S Largest Toothed Pterosaur, NHMUK R481, an Incomplete Rostrum of Coloborhynchus Capito (Seeley, 1870) from the Cambridge Greensand of England

Cretaceous Research 34 (2012) 1e9

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Cretaceous Research

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The world’s largest toothed pterosaur, NHMUK R481, an incomplete rostrum of Coloborhynchus capito (Seeley, 1870) from the Cambridge Greensand of England

David M. Martill a,*, David M. Unwin b a Palaeobiology Research Group, School of Earth and Environmental Sciences, University of Portsmouth, Portsmouth PO1 3QL, UK b School of Museum Studies, University of Leicester, 19 University Road, Leicester LE1 7RF, UK article info abstract

Article history: The assignment of a fragment of the anterior tip of a pterosaur rostrum from the Cenomanian Cambridge Received 1 June 2011 Greensand of eastern England to the ornithocheirid Coloborhynchus capito (Seeley, 1870) is conﬁrmed. Accepted in revised form 6 September 2011 The fragment represents partial left and right fused premaxillae and retains broken teeth within alveoli. Available online 29 September 2011 A width across the palate of 57.4 mm, a height at the anterior rostrum in excess of 95 mm and a tooth with a diameter of 13 mm at the base of the crown indicates a remarkably large individual, tentatively Keywords: estimated to have had a skull length in excess of 0.75 m and a wing span of up to 7 m. This fragment Pterosauria represents the largest toothed pterosaur yet reported. This ﬁnd, and several other large postcranial Coloborhynchus Cretaceous fragments from the Cambridge Greensand, suggest that ornithocheirids, toothed ornithocheiroids known e England from the earliest Early to early Late Cretaceous (Valanginian Cenomanian) achieved very large, but not Evolution giant size. Pteranodontids, edentulous ornithocheiroids currently known only from the mid Upper Gigantism Cretaceous (Coniacianeearly Campanian), reached similar dimensions, up to 7.25 m in wing span. Contrary to popular myth, however, ornithocheiroids did not attain the giant sizes (wing spans of 10 m or more) achieved by azhdarchids in the late Late Cretaceous (CampanianeMaastrichtian). Ó 2011 Elsevier Ltd. All rights reserved.

1. Introduction undertaken by Hooley (1914) and, more recently, the entire pterosaur assemblage was revised by Unwin (1991, 2001) who The Cretaceous Cambridge Greensand of eastern England was confirmed the presence of the ornithocheirids Anhanguera, excavated from the mid to late 19th century as a source of phos- Coloborhynchus and Ornithocheirus, as well as the lonchodectid phate for agricultural fertiliser (Grove, 1976), during which period Lonchodectes and the indeterminate edentulous pterosaur it yielded many fossils including vertebrates (Seeley, 1969). These Ornithostoma. almost exclusively fragmentary remains were derived from Despite the fragmentary nature of the Cambridge Greensand underlying Albian strata based on the associated derived pterosaur fossils, Owen recognised that some fragments repre- (reworked) invertebrate assemblage (Unwin, 2001). Among the sented individuals of considerable size and, for one taxon, “Pter- various fossils collected, which represent fish, ichthyosaurs, odactylus” [¼Coloborhynchus] sedgwickii, he estimated a wing span plesiosaurs, crocodiles, turtles and dinosaurs (including birds), of 22 ft (w6.55 m) (Owen, 1859a). Here we describe NHMUK R481, were many hundreds of bone fragments of pterosaurs. These a fragmentary pterosaur rostrum with teeth from the Cambridge pterosaur remains were first described by Owen (1859a, b, 1860, Greensand in the collections of the Natural History Museum, 1861) and later by Seeley in more than 20 publications spanning London. The specimen, which was first mentioned, but not figured the interval 1864 to 1901 (listed in Unwin, 2001). The most or described, by Unwin (2001), is incomplete and highly damaged: important among these were the “Index” published in 1869 and a combination of ancient reworking and recent phosphate digging. “The Ornithosauria”, which appeared in 1870. Owen erected four None the less it is noteworthy because it represents the largest new species and Seeley added many more, most of which have ornithocheirid found to date and indicates a possible upper limit to proved invalid for various reasons (see Unwin, 2001; Table 1). A the size achieved by toothed pterosaurs, thereby adding to our detailed review of the Cambridge Greensand pterosaurs was understanding of pterosaur evolution in the Cretaceous.

The following collection abbreviations are used: CAMSM, Sedgwick * Corresponding author. Museum of Earth Sciences, Cambridge, UK; CSRLV, Centro Studi e E-mail address: [email protected] (D.M. Martill). Ricerche Ligabue, Venice, Italy; IMCF, Iwaki Museum of Coal and

Table 1 3. Systematic palaeontology Metric data (in millimetres) for Coloborhynchus robustus (NSM-PV 19892, RGM 401 880) and Coloborhynchus capito (NHMUK R481, CAMSM B 54625). Abbreviations: HL, Pterosauria Kaup, 1834 humerus length; RD, depth of anterior rostral facia; RW, width of anterior rostral facia; SL, skull length; TD, maximum diameter of tooth in dental alveolus # 2; WS, Pterodactyloidea Plieninger, 1901 wing span. Data from Unwin (1991), Kellner and Tomida (2000) and Veldmeijer Ornithocheiroidea Seeley, 1891 (2003). Estimate based on restoration in Fig. 4 shown in parentheses. Ornithocheiridae Seeley, 1870 RW RD TD SL HL WS Genus Coloborhynchus Owen, 1874 NHMUK R481 57.4 (60) 56 13.6 eee CAMSM B 54625 48 44 13.0 eee Junior synonyms. Amblydectes Hooley, 1914; Siroccopteryx Mader NSM-PV 19892 25 21 8.5 616 257 4700 and Kellner, 1999; Tropeognathus (partim) Wellnhofer, 1987; a RGM 401 880 36 35 9.6 712 290 5300e5800 Uktenadactylus Rodrigues and Kellner, 2008. a Estimate based on comparison with NSM-PV 19892 (Kellner and Tomida, 2000) and SMNK 1132PAL (Frey and Martill, 1994). Type species. Coloborhynchus clavirostris Owen, 1874.

Fossils, Iwaki, Japan; NHMUK, Natural History Museum, Temporal range. Valanginian to Late Albian (Fastnacht, 2001). London, UK; NSM, National Science Museum, Tokyo, Japan; RGM, National Natuurhistorisch Museum, Leiden, Netherlands; SMNK, Staatliches Museum für Naturkunde, Karlsruhe, Germany; YORM, Yorkshire Museum, York, UK.

2. Locality and stratigraphy

The Cambridge Greensand is a thin (w0.6 m) remané deposit occurring at the base of the Chalk Formation in eastern England around the City of Cambridge (Fig. 1). It was extensively excavated between Barrington and Horningsea, Cambridgeshire in the 19th century (Grove,1976) but, with the exception of Barrington Chalk Pit (Mortimore et al., 2001), there are few exposures of this horizon available today. Although the deposit itself is generally considered to be of late Cretaceous, Cenomanian age, its macrofossil assemblage is thought to be derived largely from the underlying early Cretaceous, Albian, Gault Formation, on the basis of derived ammonites associated with the vertebrate fossils (e.g. Spath, 1923e1943; Owen, 1979; Morter and Wood, 1983). An early Cenomanian age for deposition of the Cambridge Greensand is based on an apparently autochthonous foraminiferan assemblage (Hart, 1973), but Morter and Wood (1983) could not rule out the possibility of a very late Albian age due to the presence of the bivalve Aucinella. The age of the derived vertebrates is now considered to be no older than the Cal- lihoplites auritus ammonite subzone and most likely no younger than upper Stoliczkaia dispar Zone (Cooper and Kennedy, 1977), and they are thus of late Albian age (Unwin, 2001; Fig. 2).

Fig. 2. Stratigraphic column for the AlbianeCenomanian (Early/Late Cretaceous) of Fig. 1. Outcrop map of the Gault Clay and Upper Greensand in Eastern England. The Eastern England (modiﬁed from Unwin, 2001). Cambridge Greensand Member fossils vertebrate-bearing Cambridge Greensand occurs only in the narrow region between come from hiatus phosphate horizons (indicated by pebble stipple) within the Cen- the arrows. CGS, Cambridge Greensand Member. Based on Mortimore et al. (2001). omanian, but are of Albian age. ?, uncertainty of zonal boundary. D.M. Martill, D.M. Unwin / Cretaceous Research 34 (2012) 1e9 3

Geographic range. England, Texas, Brazil, North Africa, and perhaps As preserved NHMUK R481 is 95 mm high, and 38 mm long Mongolia (Owen, 1874; Lee, 1994; Mader and Kellner, 1999; Unwin with a width of 57.4 mm (Fig. 3AeG). The restored width across the and Bakhurina, 2000; Fastnacht, 2001; Veldmeijer, 2003; Rodrigues anterior palate is approximately 60 mm (Fig. 4). In anterior aspect and Kellner, 2008). the specimen comprises a pair of fused premaxillae with a rounded triangular, boss-like termination within which are located alveoli Coloborhynchus capito Seeley, 1870. for the first pair of teeth. The rostral boss is composed of the dorsally reflected palate (“anterior rostral facies” of Fastnacht, Synonym. Coloborhynchus sp. indet. Martill, 2010, p. 307, fig. 14. 2001) and is approximately 56 mm high with an estimated width of 60 mm at its base. The right alveolus of the first tooth pair Description. The specimen described here is a highly worn fragment of contains a broken tooth with a circular cross-section and a diam- anterior rostrum with broken teeth and is deposited in the collection eter at the crown root junction of w8 mm. The left alveolus is of the Natural History Museum, London, specimen number NHMUK empty with highly abraded margins, but appears to have a long axis R481 (Figs. 3 and 4). It comprises a short length of the anteriormost extending dorsocaudally into the premaxilla. The root of the second portion of the premaxillae. The broken caudal margin is highly tooth of the right side is preserved within its alveolus and has abraded, as is the palatal surface. The lateral margins are entire, a diameter of 13 mm at the crown root junction. On the left side the although somewhat abraded in places, and there is a small oyster alveolus for this tooth has been abraded and the alveolar wall has adherent to the left lateral face (x on Fig. 3B). The anterior surface is broken through to the adjacent alveolus for the first tooth, forming abraded, and the bony compacta has been removed. Two teeth are a large cavity. Although the compacta is missing from the surface of visible in cross-section on this surface, as is a broken and worn the anterior part of the premaxillae, the exposed surface is vertical alveolus for a large tooth. The dorsal surface is broken and reveals an with respect to the remainder of the palate, and the flexion appears internal trabeculum which curves anterodorsally. Where the com- to be abrupt. The first pair of teeth would have projected anteriorly pacta is removed inter-trabecular spaces are filled with buff-coloured and somewhat ventrally while the second pair projected more phosphatic sediment typical of Cambridge Greensand fossils. anteroventrally.

Fig. 3. Fragment of rostrum of ornithocheirid pterosaur, NHMUK R481, referred to Coloborhynchus capito. A, anterior view; B, left lateral view; C, right lateral view; D, caudal view; E, close up of alveolus with broken tooth; F, dorsal view; G, palatal view. Scale bar AeD, FeG, 50 mm; E, 10 mm. 4 D.M. Martill, D.M. Unwin / Cretaceous Research 34 (2012) 1e9

the palate, and neither is there any hint of an inflated central portion, suggesting the palate was flat in this region. Both lateral faces have compacta preserved, both facies converge dorsally until they become sub-parallel forming the base of a crest, the dorsal portion of which is now missing. The lateral dorsal surface of the rostrum reveals an internal, perforated septum of bone that is vertical, becoming increasingly deflected anteriorly until it fuses with the antero-dorsal margin of the premaxillae.

4. Taxonomic assignment of NHMUK R481

The pterosaurian nature of NHMUK R481 is demonstrated by several features, principally the presence of remarkably thin compacta with a smooth external surface. Teeth with a slightly ribbed root and a sub-circular cross-section are typical for pterosaurs and the strong similarity of the specimen to the rostra of ornithocheirids such as Coloborhynchus clavirostris (Owen, 1874) and Coloborhynchus robustus (Fastnacht, 2001) confirm the pterosaurian identification. Among pterosaurs the presence of a reflected palatal surface resulting in a ‘blunt-snout’ condition is restricted to some, but not all, ornithocheirids. This, the remarkably large size of the rostrum, its relative breadth, and the size, spacing and orien- tation of the teeth are all diagnostic features of Ornithocheiridae (e.g. Unwin, 2003, p. 178). The presence of a reflected palatal surface bearing the first pair of dental alveoli and bounded on its ventrolateral corners by the second pair, is observed in Anhanguera and Coloborhynchus. In the former the degree of reflection is low (e.g. Wellnhofer, 1991, figs. 2e3), whereas in the latter the palatal surface is reflected vertically resulting in a distinctive truncated snout (Owen, 1874; Lee, 1994; Mader and Kellner, 1999; Figs. 5, 6 and 8), as observed in NHMUK R481. Several species of Coloborhynchus have been named and some of these have been assigned to distinct genera (Rodrigues and Kellner, 2008). Here we adopt a conservative approach including all Coloborhynchus-like taxa in a single genus, Coloborhynchus, and, pending a full taxonomic revision (Unwin in prep), recognise the following species: Coloborhynchus sedgwickii (Owen, 1859a); Colo-

Fig. 4. Annotated photograph of anterior aspect of NHMUK R481 highlighting the posi- borhynchus capito (Seeley, 1870); C. clavirostris Owen, 1874; tion of the dental alveoli and the border of the anterior face of the rostrum defined by a 90 C. robustus (Wellnhofer, 1987); Uktenadactylus (¼ Coloborhynchus) dorsal reflection of the palate. The dotted white line, continuous white line and black line wadleighi Lee, 1994; and Siroccopteryx (¼ Coloborhynchus) moroc- outline the shape of the anterior boss. censis Mader and Kellner, 1999. In addition, Anhanguera piscator Kellner and Tomida, 2000 and Coloborhynchus spielbergi Veldmeijer, The palatal surface of the specimen is damaged and little bone 2003 are both considered here to be junior synonyms of C. robustus. compacta remains. Poorly defined alveoli for the second pair of C. capito is represented by several fragments of the rostrum teeth are visible, as is the empty and abraded left alveolus of the including CAMSM B 54625 (the holotype), YORM 193/131F and first tooth pair. There is no hint of a ridge or groove on the surface of NHMUK R480. This species is distinguished by the presence, on the

Fig. 5. Rostrum of Coloborhynchus clavirostris Owen, 1874 (Holotype NHMUK R1822) from the Wealden Supergroup of East Sussex, southeast England. A, anterior aspect showing four anteriorly directed dental alveoli, the more ventral pair of which is located ventrolaterally; B, left lateral aspect showing anterior teeth located laterally. Scale bar, 10 mm. D.M. Martill, D.M. Unwin / Cretaceous Research 34 (2012) 1e9 5

Fig. 6. NHMUK R481 compared with holotype (CAMSN B54.625) of Coloborhynchus capito (Seeley, 1870). A, sketch of holotype of C. capito in anterior aspect (modiﬁed from Unwin, 2001); B, outline of holotype of C. capito in anterior aspect with reversed image to determine shape and size of the terminal part of the rostrum; C, NHMUK R481 in anterior aspect with location of ﬁrst two tooth pairs indicated. The dashed line is the approximated continuation of the left lateral border of the anterior rostral facies. Scale bar, 10 mm.

tip of the rostrum, of a large sagittal crest with an anterior surface 1859a; Fig. 8G). Unwin (2001) has suggested that fossils currently that has a markedly concave profile in lateral view (Fig. 8C). This assigned to C. capito and C. sedgwickii may be sexual dimorphs of feature is present in NHMUK R481. The holotype of C. capito a single species as the only difference between these two taxa (CAMSM B 54625) is a worn rostrum fragment bearing five alveoli, appears to be the presence of the anterior rostral crest in C. capito. including the characteristic first alveolus on the rostral face of the This hypothesis has recently been strengthened by the demon- palate. In anterior aspect the dorsal surface of the anterior rostral stration of sexual dimorphism in Darwinopterus (Lü et al., 2011), but facies is preserved and, using this surface as a point of reference for more complete specimens of C. capito and C. sedgwickii are required the median line, the cross-sectional shape of the anterior rostrum before this can be inferred for these species. was reconstructed by mirror imaging the specimen (Fig. 6B), NHMUK R481 differs from C. clavirostris in lacking the laterally thereby facilitating comparison with NHMUK R481. This shows that placed anterior teeth and the medially inflated palate, most strik- apart from a marked difference in size these individuals are ingly seen in lateral view (Fig. 5B). In C. wadleighi the anterior boss remarkably similar and undoubtedly conspecific. This assignment is of the premaxillae is more pronounced (Fig. 8D) and elongate supported by the similarity in proportions of the specimens, as rather than abbreviated as it is in NHMUK R481, and the same is shown by a simple scatter plot of depth vs width of the rostral tips true for Coloborhynchus moroccensis (Fig. 8A). for various ornithocheirids (Fig. 7). C. robustus Fastnacht, 2001 is known from substantial, well- The presence of a well-developed rostral crest clearly distin- preserved material from the Santana Formation of Brazil guishes NHMUK R481 from C. sedgwickii (Owen, 1859a), which (Wellnhofer, 1987; Kellner and Tomida, 2000; Fastnacht, 2001; lacks such a structure. Thus NHMUK R481 is more than twice as Veldmeijer, 2003)(Fig. 8E). In this species the anterior palate is deep as it is wide at the second tooth pair, whereas rostra of turned dorsally at an angle of about 45 increasing to nearly 90 at C. sedgwickii are of similar width and depth at this point (Owen, its upper extremity (Fig. 8E, H, I). By contrast, in NHMUK R481, the 6 D.M. Martill, D.M. Unwin / Cretaceous Research 34 (2012) 1e9

conservative than variation in the breadth and depth of the rostrum. The rostrum of CAMSM B 53625 is only 78% the size of NHMUK R481, but the maximum diameter of the teeth in the second pair of dental alveoli is much more similar (Table 1). Furthermore, teeth of comparable dimensions have also been reported in C. robustus (SMNK 2302PAL, Fastnacht, 2001) and Liao- ningopterus gui (Wang and Zhou, 2003), although in both these cases the rostrum is considerably smaller than that of NMHUK R481. Thus, while the latter is clearly larger than other known individuals of Coloborhynchus, Anhanguera and Ornithocheirus, giant size in excess of 8 m in wing span can be safely excluded. Several hundred fragmentary pterosaur postcranial bones have been recovered from the Cambridge Greensand and more than 90% of these can be assigned to Ornithocheiroidea (Unwin, 1991). The pattern of size distribution exhibited by these ornithocheiroid bones matches that seen in Fig. 7 and comparison with more complete skeletons of ornithocheirids from the Santana Formation of Brazil (Wellnhofer, 1985, 1991; Kellner and Tomida, 2000; Veldmeijer, 2003; Veldmeijer et al., 2009; Elgin and Frey, 2011) and the Jehol Biota of China (Lü and Ji, 2005; Lü, 2010) indicate that Fig. 7. Scatter plot of depth vs width of anterior rostral facies in millimetres of various most specimens represent individuals of 3e5 m in wing span. There ornithocheirid pterosaurs (data from Unwin, 1991). Symbols: diamond, Anhanguera are several teeth (Fig. 9) and fragments of humeri (CAMSM B 54053, cuvieri; square, Anhanguera fittoni; triangle, Coloborhynchus sedgwickii; circle, Colobo- B 54627) carpals (CAMSM B 54171, B 54176) and an acetabulum rhynchus robustus; star, Ornithocheirus simus; cross, Coloborhynchus capito. (CAMSM B 54253) that are comparable in size to the largest known examples of C. robustus (RGM 401 880) and presumably represent e upturned palate is vertical (Fig. 3BeC). In addition the rostral crest very large individuals of 5 6 m, or more, in wing span. The latter fi of C. robustus has a more posterior location than that of C. capito and nds may be attributable to C. capito, as this is the largest pterosaur a lower, flatter anterior profile in lateral view. in the Cambridge Greensand assemblage (Unwin, 2001), although the possibility that some (perhaps all) of them pertain to O. simus, which also achieved large sizes (Unwin, 2001), cannot be excluded. 5. The size of NHMUK R481 and its significance Prior to this study the largest known ornithocheirids included RGM 401 880, identified here as an individual of C. robustus, and It is not possible to establish the exact length of the skull or wing SMNK 1136 identified by Frey and Martill (1994) as Anhanguera sp. span for NHMUK R481 because it is too fragmentary. Nonetheless, it As discussed above, NHMUK R481 appears to represent a pterosaur is possible to gain a general idea of the size of this individual by that was considerably larger than either of these Santana Formation comparing it with the rostra of other ornithocheirids (Fig. 7), specimens and is recognised here as both the largest ornithocheirid including two relatively complete examples of C. robustus, NSM-PV and the largest toothed pterosaur yet found. NHMUK R481 is not, 19892 (Kellner and Tomida, 2000) and RGM 401 880 (Veldmeijer, however, the largest Early Cretaceous pterosaur to be found. The 2003)(Table 1). The scatter plot data falls into three distinct distal end of a remarkably large wing-phalanx one, CSRLV 12701a groups (Fig. 7). The cluster to the lower left consists of relatively (Dalla Vecchia and Ligabue, 1993), from the Santana Formation of small individuals, predominantly of Anhanguera cuvieri and Brazil represents a giant pterosaur with a wing span between 8.2 Anhanguera fittoni. A single example of C. robustus, NSM-PV 19892, is and 9.3 m. Dalla Vecchia and Ligabue (1993) were unable to resolve one of the largest pterosaurs in this cluster. Based on its wing span of the identity of this fossil beyond assignment to Pterodactyloidea. 4.7 m (Kellner and Tomida, 2000; Table 1), it can be inferred that However, following comparison with other well-preserved remains members of this cluster probably ranged up to 5 m in wing span. of ornithocheirids (Wellnhofer, 1991; Veldmeijer et al., 2009) and A more diffuse cluster located in the upper middle region of the thalassodromeids (IMCF 1052) from the Santana Formation, Pter- plot is dominated by individuals of Ornithocheirus simus and anodon (Bennett, 2001) and azhdarchoids from the Jiufotang C. sedgwickii, but also includes two examples of C. capito and a large Formation of China (e.g. Lü et al., 2006; Lü et al., 2008), it seems that individual of C. robustus (RGM 401 880). The latter is estimated to this fragment probably pertains to an azhdarchoid, rather than an have had a wing span of approximately 5.5 m (Table 1), on the basis ornithocheiroid. Large thalassodromeids have been reported from of which members of this cluster are predicted to have had wing the Santana Formation (Kellner and Campos, 2002; Veldmeijer spans of 5e6 m and in the case of the largest individuals, probably et al., 2005), thus it is most probable that CSRLV 12701a repre- well over 6 m. sents a gigantic individual of Tupuxuara or Thalassodromeus. There is a distinct gap between NHMUK R481, which lies to the Alternatively, although less likely, this fossil might represent an upper right of the plot, and the upper middle cluster. Since the azhdarchid as this clade is now recorded from the Kem Kem largest individuals of the latter were probably over 6 m in wing Formation (Ibrahim et al., 2010) which is Cenomanian, or possibly span it is reasonable to infer that NHMUK R481 was even larger, late Albian, in age (Ibrahim, 2011). with a wing span that may have reached 7 m. This size estimate This reassessment of the maximum dimensions of orni- may seem cautious, but takes into account evidence for allometric thocheirids permits a more precise understanding of the evolution growth patterns in ornithocheirids. Specimens of C. robustus of size in Cretaceous pterosaurs. With the discovery that the demonstrate that, compared to wing span, the anterior portion of holotype of Cearadactylus atrox was constructed from fragments of the rostrum experienced positive allometric growth. The rostrum an ornithocheirid (Vila Nova et al., 2010) it now appears that of RGM 401 880 is approximately 50% larger than that of NSM-PV ctenochasmatoids, which persisted until the late Early Cretaceous 19892 (Fig. 7), but the wing span of the former is less than 25% where they are represented by Pterodaustro, did not achieve wing greater than the latter (Table 1). Tooth size is also more spans in excess of 3 m. Dsungaripteroids also survived well into D.M. Martill, D.M. Unwin / Cretaceous Research 34 (2012) 1e9 7

Fig. 8. Rostra, in left lateral view and drawn to the same scale, of snub-snouted ornithocheirids compared. A, Coloborhynchus wadleighi;B,Coloborhynchus clavirostris; C, Coloborhynchus capito;D,Coloborhynchus moroccensis;E,Coloborhynchus robustus; F, NHMUK R481; G, Coloborhynchus sedgwickii;H,I,Coloborhynchus robustus. The arrows indicate the first tooth pair on the anterior rostral facies. A, redrawn from Lee (1994); B, from original specimen NHMUK R1822; C, redrawn from Unwin (2001, fig. 8a); D, redrawn from Mader and Kellner (1999, fig. 2b); E, redrawn from Fastnacht (2001, fig. 3b); F, drawn from original NHMUK R481; G, redrawn from Unwin (2001, fig. 9b); H, redrawn without occlused mandible from Kellner and Tomida (2000, fig. 5b); I, redrawn and reversed from Veldmeijer (2003, fig. 2c). 8 D.M. Martill, D.M. Unwin / Cretaceous Research 34 (2012) 1e9

Fig. 9. Two pterosaur teeth (NHMUK R35418, 35418a) from the Cambridge Greensand with basal diameters comparable to the broken tooth roots in NHMUK R481. A, root and incomplete crown (figured by Owen, 1861, pl. iv, fig. 4); B, crown missing root. Scale bar, 10 mm. the Early Cretaceous, where the largest known examples of pterosaurs was primarily a result of extrinsic environmental Dsungaripterus weii reached wing spans of 3e4 m. Among orni- factors, but rather that they reflect intrinsic anatomical, biome- thocheiroids both ornithocheirids (in the form of C. capito) and chanical or physiological constraints, the identity of which has yet pteranodontids, represented by Pteranodon, reached wing spans of to be determined. over 7 m (Bennett, 2001; this paper). Ornithocheirids achieved these maximal dimensions in the late Early Cretaceous, pter- Acknowledgements anodontids in the mid to late Late Cretaceous. Notably, neither of these lineages appear to have attained truly gigantic size (contrary We thank Sandra Chapman, Angela Milner and Lorna Steel at the to popular notions), but were contemporaneous with azhdarch- NHM, London and Matt Riley and Rob Theodore at Sedgwick Museum, oids that reached wing spans in excess of 8 m in the late Early Cambridge for their patience, Robert Loveridge (Portsmouth) for Cretaceous (Dalla Vecchia and Ligabue, 1993; this paper) and 10 m photography, Michael Fastnacht, Dino Frey, Andre Veldmeijer, Darren or more in at least two distinct Late Cretaceous species (Lawson, Naish and Mark Witton for discussion of matters pterosaurian. We are } 1975; Witton and Naish, 2008). These patterns of size distribu- grateful to Dino Frey and Attila Osi for helpful comments that much tion do not encourage the view that large size and gigantism in improved the manuscript. D.M. Martill, D.M. Unwin / Cretaceous Research 34 (2012) 1e9 9

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