Historical Biology, 2014 Vol. 26, No. 2, 229235, http://dx.doi.org/10.1080/08912963.2013.

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Alfred Nicholson Leeds and the rst fossil egg attributed to a saurian
J.J. Listona,b,c,d* and S.D. Chapmane
Yunnan Key Laboratory for Palaeobiology, Yunnan University, Kunming, Peoples Republic of China; bNational Museums Scotland, Chambers Street, Old Town, Edinburgh EH1 1JF, Scotland, UK; cSchool of Earth Sciences, University of Bristol, Wills Memorial Building, Queens Road, Bristol BS8 1RJ, UK; dInstitute of Biodiversity, Animal Health and Comparative Medicine, College of Medical, Veterinary and Life Sciences, University of Glasgow, Glasgow G12 8QQ, UK; eDepartment of Earth Sciences, The Natural History Museum, Cromwell Road, London SW7 5BD, UK (Received 22 May 2013; nal version received 26 May 2013; rst published online 18 July 2013) Discovered by the nineteenth century collector Alfred Nicholson Leeds, the rst object to be described (1898) as a fossil reptile egg is a unique nd from the Oxford Clay near Peterborough. It also comes from one of a very small number of Jurassic localities worldwide that can claim to have yielded a fossil egg. Given its historical and contemporary signicance, this object is reassessed in the light of increased understanding of such objects. Data from scanning electron microscopy, computerised tomography, synchrotron imaging, X-ray diffraction and petrographic thin sectioning prove inconclusive. However, the presence of apparent external openings resembling angusticanaliculate pores a pore type common only to certain types of dinosaur eggshell in both size and sparseness of distribution prevents its summary dismissal as not being a dinosaurian egg. Keywords: Alfred Nicholson Leeds; dinosaur egg; Oxford Clay; Callovian
a

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Introduction The Alfred Leeds collection is one of the major fossil marine reptile collections in the world (Liston 2006). It was assembled by the gentleman-farmer Alfred Nicholson Leeds (1847 1917) from specimens retrieved from , 30 pits excavated in the Middle Jurassic (Callovian) Oxford Clay in the vicinity of Peterborough, England, by a variety of brick manufacturing companies since 1874. The popularity of this clay with these companies was because of its high-carbon content, which made the bricks effectively self-ring, therefore signicantly cheaper to manufacture and so more protable to sell. From this sudden industrial increase in excavation, a steady parallel exposure of fossils for collectors resulted. The ease with which the clay could be washed from three-dimensionally preserved vertebrate skeletons meant that the fossilised animal remains could be displayed in the same way as contemporary zoological skeletons, rather than mounted in a limestone slab, as with many marine reptile fossils commonly available at the time to museums (e.g. from Holzmaden or Solnhofen). This preservation meant the skeletons lent themselves more to detailed study, and enabled workers such as Lydekker (1889), Seeley (1889), Smith (1889) and most signicantly Andrews (1910, 1913) to describe many new taxa from Alfred Leeds collection. This resulted in type material distributed between more than 650 specimens at the Hunterian Museum (University of Glasgow) and more than 350 at the Natural History Museum (London). The total number of vertebrate

specimens excavated by Alfred Leeds vastly exceeds 1000 individuals, which includes the holdings of many museums around the world that bought Alfred Leeds material (usually simply marked Oxford Clay, Peterbor rtz of Bonn. These ough) from the dealer Bernard Stu gures serve to illustrate that Alfred Leeds was no casual collector, and had a very great familiarity with fossils from the Oxford Clay. Indeed, his understanding of the distinctive nature of the fossil animals from the Oxford Clay was often greater than that of his contemporary professional peers (Young et al. 2013), and therefore it is worth respecting his opinion today, with regard to objects that he identied as unusual within the context of his prodigious experience of excavation of the Callovian clay. Among the marine reptiles that form the bulk of his collection, there are more rare items, such as the remains et al. 2010) and the rst of a pterosaur, dinosaurs (Noe large vertebrate suspension feeders, which are the pioneers of the planktivorous niche occupied today by whale sharks, basking sharks and baleen whales (Liston et al. 2013). In 1898, Alfred Leeds sold another highly unusual fossil to the (then) British Museum (Natural History): accessioned as an egg of a saurian, NHMUK PV R2903 possessed only curiosity value, until the announcement of the discovery of dinosaur eggs in the Gobi Desert by the American Museum of Natural Historys Third Asiatic expedition to Mongolia in 1922, led by Roy Chapman Andrews. In response, The Sphere in 1923 proclaimed that

*Corresponding author. Email: leedsichthys@gmail.com

The contribution of S. Chapman was authored as part of her employment and therefore copyright is asserted and retained in the contribution by The Natural History Museum. J. Liston waives his right to any copyright in the Article but not his right to be named as co-author of the Article.

230 J.J. Liston and S.D. Chapman these discoveries were nothing new, as the Oxford Clay had yielded a fossilised reptile egg in England some years earlier (A fossil reptiles eggs unearthed in England 1923; Leeds 1956, p. 76). Thus, the claim to the rst fossilised reptile egg had already been made. Eggshell fragments from dinosaur/reptile eggs had been found before; Philippe Matheron had described the rst such remains from the south of France in 1859 (Buffetaut and Le Loeuff 1994). However, this claim was new because the Oxford Clay egg appeared intact and more or less complete. In 1928, Van Straelen published a review of fossil eggs, and gured the Oxford Clay specimen together with a description; however, he was cautious about referring to it as an egg (Van Straelen 1928). Again, in 1950, dinosaur eggs were reported from Tanzania. In response, the British Museums William Elgin Swinton was commissioned by the Illustrated London News to do an article on the discovery (Swinton 1950), within which he referred to the Oxford Clay egg and suggested that it might have been laid by an amphibious dinosaur. The Alfred Leeds curio has subsequently been overlooked for decades and continues to lack a satisfactory identication. Although included in Carpenter and Alf (1994; no. 71 on p. 19) and Carpenters (1999; no. 16 on p. 260) checklists of nds, where it is listed as camptosaurid(?) egg, it has received little serious consideration since the work of Van Straelen (1928). The reason for this may be due to the perceived unlikelihood of a fossil egg occurring within the marine deposits of the Oxford Clay. However, recent work (Hayward et al. 1997) indicates the high survivability of bird eggs in marine environments, despite the rigours of the recruitment process. Hayward et al. (1997) report eggs even surviving uncracked at depths in excess of 500 m. Evans (2012) went further and also explored the implications of oating eggs as a mechanism for gene ow for terrestrial egg-laying vertebrates between land masses. This suggests that a number of mechanisms exist to effectively introduce an amniote egg to the marine environment with a real possibility of preservation (Figure 1). In 1995, NHMUK PV R2903 was selected for a series of tests being carried out on fossil eggs at Glasgow University, using a variety of available scanning technologies for non-invasive analysis (Liston and McJury 2003). These revealed the internal density contrasts of the object, and appeared to show a discrete mass of components within the object. Unexpectedly, the Oxford Clay egg produced the most promising results in terms of an object that might actually contain fossilised remains (Liston and McJury 2003). Given this result, and the global scarcity of Jurassic eggs (e.g. of 135 dinosaur egg sites globally, only 13 or 9% are Jurassic in age, Weishampel et al. 2004; Paik et al. 2012), we considered it important to

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Figure 1. Sketch illustrating the three means of delivery of a saurian egg into the Jurassic marine realm: simple recruitment as per the works of Evans (2012) and Hayward et al. (1997); transportation in the oating corpse of a maternal animal and transportation by a ying reptile or pteroportation. Copyright and courtesy of Robert Nicholls at www.paleocreations.com.

reassess this specimen from the Oxford Clay with the aim of clarifying its nature.

Materials and methods As a unique object within the collections of the Natural History Museum (London), NHMUK PV R2903 was subject to severe restrictions in terms of analyses. For example, no samples could be removed from the specimen, and gold or carbon coating of the specimens for scanning electron microscopy (SEM) was prohibited. This meant that analyses were constrained to nondestructive or non-invasive scanning methods, and scanning electron microscopy in particular required low vacuum conditions. A variety of analyses were thus carried out. First, the Vale of Leven Hospital helical medical computerised tomography (CT) scanner was used, signicantly improving on the data previously obtained at the Western Inrmarys facility (Liston and McJury 2003). The resulting data were then processed using Voxar 3D software (Toshiba Medical Visualization Systems). Secondary electron images were obtained using an FEI 200F eld-emission environmental SEM (FEG-ESEM) at the University of Glasgows School of Geographical and Earth Sciences. The microscope was operated at 20 kV, using a moderately high-beam current (not quantied on this instrument), and at a vacuum of , 50 Pa. Under these low vacuum conditions, the negative charge that accumulates on the sample surface is compensated by water vapour in the chamber, so that high-resolution images could be obtained from the sample and without the need for a conductive carbon or gold coating (e.g. Flude et al. 2012).

Historical Biology 231 The synchrotron imaging system at the European Synchrotron Radiation Facility (ESRF, Grenoble, France) was used, unfortunately causing severe re damage to the specimen after markers added to the surface combusted. This, however, produced detached pieces of shell for X-ray diffraction (XRD), high-magnication SEM and thin-sectioning work. For XRD, an INEL Inc.-manufactured curved position-sensitive detector (calibrated with silicon and silver behenate standards) was used to collect diffraction patterns from samples powdered and loaded onto a single-crystal quartz substrate. With an output of 4096 digital channels representing an arc of 1208, the simultaneous measurement of diffracted X-ray intensities at all angles of 2u across 1208 was possible. A second phase of SEM work was nally conducted at the Natural History Museum (London) using a Carl Zeiss Leo 1455-VP. pattern of lines radiate from the depressed centre of the surface. The outer perimeter of the specimen exhibits variable thickness and in some areas, it can be seen that the putative eggshell layer extends onto this surface from the convex side. Edward Thurlow Leeds (the second of Alfred Leeds ve sons) noted that ammonites immediately above the Gryphaea zone (some 3 m above the Cornbrash) in the Oxford Clay around Peterborough occur as large specimens in concretions with a thin coating of decayed shell or with a rough coat of pyrite, and are rarely well preserved (Leeds 1956, p. 79). He also noted that the higher zones of the Oxford Clay, as exposed in the Eye Pit, were richly productive in pyritised ammonites, whereas its lower zones only yielded attened shells (Leeds 1956, p. 79), not unlike the one visible on the concave surface of NHMUK PV R2903. The shell of the ammonite bears the characteristic ribbing and tuberculation of Kosmoceras jason.

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Description The greyish brown specimen measures 125.96 mm long, 108.22 mm wide and 33.32 mm high, and weighs , 980 g. The convex face of the specimen (Figure 2(a)) shows a cracked face, which looks like a mosaic of small fragments of eggshell that individually curl slightly outward from the surface of the object at their edges, revealing an apparent thickness of 0.457 mm. The clay matrix supporting the mosaic of fragments appears to be continuous; therefore, the cracking of the surface may be due to desiccation prior to recruitment and burial. The cracks are lled with sediment that binds the shell-like fragments together. This surface has been consolidated with a varnish. Shell-like fragments on the convex surface over-ride the central region of the object, indicating lithostatic compaction. The concave surface (Figure 2(b)) is primarily smooth and lacks the eggshell-like appearance of the opposite convex surface. Molluscan shell fragments, small bivalve shells and sediment with impressions of ammonite whorls cover portions of this surface of the object. Some whorl impressions are covered with a nacreous layer of shell. A

Results CT scan Colour texturing was applied to the different densities determined by the Vale of Leven helical CT scanner using Voxar 3D software (Figure 3(a) (c)), which revealed that the egg structure was partial, and conrmed not only that part of an ammonite comprised the objects concave surface, but also that fragments of a belemnite were situated within the object. Clarifying the physical structure of the specimen allowed appropriate areas to be selected for subsequent SEM analysis.

X-ray diffraction The results (Figure 4) show that the eggshell sample taken from the convex surface of the object consists of calcite; however, the shift in the main peak position relative to pure calcite indicates that a cation slightly

Figure 2. Specimen NHMUK PV R2903. (a) Convex surface; (b) concave surface and (c) convex surface after re damage. Scale bar 10 cm. Copyright image resources: NHM.

232 J.J. Liston and S.D. Chapman

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Figure 3. (a c) Images produced after colour texturing was applied to the original Vale of Leven CT scans using Voxar 3D with progressive virtual removal of matrix and shell. Note the long belemnite fragment in both (b) and (c).

Synchrotron imaging Use of the synchrotron imaging system caused severe re damage to the specimen (Figure 2(c)); however, we were still able to image the components present without their matrix, and clearly distinguish the putative partial egg from the ammonite and belemnite (Figure 5).

SEM and light microscopy Preliminary low magnication oblique SEM surface analysis indicated a triple-laminated structure (Figure 6(a)) of around 400 mm thick. Subsequent higher magnication radial sections from samples revealed the

Figure 4. XRD pattern showing results of samples from both convex and concave surfaces. Both pure and impure calcite peaks lie between 29 and 30u (G. Cressey, pers. obs.) note that the 4% peak shift between them, indicating likely Mg replacement of Ca in the calcite (G. Cressey, pers. obs.) in the putative eggshell sample.

smaller than Ca is present in small amounts throughout the structure. This cation is probably Mg and is estimated, from the peak shift, to be present at a level of , 4% replacement of Ca by Mg, relative to a standard calcite pattern. Some pyrite was also detected in this sample. In contrast, the ammonite shell debris from the concave surface contains aragonite and calcite, but this is nearly pure calcite with no apparent substitution of Ca by Mg.

Figure 5. Graphical results from synchrotron, showing distinct character of the three components: the ammonite that constitutes much of the concave surface, the fragments of the belemnite within and the third component that constitutes the convex surface.

Historical Biology 233

Figure 6. Scanning electron micrographs of the putative eggshell of NHMUK PV R2903. (a) Oblique view of the surface, showing layered structure. Scale 1.0 mm. (b) Radial view showing crystalline layer. Scale bar 10 mm. (c) Surface view showing pore. Scale bar 100 mm.

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presence of a radially orientated crystalline layer around 45 mm thick (Figure 6(b)) with individual prisms of this layer possessing a width of around 2.5 mm, the rest being obscured by clay matrix. Higher magnication SEM of the outer surface also showed sub-circular and oval pore-like openings ranging from 9 to 27 mm in diameter (Figure 6(c)). Observed gaps between these structures ranged between 157 and 211 mm. Radial petrographic thin sections show that clay adheres to the eroded outer surface (similar to Carpenter 1999, Figure AII.26), and the inner surface exhibits a secondary calcite development. Two distinct layers are visible beneath the clay. High power (Figure 7(b)) and cross polars (Figure 7(c)) reveal a lack of any internal structure.

Discussion From the distinct XRD peak discriminating the convex surface from that of the ammonite shell of the concave surface, and reconstruction of the CT and synchrotron scans (Figures 3 and 5), NHMUK PV R2903 has been shown to consist of three separate components. As revealed by the synchrotron imaging system, NHMUK PV R2903 represents an unusual assemblage of ammonite, belemnite and a third component. The nature of this third component remains uncertain, although its regular shape

would be consistent with the partial remains of one half of an egg. Although it remains somewhat problematic to model the process, by which a partial (rather than a complete) egg would be preserved on the seaoor, this result immediately falsies the hypothesis that NHMUK PV R2903 represents the rst complete fossil egg. Given that belemnites are common throughout the Oxford Clay, as also are accumulations of ammonites lying on top of each other (JJL, pers. obs.), it is not surprising that such objects would have ended up lying in an upturned bowl-like structure, such as the third component appears to be, on the seabed. The regular curvature and apparent uniformity of thickness would seem to mitigate against the third component resulting from secondary concretionary growth. So also does the outward curling of the edges of the putative shell fragments, which might be a sign of predepositional dessication prior to recruitment and burial, or a result from drying immediately subsequent to excavation, which has been xed by varnish application. A conclusive result that the third component represents the remains of part of an egg would have required identiable eggshell micro- or ultrastructure. However, neither could be identied with certainty. The low magnication SEM showed 400 mm thickness, which would be at the lower end of eggshell thickness as noted by

Figure 7. Thin sections of sample of putative eggshell with exterior to the top. (a) Under low power. (b) Under high power with plane light, note the appearance of fragments overriding each other in basal layer, although no trace of eggshell microstructure is visible. (c) Under high power with crossed polars, there is no trace of sweeping extinction or prismatic structure. Scale bar for (b) and (c) 100 mm.

234 J.J. Liston and S.D. Chapman Hirsch (1994) and Ensom (2002), and apparently lacking ornamentation commonly described for many avian and other dinosaurian eggshells (Hirsch and Packard 1987). However, the low magnication renders this image ultimately unhelpful. When SEM was repeated on samples at higher magnication, the only layer detectable was a crystalline layer with radially orientated prisms. The 45 mm thickness of this layer is much less than what would be expected in a prismatic layer in eggshell, and the 2.5 mm width of individual prisms is similarly signicantly less than the 10 20 mm range measured by Mikhailov (1997, plate 8) and Ensom (2002, plate 3, gure 8). As both these measurements are outside the ranges observed for biological calcite in eggshell, it seems most likely that this is a thin isolated pocket of post-depositional calcite growth. In contrast, the petrographic thin sections gave a whole prole of the putative eggshell with two clear layers visible: a well-demarcated inner layer and an outer heavily eroded layer. The outer layer showed no sign of prismatic structure, although this could be because of recrystallisation, consistent with the observed secondary calcite deposits. To interpret this section as eggshell, the internal layer would represent an unusually thin (, 70 mm) lower zone or mammillary layer with the outer layer representing the eroded remains of the prismatic layer after erosion of the external layer. But with this outer layer lacking any clearly denable outer perimeter and hence an absence of any possible surface ornamentation on an external layer, it is hard to identify that this layer indeed represents the prismatic layer. The absence of a clear perimeter also prohibits the measurement of a regular thickness. The structures resembling round-to-oval pore openings were particularly narrow in diameter in terms of pores observed in eggshell (Mikhailov 1997), but are consistent in both their shape and dimensions (within the range of 10 100 mm) with the openings of angusticanaliculate pores (Mikhailov 1997). However, no canals were visible in radial thin section to determine their nature beneath the openings (e.g. straight, as would be expected if these were angusticanaliculate pores). The sparse distribution of these structures, with observed gaps of 157 211 mm, also recalls the observed infrequency of angusticanaliculate pores on the surface of eggshells (Mikhailov 1997). Although estimates based on assuming that this spacing is regular make these structures appear signicantly more common than the 3 20 pores per 100 mm2 quoted by Mikhailov (1997), perhaps by as much as a factor of a hundred, it is worth noting that adjacent pores were only noted for two of the seven openings observed, so this is likely to be an overestimate. These structures, combined with the overall 3D shape, remain the most compelling arguments for the third component of NHMUK PV R2903 to be interpreted as representing the remains of part of an egg. Unsurprisingly, given the recorded infrequent distribution and low density of angusticanaliculate pores in eggshell, none were visible in radial section. However, these structures are consistent in size with the range expected for angusticanaliculate pores, and are difcult to explain as a result of dissolution or other diagenetic alterations. Mikhailov (1997) notes angusticanaliculate pores as indicative of a dry terrestrial incubating environment, and furthermore notes their occurrence in dinosauroidprismatic and ornithoid basic shell types, namely the general egg parataxa assigned to Theropoda (Prismatoolithidae and Elongatoolithidae) and Enantiornithes (Laevisoolithidae) (Mikhailov 1997, p. 39, Table 1; Carpenter 1999). He further noted that prismatoolithid eggshell appeared to be the only dinosaurian eggshell type that was common prior to the Cretaceous (Mikhailov 1997, p. 59). These observations are consistent with this specimen coming from Middle Jurassic clays that have yielded many et al. 2010). However, it must be dinosaur remains (Noe stressed that the lower zone thicknesses exhibited by these general parataxa (Prismatoolithidae circa 200 mm, Elongatoolithidae 170 300 m m, Laevisoolithidae 125 200 mm, all data from Mikhailov 1997) as well as the prism widths (10 20 mm range for these three groups, again all data from Mikhailov 1997) are considerably in excess of the gures of 70 mm (putative lower zone thickness) and 2.5 mm (putative prism width, if the radial crystalline layer is not, in fact, merely secondary calcite growth) that the third component of NHMUK PV R2903 exhibits. These numerical departures must, therefore, raise some questions over this interpretation, and in particular raise the issue that the layer visible under SEM might simply represent geologically rather than biologically directed calcite growth. In summary, the lack of a visible complete undamaged section with all layers measurable is problematic. The eroded outer surface, in particular, means that any external ornamentation present that would help with identication is also absent. The thicknesses recorded for the layers and prisms are signicantly less than the recognised ranges for known eggs. But, given the overall regular 3D shape, and in the absence of another means by which to explain the porelike openings, it still seems simplest currently to regard NHMUK PV R2903 as possibly representing a fossilised partial egg containing parts of an ammonite and a belemnite. It is hoped that further and more extensive sampling of the specimen can be permitted at some point to conclusively determine the nature of this third component. Conclusions Although the ultrastructure and microstructure could not be identied with certainty, NHMUK PV R2903 may indeed represent a recrystallised and damaged fossilised partial dinosaur egg with an angusticanaliculate pore

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Historical Biology 235 system. It has not yet been possible to disprove Alfred Leeds judgement of one of his specimens as the egg of a saurian. This is remarkable, given that by the end of the nineteenth century, fossil eggshell work was effectively unknown, and a testament to both his skills as a collector and as an observer in the eld. Acknowledgements
The authors thank both reviewers for much assistance in improving the text, Frankie Jackson and Colin Shaw (Figure 7(b),(c)), Montana State University, for providing thin-section images; Gordon Cressey and Peter Tandy, Department of Earth Sciences (Mineralogy), NHM, London, for the XRD analysis (Figure 4); Tony Wighton, NHM, London, for thin sections (Figure 7(a)); Phil Crabb, Photographic Studio, NHM, London for images of the Leeds egg; Vincent Fernandez for SYNCHROTRON access; Steve Baker, Department of Earth Sciences (Palaeontology), NHM, London, for providing access to the collections and help identifying the ammonite; Kirsty Ross, Senior Radiographer at the Western Inrmary and Vale of Leven Hospitals (Glasgow), for her innite patience in steadfastly piloting CT scanners through the weird fossil remains that JJL has for far too many years been bringing her; Caroline Smith and Martin Lee (Figure 6(a)) for steering the Lo-Vac SEM in the University of Glasgows Gregory Building; Bobby Davey for processing the DICOM scans into glorious Technicolor using the Voxar 3D software package; Bob Nicholls for fantastic eleventh hour Pteroportation work at www.paleocreations.com; Ian Rolfe for selessly transporting a fossil egg from London to Glasgow, and Eric Buffetaut for doing the same from London to Grenoble and Cambridge; and Paul Taylor and Alex Brown (NHM) for taking SEMs (Figure 6(b),(c)) and discussion. We also thank Neil Landman and Al McGowan for helpful molluscan assistance. Mike A. Taylor suggested the rotting maternal corpse as a means of facilitating egg recruitment into the marine realm.
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