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Research 32 (2011) 236e243

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Cretaceous Research

journal homepage: www.elsevier.com/locate/CretRes

A new pterodactyloid from the Santana Formation (Cretaceous) of

David M. Martill

School of Earth and Environmental Sciences, University of Portsmouth, Burnaby Road, Portsmouth PO1 3QL, article info abstract

Article history: A partial comprising fused /premaxilla and palate of a ctenochasmatoid pterosaur from the Received 13 July 2009 Santana Formation of the in NE Brazil is named as the new and Unwindia Accepted in revised form 1 December 2010 trigonus gen. et sp. nov. on account of its long slender rostrum, isodonty with raised dental alveoli and Available online 14 December 2010 dentition of seven tooth pairs restricted to the portion of the rostrum anterior to the nasoantorbital fenestra. Unwindia is assigned to the Ctenochasmatoidea, and is probably within the . Keywords: Ó 2010 Elsevier Ltd. All rights reserved. Pterosauria Ctenochasmatoidea Unwindia trigonus gen. et sp. nov. Santana Formation Brazil

1. Introduction world, surpassing the Solnhofen (Wellnhofer, 1970, 1975) and approaching the diversity of the combined Yixian Pterosaur remains are both abundant and diverse in the Santana and Jiufotang formations of the Chinese Jehol Group (Lü et al., Formation Lagerstätte of Brazil with some 12 nominal genera 2006a). A new partial pterosaur skull described here indicates the (, Araripedactylus, Araripesaurus, , Colo- presence of a new genus and species of ctenochasmatoid pterosaur borhynchus, , Criorhynchus, Pricesaurus, Santana- in the Santana Formation assemblage, a group that were wide- dactylus, Tapejara, , ) and an undescribed spread in the Early Cretaceous of . edentulous form similar perhaps to Chaoyangopterus. The genera encompass more than 20 species (Price, 1971; Wellnhofer, 1985, 1987, 1991; Buisonjé de, 1980; Leonardi and Borgomanero, 1985; 2. The new specimen Wellnhofer and Kellner, 1991; Dalla Vecchia, 1993; Kellner and Tomida, 2000; Fastnacht, 2001; Unwin, 2002; Veldmeijer, 2002, The new specimen was obtained from a fossil digger in the small 2003, 2006; Martill and Naish, 2006) although some are of town of Santana do Cariri, southern Ceará, in north east Brazil. This dubious validity (e.g. Araripedactylus dehmi Wellnhofer, 1977; Ara- region is palaeontologically rich and a thriving commercial trade in ripesaurus castilhoi' Price, 1971; Brasileodactylus araripensis Kellner, has developed, especially within the confines of what is now 1984; Pricesaurus megalodon Martins-Neto,1986). Nevertheless, the the Araripe Geopark World Heritage Site (Martill and Heads, 2007). assemblage is diverse and includes Tapejaridae, Thalassodromidae, Although not obtained from in situ, there is little reason to doubt and Ctenochasmatidae (Table 1). The Santana that the specimen comes from the Santana do Cariri region, and Formation occur either as partly crushed, but frequently from the Romualdo Member of the Santana Formation, which articulated specimens in late diagenetic , and as yields abundantly fossiliferous concretions of comparable lithology uncrushed, 3-D specimens, sometimes with soft tissues preserved (Martill, 1993; Fara et al., 2005). in eodiagenetic carbonate concretions (Martill, 1988, 1997; Martill The specimen, housed in the collection of the Museum für and Unwin, 1990). This abundance and diversity make the San- Naturkunde, Karlsruhe, Germany, SMNK PAL 6597 comprises tana Formation one of the richest sources of pterosaur fossils in the a fused left and right premaxilla and maxilla in a carbonate that has been split to reveal the right side of the rostrum (Fig. 1). The counterpart to the concretion is missing. Some bone compacta has fallen away from the specimen revealing an internal Abbreviations: NHMU, Natural History Museum, London, UK; CAMS, Sedgwick mould of crystalline . Four teeth are present in the anterior Museum, Cambridge, UK; SMNK, Staatliches Museum für Naturkunde, Karlsruhe, Germany; YORM, Yorkshire Museum, York, UK. jaws of the right side (Fig. 1b) and mechanical preparation has been E-mail address: [email protected]. employed to reveal the anterior border of the nasoantorbital

0195-6671/$ e see front matter Ó 2010 Elsevier Ltd. All rights reserved. doi:10.1016/j.cretres.2010.12.008 D.M. Martill / Cretaceous Research 32 (2011) 236e243 237

Table 1 three-dimensionality of the right side and palatal surface has Names applied to Santana Formation pterosaurs. survived. Anhanguera santanae (Wellnhofer, 1985) Anhanguera blittersdorffi (Campos and Kellner, 1985) Araripesaurus castilhoi (Price, 1971) 2.1. Systematic palaeontology Araripedactylus dehmi (Wellnhofer, 1977) nomen dubium

Brasileodactylus araripensis (Kellner, 1984) PTEROSAURIA (Kaup, 1834)

Cearadactylus atrox (Leonardi and Borgomanero, 1985) (Plieninger, 1901) Cearadactylus? ligabuei (Dalla Vecchia, 1993) LOPHOCRATIA (Unwin, 2003) CTENOCHASMATOIDEA (Unwin, 1995) robustus (Fastnach, 2001) Coloborhynchus speilbergei (Veldmeijer, 2003) UNWINDIA gen. nov. Coloborhynchus piscator (Kellner and Tomida, 2000)

Pricesaurus megalodon (Martins-Neto, 1986) Diagnosis and etymology: see for type and only species below.

Santanadactylus pricei (Wellnhofer, 1985) Unwindia trigonus gen. et sp. nov. Santanadactylus araripensis (Wellnhofer, 1985) Santanadactylus brasiliensis (Buisonjé de, 1980) Santanadactylus spixi (Wellnhofer, 1985) Derivation of name: Unwindia, after Dr David Unwin; trigonus, referring to the triangular cross-section of the rostrum. mesembrinus (Wellnhofer, 1987) Tropeognathus robustus (Wellnhofer, 1987) (Kellner, 1989) : partial rostrum with teeth, SMNK PAL 6597 Tupuxuara longicristatus (Kellner and Campos, 1988) (Fig. 1aec). Tupuxuara leonardi (Kellner and Campos, 1994) Tupuxuara deliridamus (Witton, 2009) Thalassodromeus sethi (Kellner and Campos, 2002) Type locality: Santana do Cariri region, southern Ceará, NE Brazil.

Type horizon and age: Romualdo Member of the Santana fenestra of the left side (Fig. 1c) and parts of the palate. The region Formation, Araripe Group. Late Early Cretaceous, probably latest of the nasoantorbital fenestra of the right side has suffered from (see Martill, 2007). fracturing and is not clearly discernible (Fig. 1a). Complete prepa- ration of the specimen has not occurred due to the very fragile Diagnosis: U. trigonus can be distinguished from other ptero- of the thin bone compacta. The matrix of the concretion is an saurs by the reduced dentition of the maxilla/premaxilla of only 7 ostracod limestone of light buff shade, one of several typical tooth pairs all of which lie anterior to the nasoantorbital fenestra: lithologies of Santana Formation concretions (Martill, 1997). this is apomorphic for the species. The teeth are of similar size, Crushing of the anterior premaxilla on the left side indicates that distinguishing them from the marked heterodonty of other Santana the concretion is not of the earliest diagenetic variety, but some formation tooth-bearing pterosaurs.

Fig. 1. New ctenochamatoid pterosaur Unwindia trigonus (SMNK PAL 6597) from the Romualdo Member of the Santana Formation, NE Brazil. (a) Rostrum within concretion seen in right lateral view. The line diagram above highlights the bones and teeth; (b) detail of the dentition and anterior alveoli; (c) anterior margin of the nasoantorbital fenestra (nasoantorbital fenestra) of the left side with the premaxilla/maxilla arrowed. Scale bars 10 mm. 238 D.M. Martill / Cretaceous Research 32 (2011) 236e243

Table 2 compaction and it is possible, though unlikely that one more Table of measurements. alveolus could have been accommodated in the jaw tip. Teeth are Length of specimen 221.0 mm present in the posterior most four alveoli of the right side. The teeth Length from anterior border 192.0 mm are simple, slightly laterally compressed cones slightly deflected of Nasoantorbital Fenestra to est. toward the midline of the palate. The largest tooth has a recon- anterior tip of rostrum structed height of 9 mm (Table 2), the smallest, and anterior most Height of rostrum at anterior 36.0 mm border of Nasoantorbital Fenestra tooth is a partially erupted example only 5 mm high. The dental Length of alveolus 1 indet. (3.5 mm) alveoli have a slightly oval outline with the long axis parallel with (number in brackets ¼ distance to next alveolus) the rostral long axis. The anterior two alveoli are slightly more Length of alveolus 2 5.5 mm (4.0 mm) round than those located posteriorly. The alveolar borders are Length of alveolus 3 5.0 mm (3.0 mm) Length of alveolus 4 4.0 mm (4.0 mm) slightly raised. There is a very slight lateral expansion distally of the Length of alveolus 5 5.5 mm (6.5 mm) tooth-bearing part of the rostrum and alveoli 2 and 3 are more Length of alveolus 6 5.5 mm (8.0 mm) laterally located than the posterior alveoli. Length of alveolus 7 5.0 mm The anterior border of the nasoantorbital fenestra is sharply Width of rostrum at 7th alveolus 10.0 mm rounded such that the dorsal and ventral margins must have been nearly parallel along much of its length. Although the suture 2.1.1. Description between the maxilla and premaxilla is distinct, the two bones are The preserved rostrum clearly displays the premaxilla, maxilla fused. It parallels the dorsal border of the rostrum and merges and palatines, but the only distinct boundary is that of the gently with the anterior dorsal border of the nasoantorbital premaxilla and maxilla on the lateral surfaces of the rostrum fenestra. Despite how much of the nasoantorbital fenestra is (Fig. 1). The anterior most part of the rostrum is damaged, and up to missing, the preserved portion suggests a flat oval outline rather 10 mm are estimated to be missing. Seven dental alveoli are than the rounded triangular outline of many tooth-bearing pter- present, but the anterior most alveolus is slightly damaged by odactyloids. The distance from the anterior margin of the

Fig. 2. Portion of anterior rostrum of Lonchodectes compressirostris (Owen, 1851), NHMUK 39410, from the Chalk Formation of Burham, Kent, , with the anterior most margin of the nasoantorbital fenestra in (a), dorsal, (b), right lateral and (c), palatal view. (d) Cross-section viewed from posteriorly. Scale bar ¼ 10 mm. D.M. Martill / Cretaceous Research 32 (2011) 236e243 239

Fig. 3. Anterior rostrum and dentary symphysis of Lonchodectes giganteus (Bowerbank, 1846), NHMUK 39412, from the Chalk Formation, Burham, Kent, England, in right lateral view. Scale bar ¼ 10 mm. nasoantorbital fenestra to the rostrum tip is 192 mm, and the tooth-bearing pterosaurs, especially members of the non-pter- height of the rostrum at the anterior margin of the nasoantorbital anodontid ornithocheiroids, and lonchodectid azhdarchoids. Lü fenestra is 36 mm, giving a rostral index (RI sensu Martill and Naish, et al., (2009) consider Lonchodectidae to be closer to Azhdarchoi- 2006) of 5.3. No teeth are present beneath the nasoantorbital dea than to Ctenochasmatoidea (Unwin, 2009) and fenestra. Projected anteriorly, the maxilla/premaxilla suture ctenochasmatoids. extends toward the tip of the rostrum and no downward deflection There is considerable variation in dental morphology of cte- toward the dental border is detected, suggesting that much of the nochasmatoids with basal forms, such as Cycnorhamphus and tooth-bearing part of the jaw comprises the maxilla. A few fine Pterodactylus, having relatively short conical teeth, which in the (diameter 0.5 mm) nutritive foramina are present on the lateral former, are situated entirely at the jaw tip (Wellnhofer, 1970). More maxilla above the dental battery. derived taxa, such as , and , The palatal surface displays a raised dental border and a prom- have fine, closely spaced dentition extending almost along the inent palatal ridge flanked by deep sulci. It extends anteriorly from entire jaw length (Wellnhofer, 1970; Bonaparte, 1970). A possible near the nasoantorbital fenestra to at least the 3rd tooth pair, but exception to this was the unusual Brazilian supposed cte- compaction prevents its detection more anteriorly than that. In nochasmatoid Cearadactylus atrox, with prominent, enlarged teeth lateral view the palatal ridge and dental borders are the same in the anterior jaw (Dalla Vecchia, 1993; Unwin, 2002). However, height anteriorly, but posteriorly it rises somewhat to become the holotype of this has recently been shown by Vila Nova slightly higher than the dental border just anterior to the nasoan- et al. (2010) to be a poorly repaired specimen of an orni- torbital fenestra. thocheirid close to Anhanguera and the enlarged teeth a fabrication. The cross-sectional shape at the nasoantorbital fenestra is a high Some similarities exist between U. trigonus and some of the triangle, but at alveolus 7 the premaxillae have been compacted Santana Formation ornithocheirids. In particular, ornithocheirids together to give an artificial appearance of a ‘’. The overall lacking or having only a low rostral crest such as Anhanguera pis- appearance is of a very long and slender rostrum with acute cator Kellner and Tomida (2000), have a maxilla/premaxilla suture triangular cross-section and with well-spaced, anteriorly located uniting with approximately the same angle and location on the teeth (Fig. 2). nasoantorbital fenestra as in U. trigonus. However, in A. piscator the tooth row is much more extensive and teeth are present to a loca- 3. Affinities tion halfway along the nasoantorbital fenestra (Kellner and Tomida, 2000, Fig. 4). Also, Anhanguera spp., and Coloborhynchus spielbergi The presence of teeth in Unwindia clearly rule out any affinities have a prominent ‘bull nose’ tip to the rostrum, where the anterior with the Santana Formation azhdarchoid pterosaurs Tapejara, most teeth project forwards from a nearly vertical or steeply Tupuxuara,orThalassodromeus, and the edentulous ornithocheiroid inclined palate, rather than the tapered termination seen in and nyctosaurids. An affinity with Dsungeripteridae can U. trigonus.InOrnithocheirus mesembrinus (Wellnhofer, 1987) the be ruled out on account of the presence of teeth in the anterior jaw, anterior of the rostrum is inflated into a prominent terminal crest whereas in dsungaripterids the anterior rostrum is edentulous that lacks the anteriorly projecting teeth of Anhanguera and Colo- (Unwin, 2003). The long, slender jaw of Unwindia is typical of many boryhnchus. The tooth count is also considerably higher in these 240 D.M. Martill / Cretaceous Research 32 (2011) 236e243

England. In particular, the relatively isodont dentition, slightly raised alveoli and slender build to the rostrum compare well with some species of Lonchodectes reviewed by Unwin (2001). This taxon is very poorly known, with all based on fragmentary jaw tips. Some isolated postcranial material has been assigned to Lonchodectes, but its association with type jaws is suspect. Typically Lonchodectes has a slender rostrum with well-spaced, uniform teeth and a prominent palatal ridge, although these features alone do not diagnose the taxon. Six species of Lonchodectes have been described from the English Cretaceous: Lonchodectes sagittirostris (Owen, 1874) from the Hastings Sands; Lonchodectes compressir- ostris (Owen, 1851); Lonchodectes machaerorhynchus (Seeley, 1870); Lonchodectes microdon (Seeley, 1870) and Lonchodectes platystomas (Seeley, 1870) from the , and ‘Lonchodectes’ giganteus (Bowerbank, 1846) from the Chalk. L. compressirostris also ranges up into the Chalk (Unwin, 2001). Witton et al. (2009) also reported possible lonchodectid material from the English Wealden originally collected by Gideon Mantell, but this lacks cranial material. Lü et al. (2006b) describe a lonchodectid, Yixianopterus, from the Yixiang Formation of that comprises a near complete . Of the six species of Lonchodectes, only L. platystomus, L. microdon, L. compressirostris and ‘L’. giganteus are known from rostral material and can therefore be compared with U. trigonus. Only in L. com- pressirostris and ‘L’ . giganteus is the anterior border of the nasoan- torbital fenestra seen (Figs. 3 and 4). Nevertheless, the other taxa are still informative, despite their highly fragmentary nature. The holo- type of L. microdon (CAMSM B54.486, Unwin, 2001, Fig. 11d) displays prominent raised lateral dental borders and a distinctive palatal ridge flanked by deep sulci as in Unwindia, and a similar condition is seen in examples of L. compressirostris from both the Cambridge Greensand (CAMSM B54.584, Unwin, 2001, Fig. 11b) and the Chalk (Fig. 2). The palatal surface of L. platystomus (YORM 1983/113F, Unwin, 2001, Fig. 12b) shows the median ridge disappearing between tooth pairs two and three, but it is not possible to see this in Unwindia. In specimen NHMU 39410, an example of L. compressirostris from the Chalk Formation the relationship between the dental border and the anterior margin of the nasoantorbital fenestra clearly shows that in this species the maxilla beneath the nasoan- torbital fenestra is tooth bearing, as it is in ‘L’. giganteus (Fig. 3) but in contrast to the condition seen in Unwindia. This specimen also shows the anterior margin of the palatal fenestra present as far anteriorly as the same margin of the nasoantorbital fenestra, although the situation for this character cannot be seen in Unwin- dia. The relationship of the maxillary dentition to the anterior border of the nasoantorbital fenestra is not discernable in other species of Lonchodectes. Unwindia differs from L. platystomus and ‘L’. giganteus in lacking a sagittal crest. Thus the greatest similarity is between the new specimen and L. compressirostris. However, it seems unwise to place Unwindia in Lonchodectidae due to the significant differences exhibited by the highly reduced dentition and lack of teeth beneath the nasoantorbital fenestra. Furthermore, as part of his diagnosis for Lonchodectidae, Unwin (2003) notes that the rostrum is dorsoventrally flattened, and that appears not to be the case for Unwindia, or indeed, ‘L’. giganteus, hence the use of Fig. 4. of ctenochamatoid pterosaurs showing relationship of premax/maxillary teeth to anterior border of nasoantorbital fenestra: (a) Pterodaustro; (b) Ctenochasma, quotation marks here. (c) Gnathosaurus; (d) Pterodactylus; (e) Cycnorhamphus; (f) Cearadactylus atrox; (g) Comparisons made with the tooth-bearing Chinese Early Unwindia trigonus. (a) After Chiappe et al. (2000);(bee) after Wellnhofer (1978); (f) Cretaceous pterosaurs of the , in particular with after Wellnhofer (1987), back of skull restored. Boreopterus, , Haopterus and Yixianopterus reveal some similarities (see Fig. 5). In Boreopterus the teeth are long, extremely ornithocheirids, with 12 pairs in O. mesembrinus (Wellnhofer, 1987, slender and numerous (at least 29 pairs in the premaxilla/maxilla) Fig. 2), around 16 in A. piscator (Kellner and Tomida, 2000, Figs. 4, 5 extending posteriorly to a position well beneath the nasoantorbital and 17) for C. spielbergi Veldmeijer (2003). fenestra and the rostral index is around 6.5 (6.6 in Unwindia) taken U. trigonus considerable similarities to fragmentary jaw- directly from a photograph of the specimen (Lü et al., 2006a, based taxa from the Chalk and Cambridge Greensand formations of Fig. 3.19), but estimated at 5.0 to account for some distortion of the D.M. Martill / Cretaceous Research 32 (2011) 236e243 241

Fig. 5. Non-ornithocheiroid tooth-bearing pterosaurs from the Early Cretaceous Yixian Formation, China. (a) Feilongus youngi (after Wang et al., 2005); (b) Boreopterus cuiea (after Lü and Ji, 2005); (c) smoothed line drawing of Haopterus gracilis compensating for compaction artefacts. Based on d, below; (d) H. gracilis line drawing taken from Fig. 3.21 in Lü et al. (2006a). All scaled such that the distance between the jaw tip and the nasoantorbital fenestra is equal. rostrum due to compaction. Feilongus, also with long, slender teeth, is clear from ontogenetic studies on both pterodactyloid and stem has an elongate rostrum with an R.I. of w8.6, considerably higher group pterosaurs such as spp. that, where large than that of Unwindia but, as in Unwindia, the teeth of Feilongus do data sets are available, tooth count is not necessarily variable not extend as far posteriorly as the nasoantorbital fenestra, but through ontogeny and thus tooth counts might be a reliable diag- there are considerably more (w20 tooth pairs in premaxilla/ nostic character. The of the smallest (and presumed maxilla). Similarities between Unwindia and Haopterus (Wang and youngest) specimens of Rhamphorhynchus from the Jurassic Sol- Lü, 2001) include a rostral index of w6.0 for Haopterus gracilis (6.6 nhofen Limestone of Germany (see Bennett, 1995, 1996) just 7 for Unwindia) and a similar tooth morphology. It is difficult to tooth pairs in the mandible (e.g. Rhamphorhynchus longicaudus). establish the tooth count for Haopterus, due to overlapping bones in Similarly, seven tooth pairs are present in the largest form Rham- the holotype, but at least 10 tooth pairs can be seen, extending phorhynchus longiceps. Consequently, using these and other criteria posteriorly to a location just beneath the anterior most part of the Bennett (1995) united all Rhamphorhynchus species from the Sol- nasoantorbital fenestra. Although Wang and Lü (2001) placed nhofen limestone under a single species. In the genus Pterodactylus, Haopterus in Ornithocheiridae, there is no aspect of the dental Pterodactylus micronyx and P. antiquus have similar tooth counts in morphology to suggest Haopterus is an ornithocheirid sensu Unwin the maxilla/premaxilla and no teeth are present behind the anterior (2003), and it more likely is close to Cycnorhamphus or Pter- border of the nasoantorbital fenestra (as is the case in Unwindia). odactylus (Fig. 5). The same is true for Ctenochasma elegans. In contrast, Pterodactylus The rostral index of 6.6 for U. trigonus sets it apart from most kochi has a distinctive tooth count and the posterior most tooth of other toothed pterosaurs, except Haopterus, and is significantly the maxilla is located beneath the nasoantorbital fenestra. On these higher than values obtained for ornithocheirids, although it is criteria a case can be made for removing P. kochi from Pterodactylus similar to that of Pterodactylus antiquus (RI ¼ 5.4). altogether. The dental count has not been used extensively in pterosaur A taxon that might have some bearing on the relationships of systematics except in general terms at generic and specific level, Unwindia is Yixianopterus jingangshanensis (Lü et al., 2006b) from even though it is remarkably variable over all pterosaurs (from the Yixian Formation. This taxon was tentatively placed in Lon- almost 1000 in Pterodaustro to a few or none in many groups) (but chodectidae on account of dental characters. The teeth are short, see Veldmeijer, 2009 who successfully used tooth count, tooth robust, slightly recurved cones, and are set in alveoli with slightly diameter and alveolar distance to distinguish specimens of raised margins. The tooth count appears to be 6 pairs in the Anhanguera, Brasileodactylus and Coloborhynchus). Nevertheless, it premaxilla/maxilla, all of which are located anterior to the 242 D.M. Martill / Cretaceous Research 32 (2011) 236e243 nasoantorbital fenestra and are of similar size (the first two pairs Fara, E., Saraiva, A.A.F., Campos, D.A., Moreira, J.K.R., Carvalho Siebra, D., appear slightly larger). The rostral index cannot be determined Kellner, A.W.A., 2005. Controlled excavation in the Romualdo Member of the Santana Formation (early Cretaceous, Araripe Basin, northeast Brazil): strati- fi accurately from the gure provided, but is close to 5.5. graphic, palaeoenvironmental and palaeoecological implications. Palae- The highly reduced tooth count of only seven tooth pairs, all ogeography, Palaeoclimatology, Palaeoecology 218, 145e160. anterior of the nasoantorbital fenestra in Unwindia, is a combina- Fastnacht, M., 2001. First record of Coloborhynchus (Pterosauria) from the Santana Formation (Lower Cretaceous) of the Chapada do Araripe, Brazil. Paläontolo- tion of characters not seen in any other pterosaur. Only in some gische Zeitschrift 75, 23e36. members of the Ctenochasmatoidea are the teeth of the maxilla all Kaup, J.J., 1834. Versuch einer Eintheilung der Saugethiere in 6 Stämme und der located anterior of the nasoantorbital fenestra. In the Cte- Amphibien in 6 Ordnungen. Isis 3, 311e315. Leonardi, G., Borgomanero, G., 1985. Cearadactylus atrox nov. gen., nov. sp.: novo nochasmatidae, Gnathosaurus, Ctenochasma, Pterodaustro and Pterosauria (Pterodactyloidia) da Chapada do Araripe, Ceará, Brasil. DNPM, possibly the teeth are all located anterior of the Coletânea de Trabalhos Paleontológicos, Série Geologia 27, 75e80. nasoantorbital fenestra. However, in ctenochasmatid cte- Lü, J.C., Ji, Q., 2005. A new ornithocheirid from the Early Cretaceous of Province, China. Acta Geologica Sinica 79, 157e163. nochasmatoids the teeth are numerous, long and slender and quite Lü, J.C., Ji, S., Yuan, C., Ji, Q., 2006a. Pterosaurs from China. Geological Publishing unlike those of Unwindia.InY. jingangshanensis Lü et al. (2006a,b), House, Beijing, 147 pp. [in Chinese, with English Figure Captions]. tentatively placed in Lonchodectidae by Unwin et al. (2000), there Lü, J.C., Ji, S., Yuan, C., Gao, Y.B., Sun, Z.Y., Ji, Q., 2006b. New pterodactyloid ptero- appears to be a reduced dentition of only 6, relatively uniform, saurs from the Lower Cretaceous Yixian Formation of western Liaoning. In: Lü, J.C., Kobayashi, Y., Huang, D., Lee, Y.-N. (Eds.), Papers from the 2005 Heyuan widely spaced anteriorly located teeth in the rostrum similar to Symposium. Geological Publishing House, Beijing, pp. 195e203. those of Unwindia, and a similar situation is found in Cycno- Kellner, A.W.A., 1984. Occurrência de uma mandíbula de Pterosauria (Brasileo- rhamphus suevicus (8 pairs of rostral teeth) and Cycnorhamphus dactylus araripensis, nov. gen, nov. sp.) na Formação Santana, Cretáceo da Chapada do Araripe, Ceará, Brasil. In: 33 Anais Congresso Brasileiro de Geologia, canjurensis (tooth count not known but no teeth present between vol. 2 578e590. the broken rostral tip and the nasoantorbital fenestra (see Fabre, Kellner, A.W.A., Campos, D.A., 1988. Sobre um nova pterossauro com crista sagital da 1974), both basal ctenochasmatoids according to Unwin, 2003). Bacia do Araripe, Cretáceo Inferior do Nordeste do Brasil. Annais da Academia Brasileira, Ciências 60, 459e469. Thus, U. trigonus appears to be close to Yixianopterus, and perhaps Kellner, A.W.A., 1989. A new edentate pterosaur of the Lower Cretaceous from the Cycnorhamphus on the basis of its dentition. The slender rostrum of Araripe Basin, northeast Brazil. Anais da Academia Brasiliera, Ciencias 61, Unwindia is somewhat similar to that of Lonchodectes than Cycno- 439e446. Kellner, A.W.A., Campos, D.A., 1994. A new species of Tupuxuara (Pterosauria, rhamphus which has a slightly blunt termination to the rostrum Tapejaridae from the Early Cretaceous of Brazil. Annais da Academia Brasileira, with its dorsal border subparallel to the dental border and a more Ciências 66, 467e473. triangular nasoantorbital fenestra. Accordingly, Unwindia is placed Kellner, A.W.A., Tomida, Y., 2000. Description of a new species of (Pterodactyloidea) with comments on the pterosaur fauna from the Santana within Ctenochasmatoidea, but its precise position within the Formation (-Albian), northeastern Brazil. National Science Museum group can only be resolved with the discovery of more complete Monographs, Tokyo 17, 135 pp. material. Kellner, A.W.A., Campos, D.A., 2002. The function of the cranial crest and jaws of a unique pterosaur from the Early Cretaceous of Brazil. Science 297, 389e392. Martill, D.M., 1988. The preservation of fossil fishes in concretions from the Cretaceous of Brazil. Palaeontology 31, 1e18. Acknowledgments Martill, D.M., 1993. Fossils of the Santana and Crato formations, Brazil. In: Palae- ontological Association Field Guides to Fossils, vol. 5, 159 pp. I am especially grateful to Sandra Chapman and Lorna Steel, Martill, D.M., 1997. oblique to bedding in early diagenetic concretions from the Cretaceous Santana Formation of Brazil e implications for substrate consis- NHM and Dino Frey, SMNK for granting access to specimens in their tency. Palaeontology 40, 1011e1026. care and to Bob Loveridge who very kindly took the photographs. Martill, D.M., 2007. The age of the Cretaceous Santana Formation fossil Konservat Thanks to Chris Bennett for helpful referee comments. Thanks also Lagerstätte of north-east Brazil: a historical review and an appraisal of the bio- chronostratigraphic utility of its palaeobiota. Cretaceous Research 28, 895e920. to Mark Witton, , Dave Unwin, André Veldmeijer, and Martill, D.M., Heads, S.W., 2007. 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