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Historical Biology: An International Journal of Paleobiology Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/ghbi20 Alfred Nicholson Leeds and the first fossil egg attributed to a ‘saurian’ J. J. Liston a b c d & S. D. Chapman e a Yunnan Key Laboratory for Palaeobiology, Yunnan University , Kunming , People's Republic of China b National Museums Scotland , Chambers Street, Old Town, Edinburgh , EH1 1JF , Scotland, UK c School of Earth Sciences, University of Bristol , Wills Memorial Building, Queen's Road, Bristol , BS8 1RJ , UK d Institute of Biodiversity, Animal Health and Comparative Medicine, College of Medical, Veterinary and Life Sciences, University of Glasgow , Glasgow , G12 8QQ , UK e Department of Earth Sciences , The Natural History Museum , Cromwell Road, London , SW7 5BD , UK Published online: 18 Jul 2013.

To cite this article: Historical Biology (2013): Alfred Nicholson Leeds and the first fossil egg attributed to a ‘saurian’, Historical Biology: An International Journal of Paleobiology, DOI: 10.1080/08912963.2013.809575 To link to this article: http://dx.doi.org/10.1080/08912963.2013.809575

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Alfred Nicholson Leeds and the first fossil egg attributed to a ‘saurian’ J.J. Listona,b,c,d* and S.D. Chapmane aYunnan Key Laboratory for Palaeobiology, Yunnan University, Kunming, People’s 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, Queen’s 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; final version received 26 May 2013)

Discovered by the nineteenth century collector Alfred Nicholson Leeds, the first object to be described (1898) as a fossil reptile egg is a unique find 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 significance, 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

Introduction specimens excavated by Alfred Leeds vastly exceeds The Alfred Leeds collection is one of the major fossil 1000 individuals, which includes the holdings of many marine reptile collections in the world (Liston 2006). It museums around the world that bought Alfred Leeds’ was assembled by the gentleman-farmer Alfred Nicholson material (usually simply marked ‘Oxford Clay, Peterbor- Leeds (1847–1917) from specimens retrieved from ,30 ough’) from the dealer Bernard Stu¨rtz of Bonn. These pits excavated in the Middle Jurassic (Callovian) Oxford figures serve to illustrate that Alfred Leeds was no casual Clay in the vicinity of Peterborough, England, by a variety collector, and had a very great familiarity with fossils from of brick manufacturing companies since 1874. The the Oxford Clay. Indeed, his understanding of the popularity of this clay with these companies was because distinctive nature of the fossil animals from the Oxford of its high-carbon content, which made the bricks Clay was often greater than that of his contemporary effectively ‘self-firing’, therefore significantly cheaper to professional peers (Young et al. 2013), and therefore it is manufacture and so more profitable to sell. From this worth respecting his opinion today, with regard to objects sudden industrial increase in excavation, a steady parallel that he identified as unusual within the context of his Downloaded by [JJ Liston] at 14:10 24 July 2013 exposure of fossils for collectors resulted. The ease with prodigious experience of excavation of the Callovian clay. which the clay could be washed from three-dimensionally Among the marine reptiles that form the bulk of his preserved vertebrate skeletons meant that the fossilised collection, there are more rare items, such as the remains animal remains could be displayed in the same way as of a pterosaur, dinosaurs (Noe´ et al. 2010) and the first contemporary zoological skeletons, rather than mounted in large vertebrate suspension feeders, which are the pioneers a limestone slab, as with many marine reptile fossils of the planktivorous niche occupied today by whale commonly available at the time to museums (e.g. from sharks, basking sharks and baleen whales (Liston et al. Holzmaden or Solnhofen). This preservation meant the 2013). skeletons lent themselves more to detailed study, and In 1898, Alfred Leeds sold another highly unusual enabled workers such as Lydekker (1889), Seeley (1889), fossil to the (then) British Museum (Natural History): Smith (1889) and most significantly Andrews (1910, 1913) accessioned as ‘an egg of a saurian’, NHMUK PV R2903 to describe many new taxa from Alfred Leeds’ collection. possessed only curiosity value, until the announcement of This resulted in type material distributed between more the discovery of dinosaur eggs in the Gobi Desert by the than 650 specimens at the Hunterian Museum (University American Museum of Natural History’s Third Asiatic of Glasgow) and more than 350 at the Natural History expedition to Mongolia in 1922, led by Roy Chapman Museum (London). The total number of vertebrate Andrews. In response, The Sphere in 1923 proclaimed that

*Corresponding author. Email: [email protected]

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. 2 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 reptile’s eggs unearthed in England 1923; Leeds 1956, p. 76). Thus, the claim to the first fossilised reptile egg had already been made. Eggshell fragments from dinosaur/reptile eggs had been found before; Philippe Matheron had described the first 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 figured 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 Figure 1. Sketch illustrating the three means of delivery of a ‘saurian egg’ into the Jurassic marine realm: simple recruitment Tanzania. In response, the British Museum’s William as per the works of Evans (2012) and Hayward et al. (1997); Elgin Swinton was commissioned by the Illustrated transportation in the floating corpse of a maternal animal and London News to do an article on the discovery (Swinton transportation by a flying reptile or ‘pteroportation’. Copyright 1950), within which he referred to the Oxford Clay ‘egg’ and courtesy of Robert Nicholls at www.paleocreations.com. and suggested that it might have been laid by an ‘amphibious dinosaur’. The Alfred Leeds’ curio has subsequently been reassess this specimen from the Oxford Clay with the aim overlooked for decades and continues to lack a satisfactory of clarifying its nature. identification. Although included in Carpenter and Alf (1994; no. 71 on p. 19) and Carpenter’s (1999; no. 16 on Materials and methods p. 260) checklists of finds, where it is listed as ‘camptosaurid(?) egg’, it has received little serious As a unique object within the collections of the Natural consideration since the work of Van Straelen (1928). History Museum (London), NHMUK PV R2903 was The reason for this may be due to the perceived subject to severe restrictions in terms of analyses. For unlikelihood of a fossil egg occurring within the marine example, no samples could be removed from the deposits of the Oxford Clay. However, recent work specimen, and gold or carbon coating of the specimens (Hayward et al. 1997) indicates the high survivability of for scanning electron microscopy (SEM) was prohibited. bird eggs in marine environments, despite the rigours of the This meant that analyses were constrained to non- recruitment process. Hayward et al. (1997) report eggs even destructive or non-invasive scanning methods, and surviving uncracked at depths in excess of 500 m. Evans scanning electron microscopy in particular required low Downloaded by [JJ Liston] at 14:10 24 July 2013 (2012) went further and also explored the implications of vacuum conditions. floating eggs as a mechanism for gene flow for terrestrial A variety of analyses were thus carried out. First, the egg-laying vertebrates between land masses. This suggests Vale of Leven Hospital helical medical computerised that a number of mechanisms exist to effectively introduce tomography (CT) scanner was used, significantly improving an amniote egg to the marine environment with a real on the data previously obtained at the Western Infirmary’s possibility of preservation (Figure 1). facility (Liston and McJury 2003). The resulting data were In 1995, NHMUK PV R2903 was selected for a series then processed using Voxar 3D software (Toshiba Medical of tests being carried out on fossil eggs at Glasgow Visualization Systems). Secondary electron images were University, using a variety of available scanning obtained using an FEI 200F field-emission environmental technologies for non-invasive analysis (Liston and McJury SEM (FEG-ESEM) at the University of Glasgow’s School 2003). These revealed the internal density contrasts of the of Geographical and Earth Sciences. The microscope was object, and appeared to show a discrete mass of operated at 20 kV, using a moderately high-beam current components within the object. Unexpectedly, the Oxford (not quantified on this instrument), and at a vacuum of Clay ‘egg’ produced the most promising results in terms of ,50 Pa. Under these ‘low vacuum’ conditions, the negative an object that might actually contain fossilised remains charge that accumulates on the sample surface is (Liston and McJury 2003). Given this result, and the global compensated by water vapour in the chamber, so that scarcity of Jurassic eggs (e.g. of 135 dinosaur egg sites high-resolution images could be obtained from the sample globally, only 13 or 9% are Jurassic in age, Weishampel and without the need for a conductive carbon or gold et al. 2004; Paik et al. 2012), we considered it important to coating (e.g. Flude et al. 2012). Historical Biology 3

The synchrotron imaging system at the European pattern of lines radiate from the depressed centre of the Synchrotron Radiation Facility (ESRF, Grenoble, France) surface. The outer perimeter of the specimen exhibits was used, unfortunately causing severe fire damage to the variable thickness and in some areas, it can be seen that the specimen after markers added to the surface combusted. putative ‘eggshell’ layer extends onto this surface from the This, however, produced detached pieces of ‘shell’ for convex side. X-ray diffraction (XRD), high-magnification SEM and Edward Thurlow Leeds (the second of Alfred Leeds’ thin-sectioning work. For XRD, an INEL five sons) noted that ammonites immediately above the Inc.-manufactured curved position-sensitive detector (cali- Gryphaea zone (some 3 m above the Cornbrash) in the brated with silicon and silver behenate standards) was used to Oxford Clay around Peterborough occur as large speci- collect diffraction patterns from samples powdered and mens in concretions with a thin coating of decayed shell or loaded onto a single-crystal quartz substrate. With an output with a rough coat of pyrite, and are rarely well preserved of 4096 digital channels representing an arc of 1208,the (Leeds 1956, p. 79). He also noted that the higher zones of simultaneous measurement of diffracted X-ray intensities at the Oxford Clay, as exposed in the Eye Pit, were richly all angles of 2u across 1208 was possible. A second phase of productive in pyritised ammonites, whereas its lower zones SEM work was finally conducted at the Natural History only yielded flattened shells (Leeds 1956, p. 79), not unlike Museum (London) using a Carl Zeiss Leo 1455-VP. 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. Description The greyish brown specimen measures 125.96 mm long, 108.22 mm wide and 33.32 mm high, and weighs ,980 g. Results The convex face of the specimen (Figure 2(a)) shows a CT scan cracked face, which looks like a mosaic of small fragments Colour texturing was applied to the different densities of ‘eggshell’ that individually curl slightly outward from the determined by the Vale of Leven helical CT scanner using surface of the object at their edges, revealing an apparent Voxar 3D software (Figure 3(a)–(c)), which revealed that thickness of 0.457 mm. The clay matrix supporting the mosaic of fragments appears to be continuous; therefore, the the ‘egg’ structure was partial, and confirmed not only that cracking of the surface may be due to desiccation prior to part of an ammonite comprised the object’s concave recruitment and burial. The cracks are filled with sediment surface, but also that fragments of a belemnite were that binds the shell-like fragments together. This surface has situated within the object. Clarifying the physical structure been consolidated with a varnish. Shell-like fragments on of the specimen allowed appropriate areas to be selected the convex surface over-ride the central region of the object, for subsequent SEM analysis. indicating lithostatic compaction. The concave surface (Figure 2(b)) is primarily smooth and lacks the ‘eggshell-like’ appearance of the opposite X-ray diffraction

Downloaded by [JJ Liston] at 14:10 24 July 2013 convex surface. Molluscan shell fragments, small bivalve The results (Figure 4) show that the ‘eggshell’ sample shells and sediment with impressions of ammonite whorls taken from the convex surface of the object consists of cover portions of this surface of the object. Some whorl calcite; however, the shift in the main peak position impressions are covered with a nacreous layer of shell. A 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 fire damage. Scale bar ¼ 10 cm. Copyright image resources: NHM. 4 J.J. Liston and S.D. Chapman

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 fire 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 magnification oblique SEM surface analysis indicated a triple-laminated structure (Figure 6(a)) of around 400 mm thick. Subsequent higher magnification radial sections from samples revealed the Downloaded by [JJ Liston] at 14:10 24 July 2013

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 Figure 5. Graphical results from synchrotron, showing distinct character of the three components: the ammonite that constitutes surface contains aragonite and calcite, but this is nearly much of the concave surface, the fragments of the belemnite pure calcite with no apparent substitution of Ca by Mg. within and the third component that constitutes the convex surface. Historical Biology 5

Figure 6. Scanning electron micrographs of the putative eggshell of NHMUK PV R2903. (a) Oblique view of the surface, showing layered structure. Scale ¼ 1.0 mm. (b) Radial view showing crystalline layer. Scale bar ¼ 10 mm. (c) Surface view showing ‘pore’. Scale bar ¼ 100 mm.

presence of a radially orientated crystalline layer around would be consistent with the partial remains of one half of 45 mm thick (Figure 6(b)) with individual prisms of this an egg. Although it remains somewhat problematic to layer possessing a width of around 2.5 mm, the rest being model the process, by which a partial (rather than a obscured by clay matrix. Higher magnification SEM of the complete) egg would be preserved on the seafloor, this outer surface also showed sub-circular and oval pore-like result immediately falsifies the hypothesis that NHMUK openings ranging from 9 to 27 mm in diameter (Figure PV R2903 represents the ‘first complete’ fossil egg. 6(c)). Observed gaps between these structures ranged Given that belemnites are common throughout the between 157 and 211 mm. Oxford Clay, as also are accumulations of ammonites Radial petrographic thin sections show that clay lying on top of each other (JJL, pers. obs.), it is not adheres to the eroded outer surface (similar to Carpenter surprising that such objects would have ended up lying in 1999, Figure AII.26), and the inner surface exhibits a an upturned bowl-like structure, such as the third secondary calcite development. Two distinct layers are component appears to be, on the seabed. visible beneath the clay. High power (Figure 7(b)) and cross The regular curvature and apparent uniformity of polars (Figure 7(c)) reveal a lack of any internal structure. thickness would seem to mitigate against the third component resulting from secondary concretionary growth. So also does the outward curling of the edges of Discussion the putative shell fragments, which might be a sign of pre- From the distinct XRD peak discriminating the convex depositional dessication prior to recruitment and burial, or surface from that of the ammonite shell of the concave a result from drying immediately subsequent to exca- surface, and reconstruction of the CT and synchrotron vation, which has been fixed by varnish application. scans (Figures 3 and 5), NHMUK PV R2903 has been A conclusive result that the third component represents

Downloaded by [JJ Liston] at 14:10 24 July 2013 shown to consist of three separate components. As the remains of part of an egg would have required revealed by the synchrotron imaging system, NHMUK PV identifiable eggshell micro- or ultrastructure. However, R2903 represents an unusual assemblage of ammonite, neither could be identified with certainty. The low belemnite and a third component. The nature of this third magnification SEM showed 400 mm thickness, which component remains uncertain, although its regular shape 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) ¼ 100 mm. 6 J.J. Liston and S.D. Chapman

Hirsch (1994) and Ensom (2002), and apparently lacking distribution and low density of angusticanaliculate pores ornamentation commonly described for many avian and in eggshell, none were visible in radial section. However, other dinosaurian eggshells (Hirsch and Packard 1987). these structures are consistent in size with the range However, the low magnification renders this image expected for angusticanaliculate pores, and are difficult to ultimately unhelpful. When SEM was repeated on samples explain as a result of dissolution or other diagenetic at higher magnification, the only layer detectable was a alterations. crystalline layer with radially orientated prisms. The Mikhailov (1997) notes angusticanaliculate pores as 45 mm thickness of this layer is much less than what would indicative of a dry terrestrial incubating environment, and be expected in a prismatic layer in eggshell, and the furthermore notes their occurrence in dinosauroid- 2.5 mm width of individual prisms is similarly significantly prismatic and ornithoid basic shell types, namely the less than the 10–20 mm range measured by Mikhailov general egg parataxa assigned to Theropoda (Prismatoo- (1997, plate 8) and Ensom (2002, plate 3, figure 8). As lithidae and Elongatoolithidae) and Enantiornithes (Lae- both these measurements are outside the ranges observed visoolithidae) (Mikhailov 1997, p. 39, Table 1; Carpenter for biological calcite in eggshell, it seems most likely that 1999). He further noted that prismatoolithid eggshell this is a thin isolated pocket of post-depositional calcite appeared to be the only dinosaurian eggshell type that was growth. common prior to the Cretaceous (Mikhailov 1997, p. 59). In contrast, the petrographic thin sections gave a whole These observations are consistent with this specimen profile of the putative eggshell with two clear layers coming from Middle Jurassic clays that have yielded many visible: a well-demarcated inner layer and an outer heavily dinosaur remains (Noe´ et al. 2010). However, it must be eroded layer. The outer layer showed no sign of prismatic stressed that the lower zone thicknesses exhibited by these structure, although this could be because of recrystallisa- general parataxa (Prismatoolithidae circa 200 mm, Elon- tion, consistent with the observed secondary calcite gatoolithidae 170–300 mm, Laevisoolithidae 125– deposits. To interpret this section as eggshell, the internal 200 mm, all data from Mikhailov 1997) as well as the layer would represent an unusually thin (,70 mm) lower prism widths (10–20 mm range for these three groups, zone or mammillary layer with the outer layer representing again all data from Mikhailov 1997) are considerably in the eroded remains of the prismatic layer after erosion of excess of the figures of 70 mm (putative lower zone the external layer. But with this outer layer lacking any thickness) and 2.5 mm (putative prism width, if the radial clearly definable outer perimeter and hence an absence of crystalline layer is not, in fact, merely secondary calcite any possible surface ornamentation on an external layer, it growth) that the third component of NHMUK PV R2903 is hard to identify that this layer indeed represents the exhibits. These numerical departures must, therefore, raise prismatic layer. The absence of a clear perimeter also some questions over this interpretation, and in particular prohibits the measurement of a regular thickness. raise the issue that the layer visible under SEM might The structures resembling round-to-oval pore openings simply represent geologically rather than biologically were particularly narrow in diameter in terms of pores directed calcite growth. observed in eggshell (Mikhailov 1997), but are consistent In summary, the lack of a visible complete undamaged in both their shape and dimensions (within the range of section with all layers measurable is problematic. The Downloaded by [JJ Liston] at 14:10 24 July 2013 10–100 mm) with the openings of angusticanaliculate eroded outer surface, in particular, means that any external pores (Mikhailov 1997). However, no canals were visible ornamentation present that would help with identification is in radial thin section to determine their nature beneath also absent. The thicknesses recorded for the layers and the openings (e.g. straight, as would be expected if these prisms are significantly less than the recognised ranges for were angusticanaliculate pores). The sparse distribution of known eggs. But, given the overall regular 3D shape, and in these structures, with observed gaps of 157–211 mm, also the absence of another means by which to explain the pore- recalls the observed infrequency of angusticanaliculate like openings, it still seems simplest currently to regard pores on the surface of eggshells (Mikhailov 1997). NHMUK PV R2903 as possibly representing a fossilised Although estimates based on assuming that this spacing is partial egg containing parts of an ammonite and a regular make these structures appear significantly more belemnite. It is hoped that further and more extensive common than the 3–20 pores per 100 mm2 quoted by sampling of the specimen can be permitted at some point to Mikhailov (1997), perhaps by as much as a factor of a conclusively determine the nature of this third component. 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 Conclusions with the overall 3D shape, remain the most compelling Although the ultrastructure and microstructure could not arguments for the third component of NHMUK PV R2903 be identified with certainty, NHMUK PV R2903 may to be interpreted as representing the remains of part of an indeed represent a recrystallised and damaged fossilised egg. Unsurprisingly, given the recorded infrequent partial dinosaur egg with an angusticanaliculate pore Historical Biology 7

system. It has not yet been possible to disprove Alfred Eggs and Babies. Cambridge: Cambridge University Press. Chapter Leeds’ judgement of one of his specimens as the ‘egg of a 1. p. 15–30. Ensom PC. 2002. Reptile eggshell, tiny vertebrate remains and globular saurian’. This is remarkable, given that by the end of the calcified cartilage from the Purbeck Limestone Group of southern nineteenth century, fossil eggshell work was effectively England. In: Milner AR, Batten DJ, editors. Life and environments in unknown, and a testament to both his skills as a collector Purbeck times. Spec Pap Palaeontol. 68: 221–239. Evans T. 2012. Oceanic transport of calcified eggs: taphonomic and and as an observer in the field. biogeographic implications. Hist Biol Int J Paleobiol. 24(5):467–487. Flude S, Lee MR, Sherlock SC, Kelley SP. 2012. Cryptic microtextures Acknowledgements and geological histories of K-rich alkali feldspars revealed by charge contrast imaging. Contrib Mineral Petrol. 163:983–994. The authors thank both reviewers for much assistance in Hayward JL, Folsom SD, Elmendorf DL, Tambrini AA, Cowles DL. improving the text, Frankie Jackson and Colin Shaw (Figure 1997. Experiments on the taphonomy of Amniote eggs in marine 7(b),(c)), Montana State University, for providing thin-section environments. PALAIOS. 12(5):482–488. images; Gordon Cressey and Peter Tandy, Department of Earth Hirsch KF. 1994. Upper Jurassic eggshells from the Western Interior of Sciences (Mineralogy), NHM, London, for the XRD analysis North America. In: Carpenter K, Hirsch KF, Horner JR, editors. (Figure 4); Tony Wighton, NHM, London, for thin sections Dinosaur eggs and babies. Cambridge: Cambridge University Press. (Figure 7(a)); Phil Crabb, Photographic Studio, NHM, London Chapter 10. p. 137–150. for images of the Leeds egg; Vincent Fernandez for SYNCH- Hirsch KF, Packard MJ. 1987. Review of fossil eggs and their shell ROTRON access; Steve Baker, Department of Earth Sciences structure. Scanning Microscopy. 1:383–400. Leeds ET. 1956. The Leeds collection of fossil reptiles from the Oxford (Palaeontology), NHM, London, for providing access to the Clay of Peterborough. Oxford: Blackwell. collections and help identifying the ammonite; Kirsty Ross, Liston JJ. 2006. From Glasgow to the Star Pit and Stuttgart: a short Senior Radiographer at the Western Infirmary and Vale of Leven journey around the world’s longest fish. Glasgow Nat. 24(4):59–71. Hospitals (Glasgow), for her infinite patience in steadfastly Liston JJ, McJury M. 2003. Egg candling for the 21st century: the use of piloting CT scanners through the weird fossil remains that JJL three dimensional digital imaging technology to investigate the has for far too many years been bringing her; Caroline Smith and contents of fossilised eggs. Q J Dinosaur Soc. 4(3):3–8. Martin Lee (Figure 6(a)) for steering the Lo-Vac SEM in the Liston JJ, Newbrey MG, Challands TJ, Adams CE. 2013. Growth, age and University of Glasgow’s Gregory Building; Bobby Davey for size of the Jurassic pachycormid Leedsichthys problematicus processing the DICOM scans into ‘glorious Technicolor’ using (Osteichthyes: Actinopterygii). In: Arratia G, Schultze H-P, Wilson the Voxar 3D software package; Bob Nicholls for fantastic MVH, editors. Mesozoic Fishes 5 – global diversity and evolution. eleventh hour ‘Pteroportation’ work at www.paleocreations.com; Mu¨nchen: Verlag Dr. F. Pfeil. p. 145–175. Lydekker R. 1889. On the remains and affinities of five genera of Ian Rolfe for selflessly transporting a fossil egg from London to Mesozoic reptiles. Q J Geol Soc London. 45:41–59. Glasgow, and Eric Buffetaut for doing the same from London to Mikhailov KE. 1997. Fossil and recent eggshell in amniotic vertebrates: Grenoble and Cambridge; and Paul Taylor and Alex Brown fine structure, comparative morphology and classification. Spec Pap (NHM) for taking SEMs (Figure 6(b),(c)) and discussion. We Palaeontol. 56:1–80. also thank Neil Landman and Al McGowan for helpful molluscan Noe´ LF, Liston JJ, Chapman SD. 2010. ‘Old bones, dry subject’: the assistance. Mike A. Taylor suggested the rotting maternal corpse dinosaurs and pterosaur collected by Alfred Nicholson Leeds of as a means of facilitating egg recruitment into the marine realm. Peterborough, England. In: Moody RTJ, Buffetaut E, Naish DW, Martill DM, editors. Dinosaurs and other extinct Saurians: a historical perspective. Special publications. Vol. 343. London: Geological Society. p. 49–77. References Paik IS, Kim HJ, Huh M. 2012. Dinosaur egg deposits in the Cretaceous Andrews CW. 1910. A descriptive catalogue of the marine reptiles of the Gyeongsang Supergroup, Korea: diversity and paleobiological implications. J Asian Earth Sci. 56:135–146.

Downloaded by [JJ Liston] at 14:10 24 July 2013 Oxford Clay – based on the Leeds collection in the British Museum (Natural History), London, Part I. London: British Museum (Natural Seeley HG. 1889. Note on the pelvis of Ornithopsis. Q J Geol Soc History). London. 45:391–397. Andrews CW. 1913. A descriptive catalogue of the marine reptiles of the Smith WA. 1889. On the palaeontology of sturgeons. Proc Geol Assoc. Oxford Clay – based on the Leeds collection in the British Museum 11:24–44. (Natural History), London, Part II. London: British Museum (Natural Swinton WE. 1950. Fossil eggs from Tanganyika. The Illustrated London History). News. p. 1082–1083. A fossil reptile’s eggs unearthed in England. 1923 London: The Sphere. Van Straelen V. 1928. Les Oeufs de reptiles fossiles. Palaeobiologica. p. 196. 1:295–312. Buffetaut E, Le Loeuff J. 1994. The discovery of dinosaur eggshells in Weishampel DB, Barrett PM, Coria RA, Loeuff JL, Xing X, Xijin Z, nineteenth-century France. In: Carpenter K, Hirsch KF, Horner JR, Sahni A, Gomani EMP, Noto CR. 2004. Dinosaur distribution. In: editors. Dinosaur eggs and babies. Cambridge: Cambridge Weishampel DB, Dodson P, Osmo´lska H, editors. The Dinosauria. University Press. Chapter 2. p. 31–34. Berkeley, CA: University of California Press. p. 517–606. Carpenter K. 1999. Eggs, nests, and baby dinosaurs: a look at dinosaur Young MT, De Andrade MB, Brusatte SL, Sakamoto M, Liston JJ. 2013. reproduction (life of the past). Bloomington, IL: Indiana University The oldest known metriorhynchid super-predator: a new genus and Press. species from the Middle Jurassic of England, with implications for Carpenter K, Alf K. 1994. Global distribution of dinosaur eggs, nests, and serration and mandibular evolution in predacious clades. J Syst babies. In: Carpenter K, Hirsch KF, Horner JR, editors. Dinosaur Palaeontol. 11(4):475–513. DOI:10.1080/14772019.2012.704948