André J. Veldmeijer 1, Marco Signore 2 & Hanneke J.M. Meijer 3 1 Natuurhistorisch Museum Rotterdam & PalArch Foundation 2 Dipartimento di Scienze della Terra, Università degli Studi di Napoli “Federico II” 3 Faculty of Movement Sciences, Free University of Amsterdam

Description of two () from the Lower Santana Formation,

Veldmeijer, A.J., Signore, M. & Meijer, H.J.M., 2005 - Description of two pterosaur (Pterodactyloidea) mandibles from the lower Cretaceous Santana Formation, Brazil - DEINSEA 11: 67-86 [ISSN 0923- 9308]. Published 29 December 2005

Two mandibles of pterodactyloid from the Lower Cretaceous Santana Formation (Romualdo, ) of Brazil are described and classified. The edentulous jaw is assigned to . This anterior part completes the reconstruction of the lower jaw, hitherto based on the lacking this part. The toothed is assigned to and is the first complete mandible of this .

Keywords: Anhanguera, Thalassodromeus, , Lower Cretaceous, Santana Formation, Romualdo.

Correspondence: A.J. Veldmeijer, Natuurhistorisch Museum Rotterdam, P.O. Box 23452, NL-3001 KL Rotterdam, the Netherlands, e-mail [email protected] M. Signore, Dipartimento di Scienze della Terra, Università degli Studi di Napoli “Federico II”, L.go S.Marcellino 10, 80132 Napoli, Italy. H.J.M. Meijer, Faculteit Bewegingswetenschappen,Vrije Universiteit Amsterdam, Van der Boechorststraat 9, 1081 BT Amsterdam, e-mail [email protected]

INTRODUCTION cussed only briefly, because a broader study The described specimens both originate is in progress (Veldmeijer, in press). from the Romualdo Member (Albian) of the The material is housed in the collection Santana Formation (-Albian) (Martill Oberli; casts of SAO 251093 are housed in the et al. 1993; see also Beurlen 1971; Kellner Bayerische Staatssammlung für Paläontologie & Tomida 2000; Maisey 1991; Pons et al. und historische Geologie, Munich, Germany, 1990), in Chapada do Araripe in the prov- the Staatliches Museum für Naturkunde, ince of Ceará, Brazil. Here new material of Stuttgart, Germany and Museu Nacional, Rio Thalassodromeus is described and compared de Janeiro, Brazil. with other edentulous taxa. The toothed jaw is Although it is not common to publish mate- briefly described; the focus is on the discus- rial from private collections, the decision to sion of morphological differences relative to do so is based on the fact that the collection other pterosaurs from Brazil. For a broader is unconditionally accessible. Furthermore, description of the general layout of these pter- other scientists (i.e. Wellnhofer & Kellner osaurs, see Kellner & Tomida (2000), Kellner 1991) have used the collection in their studies, & Campos (2002),Veldmeijer (2003a), and which underlines the quality of the collec- Wellnhofer (1985, 1991). Dentition patterns tion and its professional management. Finally, (i.e. the graph that visualises the size and none of the are . Access position of the teeth) are plotted and dis- to the original specimens can be arranged

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through the owner of the collection (St. Gallen, Diagnosis Thalassodromeus according to Switzerland, [email protected]) or the Natural Kellner & Campos (2002: 389) the same as for History Museum at St. Gallen (T. Bürgin, the T. sethi. “[…] anterior portion of Naturmuseum Sankt Gallen, Museumstrasse the premaxillae and dentary with sharp dorsal 32, [email protected]). and ventral edges; […]” Between […] infor- mation on other parts than the mandible. ABBREVIATIONS ad.fos. adductor fossa Species Thalassodromeus sethi ang. angular ch.t. foramen chorda tympani Holotype As for the specimen. d. dentary d.e. dorsal cutting edge Etymology note One of the authors (Kellner, d.sag.cr. dentary sagittal crest pers.com. 2002) explained that the shape of lat.cot. lateral cotyle the crest of the pterosaur reminded them of the man.sul. mandibular sulcus crown of the god Seth. It is however notewor- m.fos. Meckelian fossa thy that the crown, representing the solar disk med.cot. medial cotyle with two tall plumes, is typically worn by the pat. pathology god Amon (later Amon-Ra), or manifestations pn.for. pneumatic foramen of him, and not by Seth (for instance Baines & pre.art. prearticular Málek 1981). ret.pr. retroarticular process r. ramus Amended diagnosis Anterior part, starting ri. ridge from the anterior border of the mandibular sul- s.sh. symphyseal shelf cus, bent in dorsal direction. spl. splenial sur. surangular DESCRIPTION sym.cav. symphyseal cavity The specimen represents the symphyseal por- t.b.r. elevated tooth-bearing rim tion of the anterior mandible, but lacks the tip (approximately 10 mm). Rami are missing; the SYSTEMATIC PALAEONTOLOGY remaining part of the right one is slightly long- er than the left one. The specimen is broken Pterosauria Kaup, 1834 in two parts. The has been completely Suborder Pterodactyloidea Plieninger, 1901 prepared mechanically except at the tip where some matrix remains to protect the fragile Mandible SAO 251093 point from breaking. No sutures are discern- Figures 1, 2; Table 1 able. The symphysis of the edentulous mandible Tapejaridae Kellner, 1989 is curved dorsally, starting from the anterior border of the symphyseal shelf (s.sh.). The Thalassodromeus Kellner & Campos, dorsal aspect of the slender terminal point has 2002 a sharp cutting edge (d.e.) whereas the ventral aspect is less sharp, resulting in a tear-drop Type species and specimen Thalassodromeus cross-section. Posteriorly, the dorsal cutting sethi Kellner & Campos, 2002 - large part of edge terminates at the shelf. The shelf rapidly and mandible, DGM 1476-M, Museu de increases in width posteriorly, diverging with Ciências da Terra/Departamento Nacional de the rami. In dorsal view, two low ridges (ri.) Produção Mineral, Rio de Janeiro, Brazil. occur left and right of the middle of the shelf, 10 mm ventral to the dorsal edge, with a length

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Figure 1 Thalassodromeus sethi, mandible SAO 251093 in various aspects. Right lateral (A), ventral (B), left lateral (C) and dorsal (D) view. Cross-sections at various points (E). Scale bar = 50 mm. [illustration: A.J. Veldmeijer]

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Figure 2 Thalassodromeus sethi, mandible SAO 251093 in various aspects. Right lateral (A), ventral (B), left lateral (C) and dorsal (D) view. Scale bar = 50 mm. [photographs: E. Endenburg, reworked by J.H. van Leeuwen, courtesy Oberli]

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of approximately 30 mm. Both are orientated to the edentulous pterosaurs, some of which slightly posterodorsally. are yet unknown from Brazil. One toothed The posterior symphysis exposes an anterior exception, , is considered here cavity (sym.cav.). It appears to extend at least because the anterior mandible is edentulous as far as the anterior half of the symphyseal (Young 1964; Martil et al. 2000), curved shelf. The mandibular shelf has an oval cross- dorsally and it is compressed laterally. The section with an indented rim. The little part of symphysis in Dsungaripterus is marked by a the rami that is preserved has a slender dorsal weak ridge, that is lacking in SAO 251093. edge, which is slightly curved medially. The The medial posterior mandibular shelf width of the rami increases posteriorly and of Dsungaripterus is not present in SAO ventrally and produces a lingual concavity. In 251093. Furthermore, anterior teeth erupt in lateral view the rami are convex. Dsungaripterus where no alveoli are present As is common in pterosaurs, the mandible is in SAO 251093. Due to the small dimensions lightly built with a bone thickness often less of it in SAO 251093, it can not be determined than 1.0 mm. A posterior view clearly shows whether this jaw is part of an endentulous, the cross-section of the posterior mandible, more advanced Dsungaripterus and classifica- including the reinforcement of the hollow bone tion as such is therefore ruled out for the time by means of thin reticulate plates, forming being. large square-like pneumatized ‘cells’, which has a mandibular cavity with gives the bone optimal strength in combina- a triangular section and thus is not consist- tion with extreme weight reduction. The dorsal ent with the flattened circular cross-section edges of the rami and the symphyseal shelf are in SAO 251093. Furthermore, the symphysis more robust. Transverse bony plates here are is proportionally larger in Pteranodon. The more compact, resulting in smaller ‘cells’. A dorsal aspect of the mandible can be straight clear lateroventral border separates this area (P. longiceps) or dorsally bent (P. stern- from the more lightly constructed central and bergi) (Eaton 1910; Bennett 1994, 2001). In ventral areas. no dorsal curvature can be seen Two irregularities are visible at both lateral (Bennett 2003b; Williston 1902a,b, 1903). surfaces. On the left is a small uneven area Furthermore, all known nyctosaurs are sub- (pat.), interpreted as pathology. On the right is stantially smaller with a skull length of about a comparable irregularity, but longer antero- 30 cm. posteriorly and smaller dorsoventrally. Both Also the non-Brazilian is protrude slightly; the right one a bit more an edentulous pterosaur. These azdarchid than the left one. The surrounding surfaces pterosaurs however are very poorly repre- are slightly dented. It is tentatively suggested sented by mandibular remnants. Few mandi- that these areas may be pathological in origin. bles, although not well-preserved, serve for Other pathologies are described by Bennett comparison (Kellner & Langston 1996). The (1989, 2003a), Kellner & Tomida (2000), elongated symphysis is approximately 60 % Mader & Kellner (1999). Reports by Kellner & of the total length. The cross-section of the Tomida (2000) on the left jugal and quadrato- dentary is triangular which is not consistent jugal pathologies in Coloborhynchus piscator with the cross-section in SAO 251093. The resemble the ones observed here. dorsal rim of the anterior part of the symphysis in Quetzalcoatlus is almost flat. Ventrally, the COMPARISON AND DISCUSSION joined dentaries form a sharp that declines A detailed comparison of SAO 251093 with posteriorly. A mandibular cavity is present, but other taxa is difficult because only the sym- its extent is unknown. No statement regard- physis is known. Since SAO 251093 lacks ing the cross-section was made by Kellner & teeth and alveoli, comparisons are limited Langston (1996: 230), but they suggest a fur-

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ther investigation into the hypothesis that the Campos 1990). The type specimen includes cavity “[...] might be linked with the presence pieces of the mandible. According to the of a gular sac […]”. The transverse section authors (Kellner & Campos 2002: 391): “A of the rami are laterally convex and medially strong concavity formed by the palatine and concave, so no medial decrease in width is bordered laterally by the maxillae is present observed, as in SAO 251093. under the anterior half of the nasoantorbital The family Tapejaridae consists of the genera fenestrae. Anterior to this concavity, the pal- Tapejara, and Thalassodromeus ate is convex, forming a short ventral keel that from Brazil and from . turns into a sharp blade anteriorly. The fused Tapejara is an edentulous pterosaur genus, dentaries form a perfect counterpart to the consisting of two species from Brazil, Tapejara palate, with a developed concavity, followed wellnhoferi Kellner, 1989 and Tapejara by a short, deep sulcus (that during occlu- imperator Campos & Kellner, 1997. The sion encases the palatal keel, forming a strong mandible in Ta. wellnhoferi has a depressed interlocking mechanism) and an anterior sharp tip, a ventral keel, and a depressed shelf bor- bony blade. Between both blades there is a dered by the raised lateral rims of the dorsal gap.” Following this preliminary analysis, a margin. Thus it differs from SAO 251093 complete description is forthcoming in the near which lacks a crest, has a dorsal curvature and future (Kellner, pers.comm. 2002). lacks the depressed but has a distinct sulcus. It is clear that the mandible SAO 251093 No mandible is known of Ta. imperator, but can be regarded as Thalassodromeus, having a the depressed rostral tip suggests a dorsal con- short and deep shelf posterior to the concave, tour of the mandible similar to that of Ta. well- sharp edged blade. However, the rami in Th. nhoferi. Sinopterus dongi Wang & Zhou, 2000 sethi are straight and diverge slightly postero- is similar in gross morphology to Tapejara and laterally. This contrasts with SAO 251093 in is, therefore dissimilar to SAO 251093. which rami diverge stronger posterolaterally Tupuxuara is also an edentulous taxon from and exhibit a slight but distinct bending in Brazil but descriptions of complete mandi- lateral direction. The mediodorsal parts of the bles have not been published despite the fact rami in Th. sethi are more concave relative to that mandibles are known. However, a nearly the rami in the St. Galler mandible. Here the complete specimen of Tu. leonardii Kellner mediodorsal side of the rami is only slightly & Campos, 1994 in the Iwaki Museum of concave and slightly extends to the bottom of Coal Mining & Fossils, Japan, and the partial the shelf. In lateral view the mandible of Th. in the Prefectural Museum of Natural sethi shows a distinct rise, whereas in SAO History, Kanagawa, Japan, include mandibles. 251093 its only slightly convex. In ventral In general, the mandible is longer and smaller view Thalassodromeus anteriorly displays a than in Tapejara. In dorsal view the anterior distinct ventrally bulging area. In similar fash- third of the symphyseal shelf is flat, with ion, the anterior tip in SAO 251093 is bent in slightly raised rims, contrasting the sharp dor- dorsal direction, starting at the anterior border sal cutting edge of SAO 251093. Furthermore, of the symphyseal shelf. Although the anterior the mandibles in Tupuxuara shows no dorsal tip in Th. sethi is missing, there is no indica- curvature. The mandible in Tu. leonardii has a tion of bending at the anterior border of the comparable symphyseal shelf but anteriorly the preserved mandible, contra the reconstruction shelf fades into a flat surface. By comparison, shown by Kellner & Campos (2002). the mandibular shelf in SAO 251093 does not So far, only bones were found which show flatten into the rim. the familiar trabecular system (for instance The edentulous pterosaur, recently described in anhanguerids) or a plate-like construction as Thalassodromeus sethi Kellner & Campos, as encountered in Tu. leonardii. But the bone 2002, was first published in 1990 (Kellner & structure described here differs from these;

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Table 1 Measurements of mandible Thalassodromeus sethi (SAO 251093) in mm.

Anterior part Length 153* Height anterior part, anteriormost tip 19.4 Height anterior part, halfway length 46.8 Height anterior part, posteriormost end 51.3 Width anterior part, anteriormost tip 7.2 Width anterior part, posteriormost end 31.2 Width mandible at anteriormost end symphyseal shelf 25.1 Width symphyseal shelf at posteriormost end 14.2

Posterior part Length posterior part, right side 120* Length posterior part, left side 107* Height posterior part, anteriormost end 52.4 Height posterior part, posteriormost end, right side 45.5 Largest width posterior part, anteriormost end 32.7 Width posterior part, anteriormost end, halfway dorsal-ventral 27.6 Smallest width posterior part, anteriormost end 9.3 Width symphyseal shelf at anteriormost end 14.6 Depth symphyseal shelf at posteriormost end 10.5 Width mandibular cavity 29.4 Height mandibular cavity 16.3 Width right ramus, measured at one third height (from dorsal) 1.1** Width right ramus, measured at floor shelf 16.4**

Total Length, as preserved, right side 273* Length, as preserved, left side 260*

* Including the bending ** Measured medial-lateral a slightly comparable inner bone structure is remnant of a primitively dented . mentioned for a ? premaxilla In conclusion SAO 251093 is a small (Wellnhofer & Buffetaut 1999). Possibly, the fragment relative to the type specimen of internal structure of the lower part of the man- Thalassodromeus, but the general morphologi- dible (i.e. the ‘cells’), is comparable to the sit- cal features compare well with this taxon. The uation reported for a Romanian giant pterosau- two differ in various small details, which in rian skull (Buffetaut et al. 2001). The strength- summary are not significant enough to justify ening of the dorsal rims and, to a lesser extent the erection of a new species designation for of the sulcus, compares with the strengthening SAO 251093. The importance of the Swiss of the tooth-bearing parts in toothed pterosaurs specimen lies in the fact that it fills in the grey as reported in Coloborhynchus (Veldmeijer areas imagined by Kellner & Campos (2002) 2003a). This kind of structure might have served and thus makes the complete reconstruction to better withstand the forces exerted on these of the skull of Thalassodromeus possible and parts of the mandible during feeding. The ques- expands the diagnosis of the species. tion rises whether this might be an evolutionary

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SYSTEMATIC PALAEONTOLOGY Dentary The dorsal margin of the mandible is largely Order Pterosauria Kaup, 1834 obscured by matrix, but it can be observed that Suborder Pterodactyloidea Plieninger, 1901 the anterior part of the mandible is slightly expanded. The tooth-bearing rims are raised. Mandible SAO 200602 The depressed area between the rims has a Figures 3-5; Table 2 sulcus (man.sul.), which is flanked by tiny raised ridges. It is uncertain how far the sulcus Family Campos & Kellner, extends anteriorly and posteriorly. The dorsal 1985 margins of the rami are dentaries (d.) that form the dorsal limit of the adductor fossa (ad.fos.). Genus Anhanguera Campos & Kellner, The left and right dentaries join in a symphy- 1985 sis anterior to the Meckelian fossa (m.fos.). A deep dentary sagittal crest (d.sag.cr.) is present, Type species and specimen Anhanguera blit- starting 45 mm anterior to the dentary joint. tersdorffi, skull, MN 4805-V, Museu Nacional, This crest continuously decreases in width Rio de Janeiro, Brazil. ventrally. Fine features include small (< 1 mm) grooves and a few small (< 1 mm) perforations Remark to diagnosis No diagnostic descrip- that insert obliquely, with the grooves leading tion was provided for the anterior part of the towards them. The ventral tip of the crest is mandible in the original description of the slightly abraded. holotype, due to its absence. Teeth Amended diagnosis Mandible with smooth All anterior teeth are complete except for spoon-shaped expanded anterior part, contain- two on the right (numbers 12 and 13), which ing the largest teeth. Presence of mandibu- are broken at the alveoli. The left side has 14 lar sulcus, which is flanked by raised rims, intact teeth. The right has 15. No alveoli could extending until the anterior expansion. be distinguished at the left ramus. Three teeth of the right ramus are partially embedded and Anhanguera sp. indet. separated from the rest of the ramus (they are not illustrated). In anterior view the first alveo- DESCRIPTION lus is oriented anterodorsally. The alveoli two The mandible is complete, but broken into and three are laterodorsally and anteriorly posi- three pieces, the two rami and the anterior por- tioned. Alveolus four is placed laterodorsally tion. Remaining matrix protects the teeth from and slightly anteriorly. The subsequent alveoli, damage but a small area close to the start of numbers six to ten, are increasingly laterally the symphysis is prepared. The toothed mandi- oriented but less posteriorly. ble is long and slender with a ventrally project- The largest alveolar cross-section is the third ing dentary sagittal crest and straight diverging one. The second is only slightly smaller. In rami. The general anatomical features agree general, the teeth are curved in a medial and with previous descriptions of toothed ptero- posterior direction, although the curvature saurs (Kellner & Tomida 2000; Veldmeijer decreases posteriorly. Teeth one to four are 2002, 2003a; Wellnhofer 1985, 1987). Here more strongly curved. The teeth cross-section emphasis is placed on new features. A discus- is elliptical with the long axis oriented antero- sion on the validity of certain characters in medially and at right angles to the curvature other specimens follows. of the teeth. Right tooth nine has an anterola- teral tip facet extending laterally. The polished edges of the facet suggest that the damage is

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Figure 3 Anhanguera sp. indet., mandible SAO 200602 in various aspects. Symphysis in right lateral (A), anterior (B), left lateral (C) and ventral (D) view. Scale bar = 50 mm. [illustration: A.J. Veldmeijer] not post-mortem. Right tooth 10 has a similar portion (lat.cot.) by a diagonal ridge (ri.) that but much smaller facet. is orientated anteromedially and dorsally. This ridge is more prominent relative to another Retroarticular process ridge described for the articular (see below). The posterior quarter and retroarticular proc- The articular has a small (50 mm) pneumatic ess (ret.pr.) of the diverging rami are twisted foramen at its anteroventral corner, which is medially relative to the anterior portions. The separated from the posterior part by a shallow articular (art.) however, is orientated slightly diagonal ridge that extends anterolaterally. The more laterally relative to the surangular. The posterior part is twisted more mediodorsally inner portion of the articulation area (med. relative to the medial and lateral cotyles. The cot.), anterior and dorsal to the pneumatic posteriormost articular is slightly convex. The foramen (pn.for.), is separated from the outer ventral aspect of the retroarticular process con-

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Figure 3 (continued) Anhanguera sp. indet., mandible SAO 200602 in various aspects. Right ramus in lateral (E), ventral (F), medial (G) and dorsal (H) views; right retroarticular process in posterior view (I) and cross section (J)f o the symphysis. Scale bar = 50 mm. [illustration: A.J. Veldmeijer]

sists only of the articular. Against the lateral the articulation area dorsally, medially and lat- expansion of the surangular, the adductor fossa erally. The edges of the cotyles (cot.) expand is provided with a foramen for the transmission dorsally, laterally and medially. In medial view of the chorda tympani (ch.t.). the surangular forms the dorsal limit of the posterior mandibular opening. The anterior Surangular border is not differentiated. Ventrally, the bone In dorsal view, the surangular (sur.) overhangs does not extend more than a few millimeters.

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Figure 3 (continued) Anhanguera sp. indet., mandible SAO 200602 in various aspects. Left ramus in lateral (K), ventral (L), medial (M) and dorsal (N) views; ceratobranchials (O). Scale bar = 50 mm. [illustration: A.J. Veldmeijer]

Angular Prearticular Ventrally the angular extends posteriorly to the The prearticular (pre.art.) forms the ventral articular. In medial view the bone is slender border of the adductor fossa. It is bordered and extends far anteriorly, forming the ventral ventrally by the angular (ang.), anterodorsally border of the mandibular ramus. Possibly, the by the splenial (spl.) and posteriorly by the angular extends even further, but the exact articular. Consequently, it closes the medial course cannot be determined. In lateral view, aspect of the retroarticular process. the angular can be traced to the posterior part of the rami, displaying a small strip of bone of Splenial few mm in height. Though not traceable along In medial view the splenial covers the majority the entire length of the ramus, the anterior part of each ramus. It forms the posterior border of reveals a small strip of the angular immediately the Meckelian fossa and extends towards the posterior to the symphysis. adductor fossa, forming its anteroventral bor-

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Figure 4 Anhanguera santanae, holotype BSP 1982 I 90 in dorsal view. Scale bar = 50 mm. [illustration: E. Endenburg / A.J. Veldmeijer]

der. Ventrally, the splenial is bordered by the The mandible of arari- angular and posteroventrally by the prearticu- pensis Kellner, 1984 has no sagittal crest lar. The dorsal border is limited by the dentary. (see also Sayão & Kellner 2000, Veldmeijer 2003b, Veldmeijer et al. 2005) and is pro- Hyoid apparatus vided with a sagittal sulcus, starting at the Small rod-like bones, separated from the man- anteriormost tip, which has anteriolaterally dible in situ, are identified as ceratobranchial I side-sulcii. atrox Leonardi of the hyoid apparatus. The anteriormost ends & Borgomanero, 1985 lacks a sagittal crest of the long and slender bones diverge slightly (however, the presence cannot be ruled out laterally. Consequently, the anterior aspects are entirely as explained by Kellner & Tomida not in contact with each other. Anteriorly the 2000), on the condition that the classification rods are blunt and slightly bulbous. of Cearadactylus? ligabuei Dalla Vecchia, 1993 is accepted and following Dalla Vecchia’s COMPARISON AND DISCUSSION interpretation that the different parts belong A comparison of SAO 200602 is limited to to the same individual (but see Kellner & the toothed taxa from Brazil. Comparison Tomida 2000). Furthermore, the dentary is follows publication date. The validity of the sloping, containing the largest teeth. Recently, Anhangueridae is controversial (Unwin 2001, Unwin (2002) concluded in his review of Veldmeijer, in press) and beyond the scope of Cearadactylus that Ce. atrox is a valid spe- this work. The Cambridge Greensands mate- cies and assigned Ce.? ligabuei tentatively to rial is, in general, severely fragmented and Anhanguera. often diagnosed on ambiguous characters, so The main problem in comparing the present comparative studies would be extremely lim- mandible with Anhanguera blittersdorffi ited and, in most cases, not worthwhile. The Campos & Kellner, 1985 is the fact that validity of Criorhynchus is controversial as the holotype lacks a mandible. The mandible well (Fastnacht 2001; Kellner & Tomida 2000; in the referred specimen (Kellner & Tomida Unwin 2001), nevertheless the most recent 2000), is substantially smaller in all respects diagnosis of Criorhynchus (Fastnacht 2001), and lacks the retroarticular process. The dorsal is used here. At present Coloborhynchus, aspect however is comparable. The mandible with Co. clavirostis Owen, 1874 from the includes a posteriorly recessed and anteriorly Cambridge Greensands as type specimen, is narrow medial portion with raised tooth-- generally accepted. ing rims posterior to the spoonshaped anterior.

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Figure 5 Anhanguera sp. indet., mandible SAO 200602 in various aspects. Symphysis in right lateral (A), left lateral (B) and ventral (C) view. Scale bar = 50 mm. [photographs: E. Endenburg, reworked by J.H. van Leeuwen, courtesy Oberli]

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Figure 5 (continued) Anhanguera sp. indet., mandible SAO 200602 in various aspects. Right ramus in lateral (D), ventral (E), medial (F) and dorsal (G) views; left ramus in lateral (H) view. [photographs: E. Endenburg, reworked by J.H. van Leeuwen, courtesy Oberli]

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Figure 5 (continued) Anhanguera sp. indet., mandible SAO 200602 in various aspects. Left ramus in ventral (I), medial (J) and dorsal (K) views; ceratobranchials (L). Scale bar = 50 mm. [photographs: E. Endenburg, reworked by J.H. van Leeuwen, courtesy Oberli]

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Table 2 Measurements of mandible Anhanguera sp. (SAO 200602) in mm.

Length 403 Length retroarticular process-symphysis 238 Thickness ramus (possible) last alveolus 7.2 Width expanded anterior part between alveolus 2-3 (ventrally) 20.0 Width expanded anterior part between alveolus 3-4 (ventrally) 8.5 Width expanded anterior part between alveolus 4-5 (ventrally) 15.0 Width at symphysis 37* Width over surangular 12** Length dentary sagittal crest 115 Maximal height dentary sagittal crest 39.7 Length ceratobranchials, including the bending 144 Width, posterior aspect (one bone) 4.0 Width, halfway (both bones) 4.0 Width, anterior aspect (both bones) 7.0

* Approximate, because the right ramus is not preserved at this point. ** Approximate, because the rami are separated from each other and the symphysis.

Furthermore, a medial sulcus is also found in pair of alveoli relative to the others in another A. blittersdorffi in the depressed area. specimen of Co. robustus (SMNK 2302 PAL), Comparison with Coloborhynchus araripen- matching SAO 200602 (see below). The medi- sis (Wellnhofer, 1985) is limited to the rami, al sulcus originates between the ninth and tenth which are highly comparable. The foramina of alveoli, despite the illustration by Fastnacht that splenial are not observed in SAO 200602. (2001) which does not show this. A similar Furthermore, there is no indication of a man- situation is also seen in the referred specimen dibular sulcus but this might be due to the of Anhanguera blittersdorffi. The sulcus in the incompleteness of this specimen. referred specimen of A. bittersdorffi originates The holotype of Anhanguera santanae at the eighth alveolus, thus not as far anteriorly, (Wellnhofer, 1985) lacks a complete retroar- and continues posteriorly in ever increasing ticular process (Fig. 4). The anterior portion of distinctness and width. The sulcus originates the holotype is not preserved, thus the presence against the symphysis, and becomes clearly of a crest is uncertain, but Wellnhofer (1991) visible at the twelfth alveolus in the holotype and Kellner & Tomida (2000) assume its pres- of Co. robustus; the posterior extent of the ence. The lack of the anterior mandible in the sulcus cannot be observed in SMNK 2302 PAL holotype and in AMNH 22555 (Wellnhofer because it is not preserved. 1991) makes it impossible to determine the The mandible in Criorhynchus mesembrinus dentition pattern in these specimens. The incli- (Wellnhofer, 1987) has no anterior expan- nation of the retroarticular process and the con- sion (see also Veldmeijer 2002). The teeth figuration of the posterior part of the symphy- show no distinct variation in size, as seen in sis in SAO 200602 is similar to A. santanae. Coloborhynchus or Anhanguera, and the dorsal The present mandible differs from the holo- aspect lacks a simple sulcus as described in type of Coloborhynchus robustus (Wellnhofer the present work. Instead, Cr. mesembrinus has 1987) in that the anterior expansion in a deep and posteriorly widening sulcus. The Coloborhynchus is distinctly more robust and retroarticular process is more sharply inclined the symphysis is noticeably longer. Fastnacht medially relative to SAO 200602. Kellner (2001) notes the enlarged second and third (1996) notes that in anhanguerids the fifth and

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sixth pair of alveoli are smaller relative to the than in SAO 200602. The anterior extremity of third and seventh pair, as in SAO 200602 and the ceratobranchials is similarly expanded and Co. robustus (BSP 1987 I 47). Fastnacht (2001) the measurements are similar. notes that in Coloborhynchus the second and In summary, the configuration of SAO third pairs of alveoli are the largest, but in 200602 in dorsal view (medial portion the referred specimen of Coloborhynchus recessed relative to the tooth-bearing araripensis the third and fourth alveoli are rims) is also observed in Brasileodactylus, the biggest. In Coloborhynchus spielbergi the Coloborhynchus, Criorhynchus and ninth alveolus is larger than the second one. Anhanguera. The sulcus, however, only occurs In Coloborhynchus piscator the second and in A. santanae and A. blittersdorffi and thus third alveoli are the biggest (Veldmeijer, et al, is an important character separating these in press.) and again the ninth is larger than the otherwise similar taxa. The well-preserved second. Consequently, separating the dental Coloborhynchus specimens have either a pattern as done by Fastnacht (2001) and Unwin sulcus extending towards the anterior aspect (2001) from the rest may be premature. At the (i.e. Co. robustus) or else a weakly developed moment too little is known of the diagnostic sulcus (i.e. Co. spielbergi). The sulcus in status of dentition and this character should be Criorhynchus shows a different morphology. considered with great caution. The sulcus in Brasileodactylus extends to the Comparison with the mandible in anteriormost tip, lacks the associated ridges and Coloborhynchus piscator (Kellner & Tomida has small anteriolaterally extending side sulcii. 2000) is limited because the mandible is not Consequently, the mandible can be classified fully prepared. The difference in size, how- to Anhanguera sp. The differences between ever, is great. Although not fully ossified, the the present mandible and the compared mandible of this juvenile pterosaur is 533 mm Coloborhynchus mandibles are larger than the (Kellner & Tomida 2000), thus substantially differences between the present mandible and larger than the present specimen. The rami of the compared Anhanguera mandibles, but still the Leiden specimen of Coloborborchynchus minor. The relatively small anterior expansion, spielbergi are more strongly bent relative to the sulcus with raised rims and the relatively the present manbible and the symphysis is short symphysis, clearly distinguish the present longer. Furthermore, the Leiden mandible has specimen from the Coloborhynchus jaws. no clearly discernible sulcus. The mandible is The absence of mandibles in A. blittersdorffi 50 % larger than SAO 200602. This difference and A. santanae is sufficient reason to avoid a is probably not ontogenetic because all these specific classification. The reconstructed length specimens are adults. The second and third of the holotype of A. santanae as well as the pairs of alveoli are the largest in the Leiden width at the symphysis is comparable to SAO specimen. 200602 (in contrast to the slightly shorter and The mandible cannot be classified in the new smaller jaw of A. blittersdorffi), but this alone is genus Frey et al., 2003 because not a strong argument for a classification at spe- this genus lack a mandibular crest and no cies level. The problems distinguishing the vari- expansion is reported. ous toothed taxa from Brazil are clear (see also Comparison of the ceratobranchials is lim- Veldmeijer & Signore 2004); the systematics are ited. Taking into account the fact that compari- often based on one specimen only, despite the son is made with Brazilian pterosaurs only, fact that material is housed in various collections the only other examples of ceratobranchials all over the world. The additional description of reported are those in Coloborhynchus ararip- this seemingly unimportant material can never- ensis (Wellnhofer 1985) and Criorhynchus theless add important details to the discussion, mesembrinus (not published) but the cerato- as has been demonstrated in this paper. branchials in both of them are less complete

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ACKNOWLEDGEMENTS collections under their care. Material in Berlin We thank J. de Vos, D. Peters, T. Bürgin, W. remained inaccessible, unfortunately. The study Brinkmann and two anonymous reviewers for of various collections have been made possible reviewing the manuscript. The Natuurhistorisch due to the financial support to AJV by the Jan Museum Rotterdam and the National Museum Joost ter Pelkwijkfonds, Stichting Molengraaff of Natural History (Naturalis), Leiden are Fonds, Mej. A.M Buitendijkfonds and Mr. & thanked for their all-round support. Urs and Mrs. Endenburg; the study of the material in Sonja Oberli are thanked for the opportunity various collections in Japan was made possibly of studying the material in their collection. E. by the Netherlands Organization for Scientific Bucci is acknowledged for reworking the pho- Research (NWO). Due to the grant of the tographs and E. Endenburg for his continuous Egypt Exploration Society for studying archae- support and assistance. AJV is grateful to M. ological material in Cambridge, AJV was able Dorling, E. Frey, A.W.A. Kellner, H. Mayr, S. to study some of the type specimens from the Nabana, M.A. Norell, Y. Okazaki, M. Oshima, Cambridge Greensands, housed in th Sedgwick Y. Takakuwa, Y. Tomida, P. Wellnhofer, R. Museum, Cambridge. Wild for kindly allowing access to the

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