Theropods of the British Isles Part I

The bipedal theropods represent the most diverse group of dinosaurs including all the meat-eating ones as well as some omnivorous and herbivorous forms. Primitive theropods include the coelophysoids (small, slender and lightly built dinosaurs that thrived worldwide during the Late Triassic, and for which the best known representative is the North American Coelophysis), the ceratosaurs (including forms such as Ceratosaurus and Carnotaurus) and the tetanurans. The last group contains the vast majority of the theropods and its members are characterized among other things by a rigid tail and the total loss of the fourth and fifth digits in their hands. Megalosaurs (i.e. Megalosaurus) are early tetanurans while Spinosaurs (i.e. Spinosaurus and co) are possibly related to them. Later tetanurans are the allosauroids (large predators such as Allosaurus and Carcharodontosaurus) and the coelurosaurians, which in turn include the tyrannosauroids (such as Tyrannosaurus), the ornithomimosaurs (the ostrich-mimic forms such as Struthiomimus and Ornithomimus) and the maniraptorans. The maniraptorans with their modified wrist and generally large hands are represented by the birds (where the hands became wings) and all their closest relatives, such as the oviraptosaurs (Oviraptor and co), the deinonychosaurs (Velociraptor and co).

Fig 1.- Small coelophysoids lived during the Late Triassic in Great Britain.

Late Triassic Theropods

During the Upper Carnian (~ 220 MYA) of Scotland lived Saltopus elginensis Huene 1910, known from a poorly preserved partial skeleton including dorsal, sacral and caudal vertebrae and fragments of fore and hind limbs (BMNH R3915) found in the Lossiemouth Sandstone Formation, near Elgin, Morayshire. The exact affinities of the animal have been debated. It was either a primitive theropod or a more ancestral dinosauriform.

Theropods were definitely present during the Late Triassic period in the British Isles as proven by the discovery of fragments in the fissure fills of southern Wales. A pelvis, femur and dorsal vertebrae (BMNH PV RU P77/1 and RUP 76/1) from Pant-y-ffynon, Wales of Norian age (~210 MYA), were possibly from a coelophysoid.

The dubious ‘Zanclodon’ cambrensis Newton, 1899 of Rhaetian age (~200 MYA) is known from the mold of a large left dentary with teeth (BGS 6532/BMNH R2912) from Glamorganshire, Wales (Lilstock Fm). This one might be another coelophysoid.

Fig 2.- Larger coelophysoid such as Sarcosaurus roamed the Early Jurassic of England.

Early Jurassic Theropods

In the early Jurassic, coelophysoids are represented by the shadowy Sarcosaurus. The type species, Sarcosaurus woodi Andrews, 1921 is known from a partial pelvis, femur and vertebra (BMNH 4840/1) from Leicestershire (Lias Fm) of Early Sinemurian age (which probably is actually of earlier Late Rhaetian or Hettangian age, ~198-200 MYA). A second species, Sarcosaurus andrewsi Huene, 1932 (= Magnosaurus woodwardi) is based on a partial right tibia (BMNH R3542), originally reported by Woodward in 1908 from Warwickshire of Hettangian age (~198 MYA). Sarcosaurus was a quite large coelophysoid with an estimated length of about 3.5 m.

From the Sinemurian (~192 MYA) Upper Broadford Beds Formation of the Isle of Skye (Scotland) came an incomplete right tibia (NMS.G.1994.10.1), interpreted as belonging to a small theropod, probably another coelophysoid (Benton et al., 1995).

A partial hindlimb from Charmouth, Dorset  (BMNH 39496) that was described by Owen (1861) alongside remains of the ornithischian Scelidosaurus harrisonii was reported from the Lower Lias (Hettangian-Sinemurian). This one was found to be comparable to a megalosaur and would therefore be an early member of this group of large theropods that will dominate the Middle Jurassic period.

Finally, a tooth (BMNH 41352) from the Lias group of Lyme Regis, named ‘Megalosaurus’ lydekkeri von Huene, 1926 (= Magnosaurus lydekkeri), is from an indeterminate theropod.

Original artworks on Paleoexhibit are copyrighted to Nobu Tamura. Do not use without permission (Email: nobu dot tamura at yahoo dot com)


References:

C. W. Andrews. 1921. On some remains of a theropodous dinosaur from the Lower Lias of Barrow-on-Soar. Annals and Magazine of Natural History, series 9 8:570-576

M.J. Benton, D.M. Martill & M.A. Taylor, 1995. The first Lower Jurassic dinosaur from Scotland: limb bone of a ceratosaur theropod from Skye. Scottish Journal of Geology, 31, 177–182.

F. v. Huene. 1910. Ein primitiver Dinosaurier aus der mittleren Trias von Elgin [A primitive dinosaur from the Middle Trias of Elgin]. Geologie und Paläontologie Abhandlungen (n.s.) 8(6):317-322.

D. Naish and D. M. Martill. 2007. Dinosaurs of Great Britain and the role of the Geological Society of London in their discovery: basal Dinosauria and Saurischia. Journal of the Geological Society, London 164:493-510.

O. W. M. Rauhut and A. Hungerbühler. 2000. A review of European Triassic theropods. GAIA 15:75-88.

Sauropods of the British Isles Part III

Fig 1.- Rebbachisaurids were present on the Isle of Wight during the Early Cretaceous.

Sauropods from the Early Cretaceous (part II)

The Wessex Formation of the Isle of Wight

The Wessex formation on the Isle of Wight of Barremian age (~ 127 MYA) has a number of sauropod remains, all very fragmentary but enough to point to a high degree of diversity for this group in the British Isles during the Early Cretaceous.

The presence of Diplodocids is scarce if ever they were present at all. A chevron has been described by Alan Charig (1980) as belonging to a diplodocid, but the identification has since then been disputed.  The presence of Rebbachisaurids, on the other hand, is well attested and is represented by isolated bones, including a characteristic scapula [= shoulder blade]  (MIWG 6544), tail vertebra (MIWG 5384) and possibly teeth.  From these scant remains, it appears that the unnamed British representative of this group is most closely related to the Spanish Demandasaurus and the African Nigersaurus (see my previous post about it).

Brachiosaurids were also certainly there and remains are represented by  ‘Pleurocoelus’ valdensis Lydekker, 1889, based on teeth, dorsal and caudal vertebra found near Cuckfield, East Sussex, Hastings beds and on the Isle of Wight. Usually considered to be dubious, Pleurocoelus valdensis is, according to Ruiz-Omeñaca & Canudo (2005), a perfectly valid taxon that was also present in the Iberian Peninsula. Other possible brachiosaurid remains include  ‘Ornithopsis’ eucamerotus Hulke, 1882, based on a set of pelvis bones (BMNH R97), an unnamed taxon evidenced by a single large cervical vertebra (MIWG 7306) that might have belonged to the largest dinosaur of Europe (Naish et al., 2004), and Eucamerotus foxi Blows, 1995, described from a neural arch (BMNH R2522), two dorsals (BMNH R89-90) and another dorsal from a juvenile specimen (BMNH R2524). Oplosaurus armatus Gervais, 1852, is based on a large tooth (BMNH R964) that may belong to a brachiosaurid, but more recent analysis indicated it is more probably from a camarasaurid (Canudo et al., 2002). Chondrosteosaurus gigas Owen, 1876 is known from two neck vertebrae (BMNH 46869 & BMNH 46870) and is probably a basal titanosauriform, although we cannot say if it is a camarasaurid or a brachiosaurid.

Fig 2.- The specialized Titanosaurs were the dominant group of sauropods at the end of the Early Cretaceous.

The most evolved group of sauropods, the titanosaurs, is represented by Iuticosaurus valdensis LeLoeuff et al., 1993, known from 2 tail vertebrae (BMNH R146a & BMNH 151).

Finally, two dubious taxa are indeterminate sauropods: the Isle of Wight ‘Ornithopsis’ hulkei dorsal vertebra (BMNH R28632) was renamed Bothriospondylus elongatus by Owen in 1875 and ‘Ornithopsis’ eucamerotus by Hulke in 1882. Chondrosteosaurus magnus Owen, 1876 is based on a single partial vertebra (BMNH R98).

The Lower and Upper Greensand

The Lower Greensand Group of Aptian age (~120 MYA) delivered a pelvis and associated sacrum (BMNH R12713) from Luccombe Chine, Isle of Wight of a titanosauriform of some sort (Stroh, 1949, Blows, 1995). Also from the Lower Greensand came Dinodocus mackesoni Owen, 1884, based on a humerus (BMNH 14695) from Hythe, Kent. Possibly another indeterminate titanosauriform.

The Upper Greensand formation (Albian-Cenomanian age, ~112 MYA) in southeastern England bears a few sauropod remains, which are the youngest in the British Isles and all belong to titanosaurs. “Titanosaurus” lydekkeri Huene, 1929 (= Iuticosaurus lydekkeri), based on a vertebra (BMNH 32390) found on the Isle of Wight, was synonymized with Macrurosaurus semnus by McIntosh, 1990, but generally considered a nomen dubium.  Macrurosaurus semnus Seeley, 1876 is based on a set of 25 caudal vertebrae (SM B55630) and 15 more various bits found in Cambridgeshire and of Cenomanian age, and is an indeterminate titanosaur.

This concludes our tour of the sauropods of the British Isles.


Original artworks on Paleoexhibit are copyrighted to Nobu Tamura. Do not use without permission (Email: nobu dot tamura at yahoo dot com)


References:


W. T. Blows. 1995. The Early Cretaceous brachiosaurid dinosaurs Ornithopsis and Eucamerotus from the Isle of Wight, England. Palaeontology 38(1):187-197


W. T. Blows. 1998. A review of Lower and Middle Cretaceous dinosaurs of England. In S. G. Lucas, J. I. Kirkland, and J. W. Estep (eds.), Lower and Middle Cretaceous Terrestrial Ecosystems, New Mexico Museum of Natural History and Science Bulletin 14:29-38


J. Le Loeuff, E. Buffetaut, M. Martin, V. Martin, and H. Tong. 1993. Découverte d'Hadrosauridae (Dinosauria, Ornithischia) dans le Maastrichtian des Corbières (Aude, France) [Discovery of Hadrosauridae (Dinosauria, Ornithischia) in the Maastrichtian of Corbières (Aude, France)]. Comptes Rendus de l'Academie des Sciences, Paris, Série II 316:1023-1029


R. Lydekker. 1889. Note on some points in the nomenclature of fossil reptiles and amphibians, with preliminary notices of two new species. Geological Magazine, decade 3 6:325-326


R. Lydekker. 1893. On a sauropodous dinosaurian vertebra from the Wealden of Hastings. Quarterly Journal of the Geological Society of London 49:276-280


J. S. McIntosh. 1990. Sauropoda. In D. B. Weishampel, H. Osmólska, and P. Dodson (eds.), The Dinosauria. University of California Press, Berkeley 345-401


A. G. Melville. 1849. Notes on the vertebral column of the Iguanodon. Philosophical Transactions of the Royal Society of London 139:285-300.


D. Naish, D.M. Martill, D. Cooper & K.A. Stevens, 2004. Europe’s largest dinosaur? A giant brachiosaurid cervical vertebra from the Wessex Formation (Early Cretaceous) of southern England. Cretaceous Research, 25, 787–795.


D. Naish and D. M. Martill. 2007. Dinosaurs of Great Britain and the role of the Geological Society of London in their discovery: basal Dinosauria and Saurischia. Journal of the Geological Society, London 164:493-510


R. Owen. 1876. Monograph on the fossil Reptilia of the Wealden and Purbeck formations. Supplement no. VII. Crocodilia (Poikilopleuron) and Dinosauria? (Chondrosteosaurus). [Wealden.]. The Palaeontographical Society, London 1876:1-7


J. I. Ruiz-Omeñaca and J. I. Canudo. 2005. "Pleurocoelus" valdensis Lydekker 1889 (Saurischia, Sauropoda) en el Cretácico Inferior (Barremiense) de la Península Ibérica ["Pleurocoelus" valdensis Lydekker 1889 (Saurischia, Sauropoda) in the Lower Cretaceous (Barremian) of the Iberian Peninsula]. Geogaceta 38:43-45


H. G. Seeley. 1870. On Ornithopsis, a gigantic animal of the pterodactyle kind from the Wealden. Annals and Magazine of Natural History, series 4 5:279-283.


H. G. Seeley. 1876. On Macrurosaurus semnus (Seeley), a long tailed animal with procoelous vertebrae from the Cambridge Upper Greensand, preserved in the Woodwardian Museum of the University of Cambridge. Quarterly Journal of the Geological Society of London 32:440-444


M. P. Taylor and D. Naish. 2007. An unusual new neosauropod dinosaur from the Lower Cretaceous Hastings Beds Group of East Sussex, England. Palaeontology 50(6):1547-1564.

Sauropods of the British Isles Part II

Fig 1.- A hypothetical reconstruction of Pelorosaurus conybeari.

Sauropods from the Early Cretaceous (I)

The Hastings Beds

The Hastings Beds in East Sussex, of Berriasian-Valanginian age (~140 MYA), bore a number of fragmentary sauropod remains. A set of tail vertebrae and chevrons (BMNH R2544–2555) found near Cuckfield, East Sussex, were originally described alongside some iguanodont remains by Richard Owen as Cetiosaurus brevis (Owen, 1842). Alexander Melville, noting Owen’s mistake, renamed the sauropod vertebrae Cetiosaurus conybeari (Melville, 1849). A year later, Gideon Mantell realizing that they belong to a animal quite distinct from Cetiosaurus, changed the name into Pelorosaurus conybeari (Mantell, 1850), and added to the description, a humerus (BMNH 28626) found a few meters away from the original vertebrae material. This succession of attribution changes resulted in a taxonomical nightmare for the later generation scientists. Technically, the name C. brevis has indeed seniority over Pelorosaurus conybeari and should be considered to be the valid name. However, with the invalidation of C. medius (see Part I), C. brevis would also be the type species of the genus Cetiosaurus, making its use for the Middle Jurassic C. oxoniensis, which turned out to be a very different animal quite problematic (As far as I know, the petition to ICZN to make C. oxoniensis the type species of Cetiosaurus is still pending) . As for the general aspect of what Pelorosaurus may have looked like, all that can be said from the scant remains is that it was a brachiosaurid and would probably resemble to a smaller version of the North American Late Jurassic Brachiosaurus, with a possible size of some 15 meters in length.

Not much can be said about the three other named sauropods of the Hastings Beds. ‘Pelorosaurus’ becklesi Mantell, 1852 (= Morosaurus becklesii Marsh, 1889) based on a humerus (BMNH R1868), ulna, radius and skin impressions, probably belong to a different animal than Pelorosaurus conybeari. It may also be a brachiosaurid unless it is a more advanced titanosaur. ‘Ornithopsis’ hulkei Seeley, 1870 is based on two dorsal vertebrae, one from East Sussex (BMNH R2239), the other from the Isle of Wight Wessex Formation (BMNH R28632), and originally thought to belong to a pterosaur (thus the genus name which means “bird likeness”). Owen (1876), however, split the two findings, naming the East Sussex vertebrae Bothriospondylus magnus, then Chondrosteosaurus magnus. The remains have no distinct characteristics apart the fact that they belong to a sauropod of some sort so the name should be considered dubious.  Xenoposeidon proneneukos Taylor & Naish, 2007 is based on a single partial back vertebra (BMNH R2095). Xenoposeidon’s vertebra is so unique that its affinities within the Sauropods are quite uncertain.


Next will be the sauropods from the Wessex Formation.

 

 

Original artworks on Paleoexhibit are copyrighted to Nobu Tamura. Do not use without permission (Email: nobu dot tamura at yahoo dot com)

References:

G. A. Mantell. 1850. On the Pelorosaurus; an undescribed gigantic terrestrial reptile, whose remains are associated with those of Iguanodon and other saurians in the strata of the Tilgate Forest, in Sussex. Philosophical Transactions of the Royal Society of London 140(16):379-390.

A. G. Melville. 1849. Notes on the vertebral column of the Iguanodon. Philosophical Transactions of the Royal Society of London 139:285-300.

D. Naish and D. M. Martill. 2007. Dinosaurs of Great Britain and the role of the Geological Society of London in their discovery: basal Dinosauria and Saurischia. Journal of the Geological Society, London 164:493-510.

R. Owen. 1876. Monograph on the fossil Reptilia of the Wealden and Purbeck formations. Supplement no. VII. Crocodilia (Poikilopleuron) and Dinosauria? (Chondrosteosaurus). [Wealden.]. The Palaeontographical Society, London 1876:1-7.

H. G. Seeley. 1870. On Ornithopsis, a gigantic animal of the pterodactyle kind from the Wealden. Annals and Magazine of Natural History, series 4 5:279-283.

M. P. Taylor and D. Naish. 2007. An unusual new neosauropod dinosaur from the Lower Cretaceous Hastings Beds Group of East Sussex, England. Palaeontology 50(6):1547-1564.