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20 November 2009 A New Large-Bodied Theropod Dinosaur from the Middle Jurassic of Warwickshire, United Kingdom
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Abstract

Previously undocumented postcranial material from the Chipping Norton Limestone Formation (Middle Jurassic: Lower Bathonian) of Cross Hands Quarry, near Little Compton, Warwickshire represents a new large-bodied theropod dinosaur, distinct from the contemporaneous Megalosaurus bucklandii. Cruxicheiros newmanorum gen. et sp. nov. is diagnosed by a single autapomorphy, the presence of a proximomedially inclined ridge within the groove that marks the lateral extent of the posterior flange of the femoral caput (trochanteric fossa). C. newmanorum shows three tetanuran features: widely separated cervical zygapophyses, a swollen ridge on the lateral surface of the iliac blade and an anterior spur of the caudal neural spines. However, due to fragmentary preservation its affinities within Tetanurae remain uncertain: phylogenetic analysis places it as the most basal tetanuran, the most basal megalosauroid (= spinosauroid) or the most basal neotetanuran.

Introduction

The recognition of dinosaur fossils as the remains of gigantic fossil reptiles was based on theropod bones from the shallow- to marginal-marine Middle Jurassic Bathonian Stage of the United Kingdom (Parkinson 1822; Buckland 1824). Prior to this, theropod and other dinosaur material was recovered from these strata but remained unrecognised (Plot 1677: pl. 8: 4; Lluyd 1699: pl. 16: 1328; Woodward 1728; Brookes 1763: vol. 5, fig. 317; Delair and Sarjeant 1975, 2002). Throughout a long history of collection, most theropod remains of this provenance have been referred uncritically to either Megalosaurus (large-bodied individuals, represented by the majority of specimens) or Iliosuchus (small-bodied individuals) (Owen 1842; Phillips 1871; von Huene 1926; Steel 1970; Benton and Spencer 1995).

However, recent work (Benson 2009, 2010) has clarified the status of the core of this material, mostly collected from the Taynton Limestone Formation (Middle Bathonian) of Stonesfield, Oxfordshire, the type locality of Megalosaurus and Iliosuchus. This work established that only one large-bodied theropod taxon was present in the most productive UK Bathonian localities: Stonesfield (above) and New Park Quarry (Chipping Norton Limestone Formation, Lower Bathonian), Gloucestershire (Benson 2009). This led to the identification of unique diagnostic features (autapomorphies) of Megalosaurus and the recognition of additional Megalosaurus remains in the Chipping Norton Limestone Formation (Lower Bathonian) of Oakham Quarry, Sarsgrove and Workhouse quarries, Oxfordshire (Benson 2010). We also now recognise that no material outside of the Lower-Middle Bathonian formations of the United Kingdom can currently be referred to Megalosaurus.

The present contribution documents a collection of theropod remains from the Chipping Norton Limestone Formation (Lower Bathonian) of Cross Hands Quarry near Little Compton, Warwickshire, UK that can be referred to a new taxon of large-bodied theropod dinosaur. The stratigraphy and palaeontology of this site, part of which is preserved as a Site of Special Scientific Interest (protected status awarded by the UK government), was recently documented by Sumbler (2002). The theropod material was previously reported by Sumbler (2002) but until now had not been studied in any detail.

The Middle Jurassic saw the early history of Tetanurae, a taxonomically and ecologically diverse theropod clade (Holtz et al. 2004). Unfortunately however, Middle Jurassic dinosaur faunas are still poorly understood. Middle Jurassic theropod fossils are rare and determinate remains have only been reported from Europe, Argentina and China (Weishampel et al. 2004). Most Middle Jurassic European theropod taxa are megalosaurids (sensu Benson 2010), although Proceratosaurus (a tyrannosauroid coelurosaur; Rauhut and Milner 2008; Rauhut et al. 2010) and Poekilopleuron (a possible allosauroid; Benson 2010) are exceptions. Benson (2010) reported geographically localised distributions for Middle Jurassic theropod clades and new specimens can add detail to this biogeographic hypothesis. As such, the Cross Hands Quarry theropod remains represent an important data point for studies of dinosaur biogeography, phylogeny and faunal composition. The remains are also significant as they cast doubt on uncritical referrals of indeterminate theropod remains from the Bathonian of the United Kingdom to Megalosaurus, primarily abundant isolated teeth and bones (Benton and Spencer 1995).

Based on correspondence held at the Birmingham Museum and Art Gallery (BIRMG) and recollections of one of the present landowners (Mr. Stephen Newman) the new remains were collected from Cross Hands Quarry in the early 1960s. They were brought to the BIRMG for preparation and some formed part of a temporary display that was installed in the early 1990s. The material was recently (2008) transferred to the Warwickshire Museum Service (WARMS), prompting its study.

Institutional abbreviations.—BIRMG, Birmingham Museum and Art Gallery, Birmingham, UK; OUMNH, Oxford University Museum of Natural History, Oxford, UK; UC OBA, Department of Organismal Biology, University of Chicago, Chicago, USA; WARMS, Warwickshire Museum Service, Warwick, UK.

Systematic palaeontology

Dinosauria Owen, 1842
Theropoda Marsh, 1881
Tetanurae Gauthier, 1986
Tetanurae incertae sedis Genus Cruxicheiros nov.

  • Type species: Cruxicheiros newmanorum gen. et sp. nov.

  • Etymology: From Latin crux, cross; and Greek cheiros, hand; intended as “cross hand”, a version of the locality name.

  • Diagnosis.—As for the type and only species.

Cruxicheiros newmanorum sp. nov.
Figs. 13.

  • Etymology: After the Newman family, owners of Cross Hands Quarry, Warwickshire, United Kingdom.

  • Type material: Holotype: WARMS G15770, a partial right femur. Paratypes: WARMS G15771, additional theropod material from the type locality that probably represents the same individual as the holotype: an anterior dorsal or posterior cervical vertebra; a dorsal neural arch; a partial dorsal vertebra; the anterior half of a middle-distal caudal vertebra; a partial right scapulocoracoid; a partial left ilium; the proximal end of a left pubis; numerous rib and bone fragments.

  • Type locality: Chipping Norton Limestone Formation (Lower Bathonian Zigzagiceras zigzag Biozone; Torrens 1980) of Cross Hands Quarry (National Grid Reference [NGR] SP 269 291) near Little Compton, Warwickshire.

  • Type horizon: The Chipping Norton Limestone Formation of the Little Compton area is characterised by sandy ooidal and bioclastic limestones, yielding a marine bivalve fauna (Sellwood and McKerrow 1974; Sumbler 2002). Abundant terrestrial plant material (Sumbler 2002) indicates significant terrigenous input into a shallow marine environment and a potentially appropriate mechanism for derivation of the dinosaur remains.

  • Diagnosis.—Tetanuran theropod with autapomorphic proximomedially inclined ridge within the trochanteric fossa of the femur. Differs from the contemporaneous Megalosaurus bucklandii in posessing low proportions of the dorsal neural spines, transversely broader dorsal neural spines, a prominent posterior flange of the femoral caput and a lower ratio of anteroposterior length to mediolateral width of the pubic peduncle (1.60 in Megalosaurus and estimated between 1.00 and 1.10 in C. newmanorum). Few informative comparisons can be made between C. newmanorum and Dubreuillosaurus or Poekilopleuron from the lower Bathonian of France because of limited overlap between known material. However, Dubreuillosaurus shows two large internal chambers separated by a midline septum in pneumatic vertebrae and an unfused scapula and coracoid (perhaps due to immaturity of the holotype) (Allain 2005) whereas C. newmanorum vertebrae have more numerous internal pneumatic chambers and the scapulocoracoid suture is fused and swollen. Poekilopleuron also differs from C. newmanorum as it lacks prominent anterior spur on the caudal neural spines.

  • Remarks.—Collections records for the specimens are scant and there is no formal record of their association such as a quarry map. However, the relative sizes and the absence of repeated elements are consistent with the hypothesis that they represent a single individual. Additionally, the remains were collected over a short time period of one or maybe two field seasons (BIRMG, unpublished collections records), and all preserve an identical matrix of sandy bioclastic ooidal limestone as well as residual patches of re-deposited stalagmitic calcite. They therefore probably correspond to a point locality. Finally, diagnostic remains of large-bodied theropods other than Megalosaurus are rarely preserved in British Bathonian deposits (Benson 2010), but most of the Cross Hands Quarry specimens are demonstrably different from Megalosaurus. If the material does represent multiple individuals then it is very unlikely that it represents more than one of these rare non-Megalosaurus taxa.

    The following specimens from Cross Hands Quarry are listed in the accessions register at BIRMG: metacarpal; coracoid; 3 ribs; fibula; vertebra; 2 vertebrae; left ilium, left femur fragment; lower femur fragment; and pubic bone fragment (BIRMG, unpublished collection records). The listed metacarpal and fibula are unaccounted for and may be lost or have originally been misidentified. Additionally, four vertebrae are present whereas only three were listed. This leaves the possibility that one of the vertebrae described here was not recovered from Cross Hands Quarry but as there is no way to identify which this is, they are all described.

    An additional specimen, a fragmentary right tibia (OUMNH J.29831), was collected from Cross Hands Quarry by Mr. P. Stewart and donated to the OUMNH in 1980. This specimen is very poorly preserved and its affinities within Theropoda cannot be determined. It is not referred to Cruxicheiros newmanorum herein as the details of its collection, from where and which horizon in the quarry it was collected, cannot be established at present. The specimen should be considered as Theropoda indet.

  • Stratigraphic and geographic range.—Only known from a single locality in the Lower Bathonian of Warwickshire, United Kingdom.

  • Description

    Vertebrae.—A fragmentary posterior cervical or anterior dorsal vertebra is preserved as a transverse slice from close to the posterior end of the bone (Fig. 1A). This reveals the neural canal and several large internal chambers (camerae) in the centrum. These camerae are probably pneumatic in origin, arising from invasion of the bone by pneumatic diverticulae via large pneumatic foramina (“pleurocoels”) in the lateral surfaces of the centrum, as is common in theropods (Britt 1993). They are similar to the confirmed pneumatic camerae of Sinraptor (Britt 1993: fig. 21.1–3, 22.11) but are smaller and more numerous than those of megalosauroids (= spinosauroids) such as Marshosaurus (Britt 1993: fig 19.3–4) and Torvosaurus (Britt 1991, 1993), in which two large, main chambers are usually divided by a midline septum. Vertebrae with thin external walls and very large numbers of small internal chambers are termed “camellate”. These are known in neoceratosaurs and carcharodontosaurian allosauroids among basal theropods (Britt 1993), which therefore differ markedly from C. newmanorum.

    A dorsal centrum and transverse process (Fig. 1B) is mostly concealed by attached matrix. The centrum lacks internal pneumatic cavities, indicating that pneumatic foramina (= pleurocoels) were absent. A prominent anterior centrodiapophyseal lamina is visible. A dorsal neural spine and left transverse process (Fig. 1D) also have a large block of attached matrix in the space between them. The transverse process is oriented dorsolaterally, and the combined anterior and posterior centrodiapophyseal laminae form a stout longitudinal ridge along its ventral surface. Some matrix is still attached to the bone surface adjacent to this ridge anteriorly and posteriorly, giving the impression (Fig. 1D1) that large foramina are present. However, they are absent. The neural spine is abraded posteriorly, but a thick, rugose, ligament ossification is present on the anterior surface, as in various basal tetanurans, including Marshosaurus (DMNH 3718) and the allosauroids Aerosteon (Sereno et al. 2008), Allosaurus (Madsen 1976), Neovenator (Brusatte et al. 2008) and Tyrannosaurus (Brochu 2002). Less prominent ligament ossifications are present in other large theropods such as Torvosaurus (Britt 1991). The distal end of the dorsal neural spine of WARMS G15771 is abraded but expands transversely (Fig. 1D4), indicating that the distal end is almost complete. In the British Jurassic theropods Megalosaurus (Benson 2010) and Metriacanthosaurus (OUMNH J. 12144), and in Ceratosaurus (Madsen and Welles 2000) and Sinraptor (Currie and Zhao 1994), the neural spine is over 1.5 times the height of the centrum and substantially longer than the transverse processes. However, in Cruxicheiros the neural spine is approximately the same length as the transverse process, indicating relatively low proportions as in various other theropods including Allosaurus (Madsen 1976), Marshosaurus (DMNH 3718), Torvosaurus (Britt 1991) and Piatnitzkysaurus (Bonaparte 1986).

    The anterior half of a middle-distal caudal vertebra (WARMS G15771) has a shallowly concave anterior articular surface and short prezygapophyses. A lump of limestone matrix adheres to the dorsal surface of the neural arch anteriorly. On the posterior surface of this lump, a section through the low anterior portion of the neural spine is preserved which indicates that an anterior spur of the neural spine was present. This feature is a tetanuran synapomorphy (Rauhut 2003).

  • Pectoral girdle.—A right scapulocoracoid fragment (Fig. 2A) preserves portions of the fused scapula and coracoid. The area of fusion is swollen and heavily ossified, as in Megalosaurus (Benson 2010), and the glenoid faces posterolaterally. The ventral margin of the coracoid foramen is preserved, forming a suboval opening on both the medial and lateral surfaces of the bone, as it does in most other theropods. The specimen is highly abraded and it is not possible to determine the morphology of the coracoid tubercle or posteroventral process, although this process is clearly present. Fragments of the scapula are also preserved (Fig. 2B). These are not adequate to assess the proportions of the blade. However, they do indicate that it was mediolaterally narrow with a cross-section that tapers dorsally and ventrally (Fig. 2C2) as in other theropods.

  • Pelvic girdle.—The left ilium is 477 mm long as preserved, missing the anterior blade, posterior and medial blades, supracetabular crest, and ischial peduncle (Fig. 3A). The pubic peduncle and periphery of the iliac blade were damaged during preparation, so the bone surface is stripped away around the margins of the bone. A low, swollen median ridge is present on the lateral surface of the blade. This ridge is a tetanuran synapomorphy that is widely-distributed among basal members of the clade (Bonaparte 1986; Benson 2009; Zhao et al. 2010). Bone surface preservation is not sufficient to determine whether associated nutrient foramina were present in C. newmanorum. The supracetabular crest is damaged but seems to project ventrolaterally as a shelf, as in tetanurans other than Chuandongocoelurus and Monolophosaurus (Zhao et al. 2010). As preserved, the pubic peduncle is slightly longer anteroposteriorly (95 mm) than its maximum mediolateral width (90 mm). However, medial and posterior portions are broken off and it may have been as wide as long, or only slightly longer than wide. In Megalosaurus and most other tetanurans the pubic peduncle is at least 1.3 times as long anteroposteriorly as it is wide mediolaterally, but in the megalosaurid Eustreptospondylus (OUMNH J. 13558, J.29774; Sadleir et al. 2008) and the allosauroid Fukuiraptor (Azuma and Currie 2000) it is approximately as broad as long, resembling the condition in non-tetanurans. This may also have been the case in C. newmanorum.

    The proximal end of the left pubis is abraded. The smooth anterior margin of the obturator “foramen” is preserved, but preservation is not good enough to determine if it was open as a notch or closed posteriorly to form a foramen.

  • Hindlimb.—A right femur is preserved in three pieces. One piece comprises the head and proximal part of the shaft (Fig. 3C), another comprises a portion of the shaft including the fourth trochanter (Fig. 3D), and the third comprises the abraded distal end (Fig. 3E). The proximal and posterior surfaces of the head and posterior surface of the caput are well-preserved. The head measures 185 mm mediolaterally and 95 mm anteroposteriorly at the caput. The greater trochanter is narrower than the caput, so the outline of the head in proximal view narrows from medial to lateral as in non-coelurosaurian theropods (Hutchinson 2001). The proximal surface of the head bears a shallow, anteromedially oriented groove, the proximal articular groove, which is present in non-neotetanuran theropods and many non-theropods (Hutchinson 2001). The posterior flange of the caput is prominent and extends further posteriorly than the posterior surface of the head (Fig. 3C4), as in most basal theropods. However, this is unlike the low posterior flange of most megalosaurids, including Afrovenator (UC OBA 1) and Megalosaurus (Benson 2010). The trochanteric fossa, the groove bounding the posterior flange laterally on the posterior surface of the femur, is anteroposteriorly broad. Within this fossa, nearly parallel to the posterior flange, is a prominent ridge. This ridge is not present in any other basal theropod femur (e.g., Madsen 1976; Bonaparte 1986; Holtz et al. 2004; Sadleir et al. 2008) including the Bathonian taxa Dubreuillosaurus (MNHN 1998–13) and Megalosaurus (Benson 2010), and is considered here as an autapomorphy of C. newmanorum. The base of the lesser trochanter is preserved along with the lateral part of the anterior surface of the greater trochanter region. However, as it is very incomplete it is not possible to determine the morphology of the lesser trochanter.

Fig. 1.

Tetanuran theropod Cruxicheiros newmanorum gen. et sp. nov. axial vertebrae (WARMS 15771) from the Chipping Norton Limestone Formation, Bathonian of the United Kingdom. A. Posterior cervical or anterior dorsal vertebra in posterior view. B. Partial middle-posterior dorsal vertebra in right lateral view showing a sagittal cross-section (B1) and in dorsal view (B2). C. Middle-distal caudal vertebra in left lateral (C1) and posterior (C2) views. D. Middle-posterior dorsal neural arch in anterior (D1), right lateral (D2, D3), and posterior (D4) views. Photographs (A, B1, B2, C1, C2, D1, D2, D4) and line drawing (D3). Crossed-hatching indicates matrix and grey tone indicates broken bone.

f01_35.eps

Fig. 2.

Tetanuran theropod Cruxicheiros newmanorum gen. et sp. nov. right scapulocoracoid (WARMS 15771) from the Chipping Norton Limestone Formation, Bathonian of the United Kingdom. A. Scapulocoracoid in lateral (A1) and ventral (A2) views. B, C. Scapular fragments in medial or lateral views (B, C1) and in cross-section (C2).

f02_35.eps

Fig. 3.

Tetanuran theropod Cruxicheiros newmanorum gen. et sp. nov. pelvic bones (WARMS 15771) and right femur (WARMS 15770) from the Chipping Norton Limestone Formation, Bathonian of the United Kingdom. A. Left ilium in lateral view (A1, A2) and pubic peduncle in ventral view (A3). B. Left pubis in lateral (B1) and medial (B2) views. C. Proximal portion of right femur in proximal (C1), medial (C2), posterior (C3, C4) and anterior (C5) views. D. Shaft fragment of right femur in lateral view. E. Distal portion of right femur in medial (E1), posterior (E2) and anterior (E3) views.

f03_35.eps

Fig. 4.

Simplified cladogram showing the alternative possible phylogenetic placements of Cruxicheros newmanorum based on analysis of the modified data set of Benson (2010). Fukuiraptor was not included in this scheme as it was recently demonstrated to be a derived allosauroid and not a basal neotetanuran (Benson et al. 2010). C. newmanorum was recovered as either a basal megalosauroid (A), a basal tetanuran (B) or a basal neotetanuran (C).

f04_35.eps

Discussion

Affinities of Cruxicheiros.—Although preserved materials of Cruxicheiros are limited, various features support determination of its systematic affinities. The swollen ridge on the lateral surface of the iliac blade, anterior spur of the caudal neural spine and widely-separated cervical prezygapophyses are unique tetanuran synapomorphies (Rauhut 2003, Benson 2010). The posterior flange of the femoral caput is prominent, extending past the posterior surface of the femoral head. In megalosaurids more derived than Eustreptospondylus, such as Afrovenator (UC OBA 1) and Megalosaurus, the posterior flange is low and does not extend past the posterior surface of the head, so it is unlikely that Cruxicheiros is a derived megalosaurid. The dorsal neural spines of Cruxicheiros are unlike the tall, transversely narrow spines of Megalosaurus (Benson 2010), but resemble those of Marshosaurus (DMNH 3718), Piatnitzkysaurus (Bonaparte 1986) and some allosauroids, including Aerosteon (Sereno et al. 2008), Allosaurus (Madsen 1976), and Neovenator (Brusatte et al. 2008). This resemblance results from their relatively low dorsoventral height, combined with prominent, rugose ridges occupying the anterior and posterior surfaces and the transversely expanded dorsal ends. Contrastingly, in most basal tetanurans with similar spine proportions, such as Monolophosaurus (Zhao and Currie 1994) and some megalosaurids (Britt 1991; Sadleir et al. 2008), rugose ridges on the anterior and posterior surfaces of the dorsal neural spines are generally less prominent and the dorsal end of the spine is less expanded.

The pubic peduncle of the ilium is reconstructed as being approximately as long anteroposteriorly as it is broad mediolaterally. This is shared with non-tetanuran theropods but is also present as a local autapomorphy in the basal megalosaurid Eustreptospondylus (Sadleir et al. 2008: pl. 20: 4) and the allosauroid Fukuiraptor (Azuma and Currie 2000). This feature in Cruxicheiros suggests a possible basal position within Tetanurae. The proximal end of the femoral head bears an oblique “articular” groove. This feature is widespread among basal theropods but is lost in neotetanurans including allosauroids (Hutchinson 2001; Benson 2010).

Cruxicheiros was scored for the phylogenetic data matrix of Benson (2010). The matrix was slightly corrected based on recent descriptive work on Monolophosaurus, which is now scored as possessing a straight pubis (ch. 171.0) (Zhao et al. 2010) and narrow paradental groove (ch. 81.0) (Brusatte et al. in press). After a priori exclusion of “wildcard” taxa identified by Benson (2010) we recovered 24 most parsimonious cladograms (MPTs) of length 585 steps, an ensemble consistency index (CI) of 0.444, a retention index (RI) of 0.618, and a rescaled consistency index (RC) of 0.275. Relationships among taxa included in the original analysis were unaltered and Cruxicheiros was recovered in three positions, each represented by eight of the MPTs: the most basal megalosauroid, branching off outside of all megalosauroid clades recovered by Benson (2010); the most basal neotetanuran, branching off outside of the split between Fukuiraptor, Allosauroidea, and Coelurosauria; or the most basal tetanuran, branching off outside of the split between Megalosauroidea and Neotetanurae. These results suggest that Cruxicheiros represents a very basal tetanuran but that current materials are insufficient to determine its precise phylogenetic affinities. When Cruxicheiros was constrained to fall within a monophyletic Megalosauridae, analysis resulted in 8 trees of 586 steps. In all these trees Cruxicheiros was the sister taxon of Eustreptospondylus, the most basal megalosaurid. When Cruxicheiros was constrained to be the sister taxon of Megalosaurus bucklandii, analysis recovered 8 trees of 588 steps. This confirms that Cruxicheiros does not belong within the genus Megalosaurus. With only one additional step, Cruxicheiros falls within Megalosauridae, but the small size of this length increase may arise from incomplete knowledge of the taxon (93.9% missing data) rather than the possibility of megalosaurid affinities.

UK Bathonian theropods.—The presence of Cruxicheiros yields new data on the diversity of large-bodied theropods represented by abundant, but often fragmentary, specimens from the Lower-Middle Bathonian of the United Kingdom. This assemblage has often been interpreted as a monospecific fauna of Megalosaurus bucklandii, essentially for taxonomic convenience (e.g., Benton and Spencer 1995; Weishampel et al. 2004), and recent suggestions that two such taxa were present in key localities at Stonesfield, Oxfordshire and New Park Quarry, Gloucestershire (Allain and Chure 2002; Day and Barrett 2004; Naish and Martill 2007) are unfounded (Benson 2009). Consequently, material of Cruxicheiros is the first unequivocal report of an additional large-bodied theropod taxon from British Bathonian strata. A third taxon represented by more fragmentary remains from the Lower Bathonian of Oakham Quarry, Oxfordshire will be the subject of a later publication (RBJB, unpublished data).

Acknowledgements

We thank the Newman family of Cross Hands Quarry for providing access to their quarry and correspondence, and to David Walker (formerly BIRMG) for further information on the history of the specimens. We also thank Jennifer Clack (University of Cambridge, Cambridge, UK) and Phil Powell (OUMNH) for clarifying details of the provenance. RBJB is supported by a fellowship from Trinity College, Cambridge and thanks Leslie Noè (Thinktank Museum, Birmingham, UK) and Phil Watson (BIRMG) for making him aware of the specimens. JDR additionally thanks Phil Watson for facilitating the transfer of the material to the Warwickshire Museum. SYNTHESYS, the Palaeontographical Society and the Jurassic Foundation provided funding to examine comparative material.

References

  1. R. Allain 2005. The postcranial anatomy of the megalosaur Dubreuillosaurus valesdunensis (Dinosauria, Theropoda) from the Middle Jurassic of Normandy, France. Journal of Vertebrate Paleontology 25: 850–858. Google Scholar
  2. R. Allain and DJ. Chure 2002. Poekilopleuron bucklandii, the theropod dinosaur from the Middle Jurassic (Bathonian) of Normandy. Palaeontology 45: 1107–1121. doi: 10.1111/1475-4983.00277 Google Scholar
  3. Y. Azuma and P.J. Currie 2000. A new carnosaur (Dinosauria: Theropoda) from the Lower Cretaceous of Japan. Canadian Journal of Earth Sciences 37: 1735–1753. doi: 10.1139/cjes-37-12-1735 Google Scholar
  4. R.B.J. Benson 2009. An assessment of variability in theropod dinosaur remains from the Bathonian (Middle Jurassic) of Stonesfield and New Park Quarry, UK and taxonomic implications for Megalosaurus bucklandii and Iliosuchus incognitus. Palaeontology 52: 857–877. doi: 10.1111/j.1475-4983.2009.00884.x Google Scholar
  5. R.B.J. Benson 2010. A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods. Zoological Journal of the Linnean Society. doi: 10.HH/j.1096-3642.2009.00569.x Google Scholar
  6. M.J. Benton and P.S. Spencer 1995. Fossil Reptiles of Great Britain. 386 pp. Chapman and Hall, London. Google Scholar
  7. J.F. Bonaparte 1986. Les dinosaurs (Carnosaures, Alosauridés, Sauropodes, Cétiosaurides) du Jurassique moyen de Cerro Cóndor (Chubut, Argentine). Annales de Paléontologie (Vertebrés-Invertebrés) 72: 247–289. Google Scholar
  8. B. B. Britt 1991. Theropods of Dry Mesa Quarry (Morrison Formation, Late Jurassic), Colorado, with emphasis on the osteology of Torvosaurus tanneri. Brigham Young University Geology Studies 37: 1–72. Google Scholar
  9. B.B. Britt 1993. Pneumatic Postcranial Bones in Dinosaurs and Other Archosaurs. 320 pp. Unpublished Ph.D. thesis, University of Calgary, Calgary. Google Scholar
  10. C.A. Brochu 2002. Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Society of Vertebrate Paleontology Memoirs 7: 1–138. doi: 10.2307/3889334 Google Scholar
  11. R. Brookes 1763. The Natural History of Waters, Earths, Stones, Fossils, and Minerals, with their Virtues, Properties and Medicinal Uses. 364 pp. Newbery, London. Google Scholar
  12. S.L. Brusatte , R.B.J. Benson , X.-J. Zhao , and P.J. Currie (in press). The skull of Monolophosaurus jiangi (Dinosauria: Theropoda) and its implications for early theropod phylogeny and evolution. Zoological Journal of the Linnean Society.  Google Scholar
  13. S.L. Brusatte , R.B.J. Benson , and S. Hutt 2008. The osteology of Neovenator salerii (Dinosauria: Theropoda) from the Wealden Group (Barremian) of the Isle of Wight. Palaeontographical Society Monographs 162 (631): 1–75. Google Scholar
  14. W. Buckland 1824. Notice on the Megalosaurus or great fossil lizard of Stonefield. Transactions of the Geological Society 21: 390–397. Google Scholar
  15. P.J. Currie and X.-J. Zhao 1994. A new carnosaur (Dinosauria: Theropoda) from the Jurassic of Xinjiang, People's Republic of China. Canadian Journal of Earth Sciences 30 (1993): 2037–2081. Google Scholar
  16. J.J. Day and P.M. Barrett 2004. Material referred to Megalosaurus (Dinosauria: Theropoda) from the Middle Jurassic of Stonesfield, Oxfordshire England: one taxon or two?, Proceedings of the Geologists' Association 115: 359–366. Google Scholar
  17. J.B. Delair and W.A.S. Sarjeant 1975. The earliest discoveries of dinosaurs. Isis 66: 5–25. doi: 10.1086/351373 Google Scholar
  18. J.B. Delair and W.A.S. Sarjeant 2002. The earliest discoveries of dinosaurs: the records re-examined. Proceedings of the Geologists' Association 113: 185–197. Google Scholar
  19. J.A. Gauthier 1986. Saurischian monophyly and the origin of birds. In : K. Padian (ed.), The Origin of Birds and the Evolution of Flight. Memoirs of the California Academy of Science 8: 1–55. Google Scholar
  20. T.R. Holtz Jr ., R.E. Molnar , and P.J. Currie 2004. Basal Tetanurae. In : D.B. Weishampel , P. Dodson , and H. Osmólska (eds.), The Dinosauria , 2nd Edition, 71–110. University of California Press, Berkeley. Google Scholar
  21. F. von Huene 1926. The carnivorous Saurischia in the Jura and Cretaceous formations, principally in Europe. Revista del Museo de La Plata 29: 35–167. Google Scholar
  22. J.R. Hutchinson 2001. The evolution of femoral osteology and soft tissues on the line to extant birds (Neornithes). Zoological Journal of the Linnean Society 131: 169–197. doi:10.1111/j.1096-3642.2001.tb01314.x Google Scholar
  23. E. Lluyd 1699. Lithophylacii Britannici Ichnographia. 156 pp. Clarendon Press, Oxford. Google Scholar
  24. J.H. Jr. Madsen 1976. Allosaurus fragilis: a revised osteology. Utah Geological and Mineralogical Survey Bulletin 109: 3–163. Google Scholar
  25. J.H. Madsen Jr and S. P. Welles 2000. Ceratosaurus (Dinosauria, Theropoda) a revised osteology. Miscellaneous Publications of the Utah Geological Survey 2: 1–80. Google Scholar
  26. O.C. Marsh 1881. Classification of the Dinosauria. American Journal of Science (Series 3) 16: 411–416. Google Scholar
  27. 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 164: 493–510. doi: 10.1144/0016-76492006-032 Google Scholar
  28. R. Owen 1842. Report on British fossil reptiles. Part II. Reports of the British Association for the Advancement of Science 1841: 60–204. Google Scholar
  29. J. Parkinson 1822. Outlines of Oryctology. 346 pp. Published by the author, London. Google Scholar
  30. J. Phillips 1871. Geology of Oxford and the Valley of the Thames. 523 pp. Clarendon Press, Oxford. Google Scholar
  31. R. Plot 1677. The Natural History of Oxfordshire, Being an Essay Towards the Natural History of England. 358 pp. Published by the author, Oxford. Google Scholar
  32. O.W.M. Rauhut 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology 69: 1–213. Google Scholar
  33. O.W.M. Rauhut and A.C. Milner 2008. Cranial anatomy and systematic position of the Middle Jurassic theropod dinosaurs Proceratosaurus from England. Journal of Vertebrate Paleontology 28: 130A. Google Scholar
  34. Rauhut O.W.M ., A.C. Milner , and S. Moore-Fay 2010. Cranial osteology and phylogenetic position of the theropod dinosaur Proceratosaurus bradleyi (Woodward, 1910) from the Middle Jurassic of England. Zoological Journal of the Linnean Society , doi: 10.1111/j.1096-3642.2009.00591.x Google Scholar
  35. R.W. Sadleir , P.M. Barrett , and H.P. Powell 2008. The anatomy and systematics of Eustreptospondylus oxoniensis, a theropod dinosaur from the Middle Jurassic of Oxfordshire, England. Palaeontographical Society Monographs 160 (627) [for 2006]: 1–82. Google Scholar
  36. B.W. Sellwood and W.S. McKerrow 1974. Depositional environments in the lower part of the Great Oolite Group of Oxfordshire and North Gloucestershire. Proceedings of the Geologists' Association 85: 190–210. Google Scholar
  37. P.C. Sereno , R.N. Martinez , J.A. Wilson , D.J. Varicchio , O.A. Alcober , and H.C.E. Larsson 2008. Evidence for avian intrathoracic air sacs in a new predatory dinosaur from Argentina. PLoS ONE 3 (9): 1–20. doi:10.1371/journal.pone.0003303 PMid: 18825273, PMCid:2553519 Google Scholar
  38. R. Steel 1970. Saurischia. Handbuch der Paläoherpetologie 14: 1–84. Google Scholar
  39. M.G. Sumbler 2002. Cross Hands Quarry, Warwickshire. In : B.M. Cox and M.G. Sumbler (eds.), British Middle Jurassic Stratigraphy. Geological Conservation Review Series 26: 202–204. Google Scholar
  40. H.S. Torrens 1980. Bathonian Correlation Chart. In : J.C.W. Cope , K.L. Duff , C.F. Parsons , H.S. Torrens , W.A. Wimbledon , and J.K. Wright (eds.), A Correlation of Jurassic Rocks in the British Isles, Part Two: Middle and Upper Jurassic. Geological Society of London Special Report 15: 21–45. Google Scholar
  41. D.B. Weishampel , P.M. Barrett , R.A. Coria , J. Le Loeuff , X. Xu , X.-J. Zhao , A. Sahni , E.M. Gomani , and CR. Noto 2004. Dinosaur Distribution. In : D.B. Weishampel , P. Dodson , and H. Osmólska (eds.), The Dinosauria , 2nd Edition, 517–606. University of California Press, Berkeley. Google Scholar
  42. J. Woodward 1728. An Attempt Towards a Natural History of the Fossils of England: in a Catalogue of the English Fossils in the Collection of J. Woodward. F. Fayram, J. Senex, J. Osborn, and T. Longman, London. Google Scholar
  43. X.-J. Zhao , R.B.J. Benson , S.L. Brusatte , and P.J. Currie 2010. The postcranial skeleton of Monolophosaurus jiangi (Dinosauria: Theropoda) from the Middle Jurassic of Xinjiang, China, and a review of Middle Jurassic Chinese theropods. Geological Magazine 147:13–27. doi:10.1017/S0016756809990240. Google Scholar

Appendices

Appendix 1

Scores for Cruxicheiros newmanorum for the data set of Benson (2010). Cruxicheiros newmanorum

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Roger B.J. Benson and Jonathan D. Radley "A New Large-Bodied Theropod Dinosaur from the Middle Jurassic of Warwickshire, United Kingdom," Acta Palaeontologica Polonica 55(1), (20 November 2009). https://doi.org/10.4202/app.2009.0083
Received: 8 July 2009; Accepted: 1 November 2009; Published: 20 November 2009
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