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Dryosaurus and , intercontinental genera of Upper ornithopod

Article · January 1980

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Dryosaurus and Camptosaurus, intercontinental genera of Upper Jurassic ornithopod dinosaurs by PETERM. GALTON *

Abstract. - The IZimmeridgian (Upper Jurassic) ornithopod dinosaurs Dryosaurus and Camptosaurus occur in western (D. altus, C. dispar), western Europe (C. prestwichii, C. sp.) and East Africa (D. lettowuorbecki). The hypsilophodontids Dryosaurus altus and D. lettowvorbeclci are almost conspecific and this extreme similari-ty indicate the presence of a land connec.tion between northern Laurasia and southern Gondwanaland sometime in the Icimmeridgian (middle Upper Jurassic). The only correct referrals of European specimens to the North American iguanodontid Camptosaurus are .the nearly comple-te slieleton of C. prestwichii (HULICE)from England and a newly described femur from Portugal. The skeletons of the North American and European of Camptosaurus differ in several respects and this probably indicates the presence of a land connection sometime in the (lower Upper Jurassic). The land connections were probably different with that for Dryosaurus being across the western end of the central Atlantic and that for Camptosaurus bei,nng across the North Atlantic epicontinental seaway.

Dryosaurus et Camptosaurtas, genres intercontinentaux de Dilnosaures ornithopodes du Jurassique superieur RisumC. - Les Dinosauriens ornithopodes kimmeridgiens (Jwassique supkrieur) Dryosaurus el Camptosaurus sont connus dans l'0uest de I'Amerique du Nord (D. altus, C. dispar), en Europe occidentale (C. prestwichii, C. sp.) et en Afrique orientale (D. lettowvorbecki). Les Hypsilo- phodontides Dryosaurus altus et D. lettowvorbecki sont presque conspecifiques et cette extr&me similaritk indique la prdsence d'une connexion terrestre entre la Laurasie au Nord et le Gondwana au Sud a un moment du Kimmeridgien (milieu du Jurassique supirieur). Les seuls specimens europeens rapport& avec raison au genre d'lguanodontide nord-amkricain Cumptosaurus sont le squelette presque complet de C. presfwichii (HULKE)d'Angleterre et un femur du Portugal recemment decrit. Les squelettes des espkces nord-amkricaines et europkennes de Cumptosaurus different 2? plusieurs points de vue et ceci indique probablement la presence d'une connexion terrestre a un momenl de l'oxfordien (dkbut du Jurassique superieur). Ces connexions terrestres ktaient probablement differentes, celle pour Dryosaurus se trouvant au travers de l'extremitd occidentale de l'Atlantique central, et celle pour Camptosaurus se pla~anth. travers le bras de mer epicontinental nord-atlantique.

Druosaurus. horizon and continent then it would all be referred to Dryosaurus altus. Janensch [l9551 described much of the ornithopod The skulls of the two species of Dryosaurus are very material from the Tendaguru Beds (Upper Jurassic, Upper distinct from those of other hypsilophodontids and show = Lower [Hallam, 19751) of only a few diflerences (fig. 1A-D), some of which (posterior Tanzania, East Africa. He referred it all to region, fig. 1A, C ; width fig. lB, D) result from crushing Zettow-vorbecki POMPECICJ[l9201 that, as shown by Galton of the Morrison skull. In addition, two skulls of Hypsilo- [l977 a], is referable to the genus Dryosaurus MARSH phodon foxii show greater differences in sutural pattern (1894) that was previously unknown outside of the anteroventral to the orbit (fig. 3 A, B ; Galton [1974]) than Morrison Formation (Upper Kimmeridgian) of the western do those of Dryosaurus (fig. lA, C). The main character interior of the of America. The correct that distinguishes the skull of Dryosaurus altus from that name for the Tendaguru hypsilophodontid is Dryosaurus of D. lettowuorbecki is the difference in length of the supsa- Zettowvorbecki (VIRCHOW,1919) [Galton in press a]. The orbital (fig. lA, C). The crowns of the teeth are lozenge- type material (YPM 1876, 1884, not YPM 1915 as given shaped (figs. 1E-H, 2E-G) with the central ridge of a by Galton [l977 a, b]) of altus MARSH(1878), maxillary tooth (fig. lE, G) being as prominent as that of the type species of the genus Dryosaurus MARSH(1894), a dentary tooth (figs. IF, I-I, 2E-G). The deltopectoral will be described elsewhere with supplementary infor- crest of the humerus is low (fig. 11-L). In the pelvic girdle mation on D. Zettowvorbecki [Galton, in press a]. the main body of the ilium is low with an obliquely trun- Hypsilophodontid dinosaurs are particularily suitable cated posterior end (fig. 1M-P) and the brevis shelf is for paleobiogeographic work because the forms of the broad (Q. IN, 0) ; the obturator process of the ischium skull, teeth, pelvis and femur is distinctive for each genus. is proximally placed and the distal half is curved and bar- Dryosaurus is especially important because both species shaped (fig. IM, N) ; and the anterior process of the are well represented (figs. 1A-Y, 2E-G), so that almost pubis is slender with a variably developed ventrolateral the complete anatomy plus some data on individual edge (figs. lM, N, Q, R). The femora (fig. IS-W) possess variation (fig. 1I-W) is documentable for each species the following suite of characters : a deep cleft separating [Galton, in press a]. The diflerences between the two the lesser and greater trochanters (fig. IS, T) ; the deep species of Dryosaurus (figs. 1A-Y, 2E-G) are minor [Galton, depression level with the proximally placed fourth tro- in press a] when one considers three points. Firstly, there chanter is set well anteriorly on the shaft (fig. IS, T) ; is an extremely long distance between the two localities distally (fig. IS-W) the medial edges of the medial condyles even when the continents are reassembled in their Upper are squared off, there is an anterior intercondyle groove, Jurassic positions (over 11,000 km ; see maps in Hallam and the posterior lateral condyle is thin. In the pes, the [1975, 19771, Galton [l977 a, b]). Secondly; -%hew-a-r-e- first m-eta"carsar is-rudimentary -sethere were only three almost as many differences between individuals of the movable toes (fig. lX, Y). Lower foxii (fig. 3 ; Galton An incomplete tibia (fig. 25, K) from the Lower Oxford [1974]), all the material of which came from a bed less Clay (, upper Middle Jurassic) near Peterborough, than 1 m thick in a 0,s km length of sea cliff in southern England. Thirdly, each species shows a similar range of individual variation for those elements that are well repre- * Department of Biology, University of Bridgeport, Bridgeport, sented (e.g. humerus, ilium, pubis, femur, fig. 1 I-W, Ct. 06602, U.S.A. Galton in press a). If this material all came from the same Note presentee a la seance du 5-9 septembre 1978. P. M. GALTON

FIG. 1. - Comparative anatomy of hypsilophodontid dinosaurs Dryosaurus alfus (MARSH,1878) from Morrison Formation of western U.S.A. (A, B, G-J, M, 0, Q, S, U, V, X ; identifying letters underlined to facilitate comparisons) ; Dryosaurus Ieffowvorbeeki (Vr~c~ow,1919) from Tendaguru Beds of East Africa (C-F, K, L, N, P, R, T, W, Y) ; and holotype of eanalieulafus (GALTON,1975) from Wealden (Lower Cretaceous) of England (Z) in lateral view (A, C, E, G, I-R), medial view (F, H, S, T, Z), dorsal view (B, D, N, 0, X, Y) and/or distal ends (I, K, L, S-W, Z). All bones drawn as if from left side except medial views with distal views that are from right side. A, skull of CM 3392 with quadrate YPM 1876 ; B, skull CM 3392, C, D, skull HMN dyA, modified after Janensch [l9551 ; E, slightly worn maxillary tooth after Janensch [l9551 ; F, moderately worn dentary tooth after Janensch [l9551 ; G, unworn maxillary tooth YPM 1876 ; H, unworn dentary tooth YPM 1876 ; I, hume- rus YPM 1876 ; J. humerus AMNH 834 ; K, humerus UT 1495122 ; L, humerus UP 1495123 ; M, pelvic girdle AMNH 834 ; N, reconstruction of pelvic girdle (HMN dyI, dyII, dy35) with dorsal view of ilium (HMN dyII) : 0, ilium DNM 1016 in dorsal and lateral views ; P, ilium UT 149519 ; Q, pubis YPM 1884 ; R, pubis UT 149516 ; S, femur YPM 1876 ; T, femur HMN dy 36 ; U, femur CM 21786 ; V, femur AMNH S34 ; W, femur UT 1495114 ; X, pes CM 21786 ; Y, pes HMN dyV ; Z, femur BMNH R185, after Galton 119751. Abbreviations :a, anterior intercondylar groove ; ap, anterior process of pubis ;b, brevis shelf ; d, deltopectoral crest ; f, fourth trochanter ; g, greater trochanter ; i, ilium ; is, ischium ; 1, lesser trochanter ; m, depression for M. caudi-femoralis longus ; o, obturator process ; p, pubis ; S, supraorbital; 1, vestigal first metatarsal. Scale lines represent 5 cm (A-D, 1-2) or 1 cm (E-H), from Galton [l977 b].

England is similar to that of the holotype of Dryosaurus Gilmore [l9091 provisionally accepted C. prestwichii alfus but this specimen is probably generically and speci- (HULRE)as a valid species of Camptosaurus on the basis fically indeterminate [Galton, 1977 b] and it may be of the original description in which most of the skeleton referable to (( Camptosaurus s Zeedsi LYDEKKER(1889) was not illustrated. Camptosaurus is one of the few dino- (fig. 3G). saurian genera cited [Charig, 19731 from the Upper Jurassic of both sides of the North Atlantic. A redescrip- Camptosaurus. tion of the holotype shows that the Cumnor ornithopod is correctly referred to the American genus Camptosaurus Outside of western North America and__E_a&Africa, as C. presiwichii [Galton and Powell, in press]. Confir- only two ornithopod skeletons have been reported from mation of the presence of the-Morrison genus Campto- the Upper Jurassic but, because Sanpasaurus yaoi YOUNG saurus in England provides an important piece of evidence (1944) from China is based on bones of a sauropod for the presence of a land connection between North [Rozhdestvenskii, 19671, the skeleton from the Lower America and western Europe sometimes in the Late Kimmeridge Clay of Cumnor near Oxford, England is Jurassic. unique. This skeleton is the holotype of presfwi- The skeleton of Camptosaurus prestwichii is similar to chii HULKE(1880), the type species of Cumnoria SEELEY those of the C. dispar (MARSH,1879) from the Morrison (ISSS), and it has been referred [LYDEI~IER1889, 18901 Formation [see Gilmore, 19091. The skull is low and broad, to the North American genus Camptosaurus MARSH(1885). that is an unusual condition for an ornithopod, and the FIG. 2. - A-D, teeth of Camptosaurus, lateral views of maxillary tooth crowns (A, B) and medial views of dentary tooth crowns (C, D) of C. dispar (YPM 7416, USNM 4281) from the Morrison Formotion of western U.S.A. (A, C) and C. prestwichii (OUM 5.3303) from the Lower Kimmeridge Clay of Cumnor near Oxford (B, D) ; E-G, medial view of right dentary teeth of Dryosaurus ; E, F, D. altus from Morrison Forma- tion of western U.S.A. E, YPM 1876, F, CM 3392 ; G, D. lettowvorbecki (HMN dy 14) from Tendaguru Beds of East Africa ; H, I right femur of Camptosaurus (SGP) from Kimmeridgian of Portugal in H, medial view and I, anterior view with distal end ; J, I<, left tibia (CUM 5.46889) of hypsilophodontid from Oxford Clay of England in J, posterior and K, lateral views. Scale lines represent 1 cm (A-G) or 5 cm (H-10.

FIG.3. - Individual variation in Hypsilophodon foxii from the Lower Cretaceous near Cowleaze Chine, Isle of Wight, England. A, B, reconstruction of skull with maxilla and jugal based on A, R2477 ; B, R197 ; C, D, scapula and coracoid : C, R5830 ; D, R196 ; E, F, humerus : E, R196 ; F, R5830 ; G, H, pelvic girdle : G, R195 ; H, R196 ; I, ischium of R5830 ; J, ilium of R2477, K, metatarsus of R5830 ; L, pes of R196. Abbreviations : j, jugal ; m, maxilla. All bones from left side, E and F in lateral view with proximal and distal ends, K and L in dorsal view, rest in lateral view, originals in BMNH, scale lines represent 2 cm, from Galton [l977 b].

MEM. Soc. GEOL. DE FRANCE.- N. S. - T. LIX. MEM. NO 139. - 14

-. P. M. GALTON

A, C, E after Gilmore [1909], K after Gilmore [1912], F, I, L holotype YPM 1877, scale lines represent 10 c,m, from Galton and PoweU (in press).

outlines of the bones are similar (fig. 4A, B). The main pl. 7, figs. 2-61 referred to Iguanodon prestwichii are pro- difference is that the intramaxillary cavity (situated bably iguanodontid but are generically and specifically ventral to the antorbital cavity) is proportionally very indeterminate. The caudal vertebra (Sauvage 1897-1898, much smaller in C. prestwichii than it is in C. dispar pl. 7, figs. 7-12) is that of a theropod dinosaur and it is the (maximum antero-posterior length about one maxillary holotype of the coelurid Caudocoelus sauuagei HUENE crown width-in the former and about five crown widths (1932). Two associated sacral vertebrae from the Kimme- in the latter). The jaws each bear fifteen or. sixteen teeth, ridgian of Porto-de-Mos (Leira) Portugal (Sauvage 1897- the form of which is very similar in both species (fig. 2A-D).. 1898, pl. 7, fig. 1). are regarded as those of a juvenile The vertebral count is the same with nine cervicals, sixteen stegosaur (Dacentrurus (= Omosaiirus) lennieri) by Lappa- dorsals, six or seven sacrals and about forty five caudals. rent and Zbyszewski [1957, .p. 51.1.Apart from Campto- The humerus has a low deltopectoral crest (fig. 4 K-M). . saurus prestwichii, the other species. reported from Europe " In the pelvic girdle (fig. 4C-E) the deep ilium has a narrow [Gilmore, 1909 ; 'Steel, 1969 ; Charig, 19731 are incorrectly . brevis shelf posteriorly ; thehubis has an evenly expanded '. referred to this genus [Galton and Powell, in press]. blade-like anterior process and the distal ends of thelong Campfosaurus inkeyi NOPCSA (1900) from the Upper posterior processes were united together ; arid the ischium Cretaceous of Hungary is based on a dentary of has an anteriorly 'placed obturator process and a bar- (NOPCSA1904, = ). Campfosaurus ualdensis shaped distal end. Distally the femur ,(fig. 4F, H) has a -LYDEKI

. ..

.- - - UPPER JURASSIC ORNITHOPOD DINOSAURS 107 is extremely similar to that of D. letiowvorbecki (VIRCHOW) sible to restore the coast lines in this area accurately as from the Tendaguru Beds (Upper Kimmeridgian) of a result of the complexity of movements of the many Tanzania, East Africa (figs. 1A-Y, 2E-G ; Galton in press a). small areas involved in the Alpine System orogeny [Dewey The anatomy of Camptosaurus prestwichii (HULKE)from et al., 19731. If a European route was involved, then a the Lower Kimmeridgian (about 141 m.y., van Hinte land bridge emerged from the shallow sea between Europe [1976]) of England also differs in several respects from and North Africa for a brief period or periods in the Upper that of the slightly more recent C. dispar (MARSH)[figs. 2A- Jurassic. However, the maximum extent for Jurassic D, 4, Galton and Powell, in press]. The difference in seas was reached during the Callovian, Oxfordian and build probably represents a relatively short term evolu- Kimmeridgian, when between a quarter and a third of tionary change and a gracile and a massive species of the total continental area was covered [Hallam, 1969, Iguanodon lived together in the Early Cretaceous of 19751, and the continental area showing most submergence western Europe [Casier, 19601. However, the reduction was western Europe and northern Africa. It is considered in size of the intramaxillary cavity with additional ossi- improbable that an area characterized by such extensive fication lateral to the tooth row is an advanced character submergence during the Callovian, Oxfordian and Kimme- for an ornithopod [see Galton, 19741. Consequently, it is ridgian would have been the location of the Morrison- surprising that this reduction occurs in Camptosaurus Tendaguru land route [Galton, 1977 a, b]. prestwichii rather than in the more recent C. dispar. The The primitive cetiosaurid sauropod Amygdalodon is higher degree of evolutionary divergence of the two known from the Middle Jurassic of Patagonia, Argentina species of Camptosaurus is not a function of distance, [see Casamiquela, 19631 and Bonaparte [1978, p. 341 because the Upper Jurassic positions are only half as far reports the discovery of a group of Middle Jurassic loca- apart as were those of Dryosaurus [see maps in Cox, 1974 ; lities (Bathonian-Callovian) with (( an almost complete Galton, 1977 a, b ; Hallam, 19771, but it may be that skeleton of a carnosaur (not ) and four incom- the two species of Camptosaurus were isolated from each plete sauropods (cetiosaurids and ? ?). No dinosaurs have other for 'a longer period of time than was the case for yet been reported from the Jurassic deposits of Central the species of Dryosaurus. America or northern South America [Charig, 19731, but The presence of the hypsilophodontid dinosaur Dryo- the possibility exists that the Morrison-Tendaguru land saurus and other similarities in the Morrison and Tenda- connection involved the route North America-Soutll guru faunas [Galton, 1977 a, b] clearly establishes the America-Africa [Galton, 1977 a, b]. The southern end of availability of a land route between Laurasia and Gond- the South Atlantic opened first, but a land route was wanaland sometime in the Upper Jurassic. In calculating still present between South America and Africa in the the Morrison and Tendaguru endemics, I erroniously used Lower Cretaceous (, Buffetaut and Taquet [1977]) the world distribution of the genera and families rather with the earliest connection between the Central and than the distribution between these two faunas (Galton, South Atlantic occurring slightly more recently in the 1977 a, b). The revised fauna1 comparisons (table I) AIbian [Reyment, 19691. If the Morrison-Tendaguru connection was via South America then the Central GENERA FAMILIES Atlantic linking the Pacific to Tethys was a barrier for Morrison fauna (N or N,) Dryosaurus. Hallam [l9771 notes that the Central Atlantic Tendaguru fauna (N or NI) was probably represented in the Lower and Middle Jurassic Morrison endemics (E) (latest Aalenian and early Bajocian) by a well-developed Tendaguru endemics (E) Taxa in common (C) epicontinental seaway that then became restricted or Simpson's coefficient (C/N,) X 100 intermittently closed until the Mid-Callovian. However, Coefficient of difference (1-CIN,) X 100 true oceanic crust was not created in the Central Atlantic, Degree of (E/N) X 100 Morrison by the movement apart of North America and South endemism Tendaguru America plus Africa, until late Middle Jurassic (Upper TABLEI. - Comparisons between Morrison and Tendaguru dinosaur Callovian) or, more definitely, early times faunas (for table of genera and families see Galton [l977 a, b]). (Oxfordian about 149 m.y., van Hinte [1976]). A barrier to gene flow for Camptosaurus was the North Atlantic provide evidence for a land connection between Laurasia epicontinental seaway that, on the basis of evidence from and Gondwanaland sometime during the Middle or Upper marine molluscs, was established by the Pliensbachian Jurassic. However, the absence of three dinosaurs well (183 m.y.) at the latest and persisted tl~roughout the represented in the Tendaguru fauna, the sauropods Jurassic [Hallam, 19771. It is apparent that the excessi- Dicraeosaurus and Tornieria plus the stegosaur Kentro- vely long times involved (40 m.y. for Camptosaurus, saurus, from the much better sampled Morrison Forma- 10 m.y. for Dryosaurus) indicate that the dispersal of tion indicates that there were barriers to the dispersal these genera did not occur prior to the origin of the sea- of these dinosaurs during the Middle and Upper Jurassic. ways. A land route across the North Atlantic epiconti- A North America-Europe-Africa route is usually assumed nental seaway was probably available for the dispersal with the connection located in southwest Europe and of Camptosaurus some time in the Oxfordian (about northern Africa [Charig, 1971 ; Colbert, 1973 a, b ; Cox, 144 m.y.) and for Dryosaurus across the western end of the 1974; Hallam, 1972, 19751. Support for--pa-&--ef-&his- Central _At&.nnic sometime .in IGmmeridgian (about route is provided by the occurrence of certain Morrison 140 m.y.). genera in western Europe : the sauropods Acknowledgements : I thank the following people for and from the Upper Kimmeridgian of all their help while studying the specimens figured in their Portugal [Lapparent and Zbyszewski, 1957 ; referral to respective institutions indicated by the abbreviations : these genera may be incorrect] plus the ornithopod AMNI-I, American Museum of Natural History, New Camptosaurus. However, Jurassic deposits in the critical York (E. S. Gaffney) ; BMNH, British Museum (Natural region (southern Spain and Morocco) are largely marine History), London (A. J. Charig, A. C. Milnar, C. A. Wal- [Hallam, 1972 ; Arkell, 19561 and, in addition, it is impos- ker, M. Holloway) ; CM, Carnegie Museum, Pittsburgh 108 P. M. GALTON

(C. Black, D. S. Berman) ; CUM, Cambridge University Yale for providing (along with P. Dodson, University of Sedgewick Museum (C. L. Forbes); DNM, Dinosaur Pennsylvania, Philadelphia) copies of unpublished figures National Monument, Jensen, (R. Icing, J. Adams) ; prepared under the direction of Prof. 0. C. Marsh that HMN, Humboldt Museum fiir Naturltunde, East Berlin were used as a basis for figures 4F, I and L. H. P. Powell (W.-D. Heinrich, J. Helms, I-I. Jaeger) ; OUM, Oxford of Oxford kindly supplied pencil drawings of Camptosaurus University Museum (H. P. Powell) ; SGP, Services Geo- presfwichii, a full description of .vvhicl~will be published logiques du Portugal, Lisboa (G. Zbyszewski) ; USNM, elsewhere [Galton and Powell, in press]. Figures 1, 3 and 4 United States National Museum, Washington D. C. were drawn by Joan Nicoletti of the University of Bridge- (N. Hotton, 111, R. Purdy) ; UT, Universitat der Tiibingen port and the manuscript was typed by C. Saslafsky of (F. Mrestphal) ; YPM, Peabody Museam Yale University, Orange. This research was supported by USA NSF grant New Haven (J. H. Ostrom). I thank J. H. Ostrom of nos. DEB 76-09769 and DEB 77-24045.

References

ARKELLW. J.'(1956). - Jurassic Geology of the World. Oliver and HALLAMA. (1977). - Biogeographic evidence bearing on the creation Boyd, Edinburgh. of Atlantic seaways in the Jurassic. Milwaulcee Pub. Mus. BONAPARTEJ. (1978). - A-gentina. Soc. Vert. Paleo. News Bull. Spec. Publ. Biol. GeoL, no 2, p. 23-34. no 114, p. 34-35. HUENE F. von (1932). - Die fossile Reptil-Ordnung Saurischia, BUFFETAUTE. and TAQUETPh. (1977). - The giant crocodilian ihre Entwicklung und Geschichte. Monogr. Geol. Palaont., Sarcosuchus in the early 'cretaceous of Brazil and Niger. V. 4, p. 1-361. Palaeontology, v. 20, p. 203-208. HULKEJ. W. (1880). - Iguanodon p~esfwichii,a new species from CASAMIQUELAR. M. (1963). - Consideraciones acerca de Amygdalo- the Kimmeridge Clay founded on numerous fcssil remains don cabrera (Sauropoda, Cetiosauridae) del JurAsico Medio lately discovered at Cumnor, near Oxford. Q. JI geol. Soc. de la Patagonia. Ameghiniana, v. 3, p. 79-95. Lond., v. 36, p. 433-456. CASIERE. (1960). - Les de Bernissart, Brussels, 134 p. JANENSCH W. (1955). - Der Dysalotosaurus der CHARIGA. J. (1971). - Faunal provinces on land : evidence based Tendaguru-Schichten. Palaeontographica, Suppl. 7, E.R. 3, on the distribution of fossil tetrapods, with special refe- p. 105-176. rence to the reptiles of the and Mesozoic, p. 111- LAPPARENTA. F. de and ZBYSZEWSKIG. (1957). - Les dinosauriens 128, in F. A. MIDDLEMISSet al. (eds). Faunal provinces du Portugal. Mems Servs geol. Port (N.S.), no 2, p. 1-63. in space and time. Geol. J. Spec. Issue, no 4, Liverpool. LYDEKKERR. (1889). - On the remains and affinities of five genera CHARIGA. J. (1973). - Jurassic and Cretaceous dinosaurs, p. 339- of Mesozoic reptiles. Q. Jl geol. Soc. Lond., v. 45, p. 41-59. 352. In HALLAMA. (ed.). Atlas of Palaeobiogeography. LYDEKKERR. (1890). - Catalogue of the fossil Reptilia and Amphi- Elsevier, London. bia in the British MJS~U~,Part 4, London, 295 p. COLBERTE. H. 11973 a). - Wandering lands and . Dutton, MARSH0. C. (1878). - Principal characters of American Jurassic New ~ork,32j p. dinosaurs. Amer. J. Sci., (3), v. 17, p. 86-92. COLBERTE. H. (1973 b). - Cantinental drift and the distribution MARSH0. C. (1879). - Notice of new Jurassic reptiles. Amer. J. of fossil reptiles. p. 395-412. In TARLINGD. H. and Sci., (3), v. 18, p. 501-505. RUNCORNS. K. (eds) Implications of continental driIt to MARSH0. C. (1885). - Names of extinct reptiles. Amer. J. Sci., the earth sciences I. Academic Press, London. (3), v. 29, p. 169. Cox C. B. (1974). - Vertebrate palaeodistributional patterns and MARSH0. C. (1894). - The typical Ornithopoda of the American continental drift. J. Biogeogr., v. 1, p. 75-94. Jurassic. Am. J. Sci., (3), v. 48, p. 85-90. DEWEYJ. F., PITMANW. C., RYANW. B. F. and BONNINJ. (1973). - Plate Tectonics and the evolution of the Alpine System. NOPCSAF. (1900). - Dinosaurierreste aus Siebenburgen (Schadel Bull. Geol. Soc. Am., v. 84, p. 3137-3180. von Limnosaurus transsylvanicus nov. gen. et. spec.). GALTONP. M. (1974). - The ornithischian dinosaur Hypsilophodon Denlcschr. Akad. Wiss. Wien., v. 67, p. 555-591. from the Wealden of the Isle of Wight. Bull. Brit. Mus. NOI'CSAF. (1904). - Dinosaurierreste aus Siebenburgen I11 (Weitere (Naf. Hist.) Geol., v. 25, p. 1-152. Schadelreste von Mochlodon). Denkschr. Akad. Wiss. Wien., GALTONP. M. (1975). - English hypsilophodontid dinosaurs (Rep- V. 74, p. 229-263. tilia : ). Palaeontology, v. 18, p. 741-752. POMPECKJJ. F. (1920). - Das angebliche Vorkommen und Wandern GALTONP. M. (1977 a).- The ornithopod dinosaur Dryosaurus des Parietalforamens bei Dinosaurien. Sitz. Ber. naturforsch and a Laurasia-Gondwanaland connection in the Upper Fr. Berlin for 1920, p. 109-129. Jurassic. Nature Lond., v. 268, p. 230-232. REYMENTR. A. (1969). - Ammonite biostratigraphy, continental GALTONP. M. (1977 b). - Upper Jurassic ornithopod dinosaur drift and oscillatory transgressions. Nature, Lond., v. 224, Dryosaurus and a Laurasia-Gondwanaland connection. p. 137-140. Milwaukee Pub. Mus. Spec. Publ. Biol. Geol., no 2, p. 41-54. ROZHDESTVENSKIIA. I<. (1967). - New Iguanodonts from central GALTONP. M. (in press). - The hypsilophodontid dinosaur Dryo- Asia. Int. Geol. Rev., v. 9, p. 556-566. saurus from the Upper Jurassic of western North America SAUVAGEH. E. (1897-98). - Vertkbrks fossiles du Portugal. Contri- and East Africa. Paluont. Zeif. butions Q 1'6tude des poissons et des reptiles du Jurassique GALTONP. M. and POWELLH. P. (in press). - The ornithischian et du Crktacique. Mems. Servs. geol. Port. for 1897-98, dinosaur Campfosaurus presfwichii from the Upper Jurassic p. 1-46. of England. Palaeontology. SEELEY H. G. (1888). - On Cumnoria, an iguanodont founded GILMOREC. W. (1909). - Osteology of the Jurassic reptile Campto- upon the Iguanodon prestwichi, HULKE.Rept. Br. Assn. saurus with a revision of the species of the genus, and for 1887, p. 698. description of two new species. Proc. U.S. Natn. Mus., STEELR. (1969). - Handbook of Paleontology. Part 15, Ornithischia. V. 36, p. 197-332. - . - Fischer, Portland, S4 p. HALLAMA. (1969). - Tectonism and eustasy in the Jurassic. Earth- VANHINTE J. E. (1976). - A Jurassic time scale. Bull. Amer. AssOC. Sci. Rev., v. 5, p. 45-68. Petrol. Geol., v. 60, p. 489-497. HALLAMA. (1972). - Continental drift and the fossil record. Sci. VIRCHOWH. (1919). -Atlas und Epistropheus bei den Schildkroten. Amer., v. 227 (5), p. 56-66. Sitz. Ber. naturforsch. Fr. Berlin for 1919, p. 303-332. HALLAMA. (1975). - Jurassic environments. Cambridge University YOUNGC. C. (1944). - Reptilian remains from Weiyuan, Szechuan, Press. China. Bull. geol. Soc. China, v. 24, p. 187-209.

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