wall of the brain cavity (Cat.-Nr. dd 316).2/3 nat. size. fe.ep Fenestra epioptica, fe.ov Fenestra ovalis, hy Groove for the hypophysis, p.oe Parietal opening, pp.l Postparietal opening, v.c.m. groove and foramina of the Vena cerebralis media, ve Vestibulum, I-XII I.-XII. Cranial nerves. The upper foramen perforates the supraoccipital not far beneath its upper margin and opens on the external occipital surface4. It is not improbable that a foramen in the back wall of the upper temporal hollow, where the Parietal, Prootic, and Laterospenoid meet, represents the exit of a channel of the upper foramen, which branches within the supraoccipital. With Barosaurus africanus (E. FRAAS) the sulcus can be demonstrated in a large, nearly full-grown skull and a small skull (Fig. 2); 4 [Ed.]—Hinterhauptsfläche, lit. ʻposterior head surfaceʼ 184 Werner Janensch, it has a length of approximately 22 and 19 mm and is much more constricted and more sharply incised than in Brachiosaurus. The lower foramen could not be determined in the larger skull, in the smaller skull it is deeper, but it is assumed the lower end of the sulcus does not widen; also the opening on the external wall was recognizable over the foramen trigemini on one side. The upper foramen pierces with a funnel-shaped enlargement. In the large head, the opening of a 2 mm wide foramen is possible in the exposed sutural surface of the upper margin of the supraoccipital 3 cm distance from the median, perhaps also still farther laterally present for a branching canal. On the small skull, only one opening lies on the suture of the supraoccipital and the suture5 of the parietal is the same distance from the middle. The third genus Dicraeosaurus in the species D. hansemanni JANENSCH possesses a sharply incised sulcus of approximately 27 mm in length (s. W. JANENSCH 1935, fig. 135). At its upper end a small subovate pore penetrates into the thick brain cavity wall, down, 1/2 to 1 cm above the for. trigemini; the sulcus ends in an oblong pore. That the lateral openings of the pores can not be found, may be a consequence of conservation disturbance to the bottom, as it may be that they are always missing. Also with other sauropods, the sulcus and upper foramen come forwards, as F. V. HUENE (1906) first described in the skull of Megalosaurus bucklandi and Cetiosaurus oxoniensis PHILL. from Stonesfield. The endocast of Camarasaurus depicted by F. OSBORN 6 & C. MOOK (1921) shows a projection high above the foramen trigemini, likely corresponding to the foramen at the upper end of the sulcus in Brachiosaurus, although this is not self evident. An indentation is in the appropriate place in the wall of the brain cavity in Diplodocus (F. OSBORN, 1912, Fig. 16 and F. VON HUENE, 1914, Pl. 8, Fig. 3).A sulcus leading to this depression is not to be seen in these cases, based on their position I would however like to equate the indentations with the upper foramen with which the sulcus ends in the Tendaguru Sauropods. These cases mentioned from the literature were brought in connection with the saccus endolymphaticus, a view I cannot follow because of the lack of a local relationship with the ear capsule. With recent reptiles I have not found a description where the appropriate sulcus is indicated. I could determine, meanwhile, 5 [Ed.]—Spange, lit. ʻbuckleʼ or ʻhaspʼ 6 [Ed.]—Here and elsewhere, F. Osborn refers to H.F. Osborn On the traces of cranial veins in Saurischians and Ornithischians, etc. 185 that in a cut through the medial plane the skull of a Crocodilus niloticus from the Tendaguru area exhibits a 39 cm long (Nose-Condyle) sulcus in the same place as in the sauropods; it lies beside the dorsally running suture between the prootic and the laterosphenoid and is quite easily recognized. Their lengths amount to 7 mm on the right and 8 mm on the left. Also directly next to each other two small foramina are at the upper and and likewise a very small pore at the lower end, about 7 mm above the foramen trigemini, impressed into the skull wall under the margin of the laterosphenoids. A second smaller crocodile skull exhibited the sulcus also. In order to determine the meaning of the sulcus, I examined the alcohol preserved preparation of an average skull of Crocodylus americanus in the exhibition collection of the Berlin Zoologicial Museum7. It appears here that the contents of this sulcus are a tissue that cannot be recognized by the shape of the container. It is questionable to me whether the investigation of a fresh head would obtain a different result; nevertheless it would be very desirable, in order to finally clarify the question. Now the attempt is made to come to the solution of the question by comparing anatomical details coming from use of the zoological literature. The sulcus and associated foramina cannot be placed in a relationship with any cranial nerves under any circumstances. Also, the sulcus does not lead back to the Ductus endolymphaticus. Apart from the fact that the lower foramen would be inexplicable, the sulcus lies much too far away from the inner upper chamber; in Brachiosaurus I regard the foramen medially perforating the bony wall as the outlet of the D. endolymphaticus, according to its position as conditions of the inner ear clearly show. If, then, the only remaining possibility is to regard the sulcus and the foramina as a passage for blood vessels, then the arteries might be insufficient for it. The lumen of the foramina of Brachiosaurus corresponding with powerful arteries with such a peculiar route is not likely based on conditions in living reptiles. If we regard the venous system of the head in reptiles, then it seems to me that the local relationship of the lower foramina of the sulcus to the foramen of the trigeminus, 7 I am very much obligated to thank Dr. AHL, the manager of the reptile department of the Berlin Museum, and the head preparator Mr.
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