smith, 2006

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Rhomaleosaurid plesiosaurs, a group of Mesozoic aquatic reptiles, are known from a number of well preserved skeletons. The research outlined here seeks to update our understanding of pliosaur cranial anatomy, assess the status of the genus Rhomaleosaurus , and test the monophyly of the Rhomaleosauridae. Lower Jurassic pliosaurs may be subdivided by stage. Hettangian fauna - Rhomaleosaurus megacephalus (Figs 1A, B, C and 3D, E), Eurycleidus arcuatus (Fig 3C), Macroplata tenuiceps, and Archaeonectrus rostratus. An additional taxon is recognised by a unique symphysis morphology (Fig 3F). A new reconstruction of the skull of R. megacephalus is presented here (Fig 1). Casts of the holotype, the original of which was destroyed in the Bristol Museum during an air raid, are also providing vital information. Toarcian fauna - Rhomaleosaurus cramptoni (Figs 2A, B), R. zetlandicus (Fig 3B), R. propinquus, R. thorntoni (Fig 3A), R. longirostris, and R. victor (background image). The holotype of the genus (R. cramptoni) (Fig 2A) is currently under preparation and a thorough interpretation of the whole skull including the palate will follow. There is a considerable diversity in skull anatomy between these forms (compare Fig 1B with 2B), and variation in symphysis morphology is equally great (Fig 3). Indeed, A review of pectoral girdle anatomy also reveals striking variation between species pertaining to Rhomaleosaurus. (Fig 4) Cranial Anatomy and Systematics of Lower Jurassic Pliosaurs Adam Stuart Smith School of Biology and Environmental Science, University College Dublin, Belfield, Dublin 4, Ireland. <[email protected]> <www.plesiosauria.com> Acknowledgements Many thanks to Sandra Chapman, Arthur Cruickshank, Gareth Dyke, Mark Evans, Richard Forrest, Leona Leonard, Nigel Monaghan, Camilla Nichol, Leslie Noé, Roger Osborne, Jon Radley, Remmert Schouten, Julia Sigwart, Paolo Viscardi, Matt Williams, Patrick Wyse-Jackson, who have all contributed valuably to this ongoing project, and this poster. Portions of this research were made possible by a grant from Palaeontographical Society. Preliminary morphometric analyses based on skull proportions (Fig 5) and a preliminary cladistic analysis (Fig 6) show morphological distinctions, notably between Toarcian and Hettangian taxa Above Fig 1. Rhomaleosaurus megacephalus (NMING F10194). A dorsal view of Skull. B. Interpretation of A. C. Interpretation of palate, with annotations indicating characters at variance with Cruickshank (1994). Left Fig 2. Rhomaleosaurus cramptoni (NMING F8785). A. Dorsal view of skull. B. preliminary Interpretation of A. with annotations indicating major characters shared with other British Toarcian pliosaurs, but differing from R. megacephalus. Characters shared with R. megacephalus: postfrontal ridge and lack of nasals Fig. 4. Reconstructed pectoral girdles of A. R. thorntoni (BMNH R4853) and B. one half of R. megacephalus (composite of left and right elements of NMING F10194). The girdle of R. thorntoni is much larger, broader and less elongate than R. megacephalus, the latter also has a clear ‘V’ shaped posterior median coracoid embayment. 2 2.2 2.4 2.6 2.8 3 4 4.5 5 5.5 6 6.5 7 Length/ orbit length Length/ premax length R. megacephlalus (dublin) macrocephalus (cast) F8780 R. propinquus R. zetlandicus R. cramptoni F8785 Eurycleidus arcuatus F8749 Hettangian Toarcian References Cruickshank, A. R. I. 1994. Cranial anatomy of the Lower Jurassic pliosaur Rhomaleosaurus megacephalus (Stutchbury)(Reptilia: Plesiosauria). Philosophical Transactions of the Royal Society of London, Series B, 343, 247-260. Clear lateral palatine vacuity Vomers extend far posterior of internal nares Jugal exposed and contacts suborbital fenestra Large ectopterygoid 30CM 30CM Short premaxillary rostrum Large dorso-median foramen between nares Frontals separated on midline Borders of Premaxillae run parallel Prefrontal contacts postfrontal 30CM Conclusions R. propinquus, R. zetlandicus, R. thorntoni and R. cramptoni. are united via cladistic analysis and may be regarded as Rhomaleosaurus sensu stricto. Their interrelationships await the ongoing preparation of R. cramptoni. R. megacephalus and R. victor differ significantly from Rhomaleosaurus sensu stricto and from each other (Fig 6). Lower Jurassic pliosaurs form a sister relationship with the Pliosauridae, however, a traditional Rhomaleosauridae is not supported due to the exclusion of R. victor. The genus name ‘Thaumatosaurus’ is often applied to the holotype of ‘R’. victor and could be resurrected? ‘R’. megacephalus Pliosauridae Rhomaleosaurus sensu stricto A traditional monophyletc Rhomaleosauridae including ‘R’. victor is not supported. Fig. 3. Mandibular symphyses of various species currently pertaining to the genus Rhomaleosaurus, and the species Eurycleidus arcuatus. The width of each symphysis has been standardised to allow direct comparison between species (so not to scale). The top line represents the minimum symphysis length (R.thorntoni) and the bottom line represents the maximum (WARWKS G10875). Areas shaded black help visualise variation. Light grey shading = inferred missing bone. 20CM Fig. 5. See text. Fig. 6. Majority Rule consensus tree (heuristic search, 31 characters) A A B B A C B A B C D E F

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Fig. 6. Majority Rule consensus tree (heuristic search, 31 characters) Fig. 4. Reconstructed pectoral girdles of A. R. thorntoni (BMNH R4853) and B. one half of R. megacephalus (composite of left and right elements of NMING F10194). The girdle of R. thorntoni is much larger, broader and less elongate than R. megacephalus, the latter also has a clear ‘V’ shaped posterior median coracoid embayment. 4 4.5 5 5.5 6 6.5 7 Fig. 5. See text. Large dorso-median foramen between nares A 20CM 30CM 2.2

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Page 1: Smith, 2006

Rhomaleosaurid plesiosaurs, a group of Mesozoic aquaticreptiles, are known from a number of well preserved skeletons.The research outlined here seeks to update our understandingof pliosaur cranial anatomy, assess the status of the genusRhomaleosaurus, and test the monophyly of theRhomaleosauridae. Lower Jurassic pliosaurs may besubdivided by stage.

Hettangian fauna - Rhomaleosaurus megacephalus (Figs 1A,B, C and 3D, E), Eurycleidus arcuatus (Fig 3C), Macroplatatenuiceps, and Archaeonectrus rostratus. An additional taxon isrecognised by a unique symphysis morphology (Fig 3F). A newreconstruction of the skull of R. megacephalus is presented here(Fig 1). Casts of the holotype, the original of which was destroyedin the Bristol Museum during an air raid, are also providing vitalinformation.

Toarcian fauna - Rhomaleosaurus cramptoni (Figs 2A, B), R.zetlandicus (Fig 3B), R. propinquus, R. thorntoni (Fig 3A), R.longirostris, and R. victor (background image). The holotype ofthe genus (R. cramptoni) (Fig 2A) is currently under preparationand a thorough interpretation of the whole skull including thepalate will follow.

There is a considerable diversity in skull anatomy between theseforms (compare Fig 1B with 2B), and variation in symphysismorphology is equally great (Fig 3). Indeed, A review of pectoralgirdle anatomy also reveals striking variation between speciespertaining to Rhomaleosaurus. (Fig 4)

Cranial Anatomy and Systematics of Lower Jurassic Pliosaurs

Adam Stuart SmithSchool of Biology and Environmental Science, University College Dublin,

Belfield, Dublin 4, Ireland. <[email protected]> <www.plesiosauria.com>

AcknowledgementsMany thanks to Sandra Chapman, Arthur Cruickshank, Gareth Dyke, Mark Evans, Richard Forrest, Leona Leonard, Nigel Monaghan, Camilla Nichol, Leslie Noé,Roger Osborne, Jon Radley, Remmert Schouten, Julia Sigwart, Paolo Viscardi, Matt Williams, Patrick Wyse-Jackson, who have all contributed valuably to this ongoingproject, and this poster. Portions of this research were made possible by a grant from Palaeontographical Society.

Preliminary morphometricanalyses based on skullproportions (Fig 5) and apreliminary cladistic analysis(Fig 6) show morphologicaldistinctions, notably betweenToarcian and Hettangian taxa

Above Fig 1.Rhomaleosaurusmegacephalus (NMINGF10194). A dorsal view ofSkull. B. Interpretation of A.C. Interpretation of palate,with annotations indicatingcharacters at variance withCruickshank (1994).

Left Fig 2. Rhomaleosauruscramptoni (NMING F8785).A. Dorsal view of skull. B.preliminary Interpretation ofA. with annotations indicatingmajor characters shared withother British Toarcianpliosaurs, but differing fromR. megacephalus.Characters shared with R.megacephalus: postfrontalridge and lack of nasals

Fig. 4. Reconstructed pectoral girdles of A. R. thorntoni (BMNH R4853)and B. one half of R. megacephalus (composite of left and right elementsof NMING F10194). The girdle of R. thorntoni is much larger, broader andless elongate than R. megacephalus, the latter also has a clear ‘V’ shapedposterior median coracoid embayment.

2

2.2

2.4

2.6

2.8

3

4 4.5 5 5.5 6 6.5 7

Length/ orbit length

Len

gth

/ p

rem

ax l

en

gth

R. megacephlalus (dublin)macrocephalus (cast) F8780R. propinquusR. zetlandicusR. cramptoni F8785Eurycleidus arcuatus F8749

Hettangian

Toarcian

ReferencesCruickshank, A. R. I. 1994. Cranial anatomy of the Lower Jurassicpliosaur Rhomaleosaurus megacephalus (Stutchbury)(Reptilia:Plesiosauria). Philosophical Transactions of the Royal Society of London,Series B, 343, 247-260.

Clear lateralpalatine vacuity

Vomers extend far posterior of internal nares

Jugal exposed and contacts suborbital fenestra

Largeectopterygoid

30CM

30CM

Short premaxillaryrostrum

Large dorso-medianforamen betweennares

Frontalsseparated on midline

Borders of Premaxillaerun parallel

Prefrontalcontactspostfrontal

30CM

ConclusionsR. propinquus, R. zetlandicus, R.thorntoni and R. cramptoni. are unitedvia cladistic analysis and may beregarded as Rhomaleosaurus sensustricto. Their interrelationships awaitthe ongoing preparation of R.cramptoni.

R. megacephalus and R. victor differsignificantly from Rhomaleosaurussensu stricto and from each other (Fig6). Lower Jurassic pliosaurs form asister relationship with thePliosauridae, however, a traditionalRhomaleosauridae is not supporteddue to the exclusion of R. victor. Thegenus name ‘Thaumatosaurus’ is oftenapplied to the holotype of ‘R’. victorand could be resurrected?

‘R’. megacephalus

Pliosauridae

Rhomaleosaurus sensu stricto

A traditionalmonophyletcRhomaleosauridaeincluding ‘R’. victoris not supported.

Fig. 3. Mandibular symphyses of various species currently pertaining tothe genus Rhomaleosaurus, and the species Eurycleidus arcuatus. Thewidth of each symphysis has been standardised to allow directcomparison between species (so not to scale). The top line represents theminimum symphysis length (R.thorntoni) and the bottom line representsthe maximum (WARWKS G10875). Areas shaded black help visualisevariation. Light grey shading = inferred missing bone.

20CM

Fig. 5. See text.

Fig. 6. MajorityRule consensustree (heuristicsearch, 31characters)

A

A B

BA

CB

A B C D E F