Martill, David M.

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ZittelianaAn International Journal of Palaeontology and GeobiologySeries B/Reihe BAbhandlungen der Bayerischen Staatssammlung für Pa lä on to lo gie und GeologieB28DAVID W. E. HONE & ERIC BUFFETAUT (Eds)Flugsaurier: pterosaur papers in honour of Peter WellnhoferCONTENTS/INHALTDedication 3PETER WELLNHOFERA short history of pterosaur research 7KEVIN PADIANWere pterosaur ancestors bipedal or quadrupedal?:Morphometric, functional, and phylogenetic considerations 21DAVID W. E. HONE & MICHAEL J. BENTONContrasting supertree and total-evidence methods: the origin of the pterosaurs 35PAUL M. BARRETT, RICHARD J. BUTLER, NICHOLAS P. EDWARDS & ANDREW R. MILNERPterosaur distribution in time and space: an atlas 61LORNA STEELThe palaeohistology of pterosaur bone: an overview  109S. CHRISTOPHER BENNETTMorphological evolution of the wing of pterosaurs: myology and function 127MARK P. WITTONA new approach to determining pterosaur body mass and its implications for pterosaur fl ight 143MICHAEL B. HABIBComparative evidence for quadrupedal launch in pterosaurs  159ROSS A. ELGIN, CARLOS A. GRAU, COLIN PALMER, DAVID W. E. HONE, DOUGLAS GREENWELL & MICHAEL J. BENTONAerodynamic characters of the cranial crest in Pteranodon 167DAVID M. MARTILL & MARK P. WITTONCatastrophic failure in a pterosaur skull from the Cretaceous Santana Formation of Brazil 175MARTIN LOCKLEY, JERALD D. HARRIS & LAURA MITCHELLA global overview of pterosaur ichnology: tracksite distribution in space and time 185DAVID M. UNWIN & D. CHARLES DEEMINGPterosaur eggshell structure and its implications for pterosaur reproductive biology 199DAVID M. MARTILL, MARK P. WITTON & ANDREW GALEPossible azhdarchoid pterosaur remains from the Coniacian (Late Cretaceous) of England 209TAISSA RODRIGUES & ALEXANDER W. A. KELLNERReview of the pterodactyloid pterosaur Coloborhynchus 219JUNCHANG LÜ, LI XU & QIANG JIRestudy of Liaoxipterus (Istiodactylidea: Pterosauria), with comments on the Chinese istiodactylid pterosaurs 229DAVID M. MARTILLFirst pterosaur remains from the Exu Formation (Cretaceous) of the Araripe Basin, Brazil     243ERIC BUFFETAUTLate Cretaceous pterosaurs from France: a review 249Zitteliana München, 31.12.2008 ISSN 1612-4138B 28 255 SeitenEditors-in-Chief/Herausgeber: Michael Krings, Gert WörheideProduction and Layout/Bildbearbeitung und Layout: Martine FockeBayerische Staatssammlung für Paläontologie und GeologieEditorial BoardA. Altenbach, MünchenB.J. Axsmith, Mobile, ALF.T. Fürsich, ErlangenK. Heißig, MünchenH. Kerp, MünsterJ. Kriwet, StuttgartJ.H. Lipps, Berkeley, CAT. Litt, BonnA. Nützel, MünchenO.W.M. Rauhut, MünchenB. Reichenbacher, MünchenJ.W. Schopf, Los Angeles, CAG. Schweigert, StuttgartF. Steininger, EggenburgRichard-Wagner-Str. 10, D-80333 München, Deutschlandhttp://www.palmuc.de/zittelianaemail: zitteliana@lrz.uni-muenchen.deFür den Inhalt der Arbeiten sind die Autoren allein ver ant wort lich.Authors are solely responsible for the contents of their articles.Copyright © 2008 Bayerische Staassammlung für Pa lä on to lo gie und Geologie, MünchenDie in der Zitteliana veröffentlichten Arbeiten sind urheberrechtlich geschützt. Nachdruck, Vervielfältigungen auf photomechanischem, elektronischem oder anderem Wege sowie die Anfertigung von Übersetzungen oder die Nut zung in Vorträgen, für Funk und Fernsehen oder im Internet bleiben – auch auszugsweise – vorbehalten und bedürfen der schriftlichen Ge neh mi gung durch die Bayerische Staatssammlung für Paläontologie und Geologie, München.ISSN 1612-4138Druck: Gebr. Geiselberger GmbH, AltöttingCover Illustration: Modell eines Rhamphorhynchus aus dem Oberjura von Eichstätt. Entwurf: P. Wellnhofer, Modell: R. Liebreich, Foto und Collage: M. Schellenberger, L. Geißler, BSPG München.Umschlagbild: Reconstitution of a Rhamphorhynchus from the Upper Jurassic of Eichstätt, Bavaria. Concept: P. Wellnhofer; design: R. Liebreich; photograph and collage: M. Schellenberger, L. Geißler, BSPG Munich. 243AbstractA fragmentary pterosaur proximal humerus from the basal member of the Exu Formation (Cretaceous) of Piauí, Brazil rep-resents the fi rst pterosaur element from this formation and may even be the fi rst macrofossil from this dominantly arenaceous horizon. The fragment is tentatively assigned to Lophocratia UNWIN, 2003. The preservation is inconsistent with it having been reworked from the famously pterosaur-rich Santana For-mation which under lies the Exu Formation in the region.Key words: Cretaceous, Pterosauria, Exu Formation, BrazilZusammenfassungDer fragmentarische proximale Teil des Humerus eines Pterosauriers aus dem basalen Glied der Exuformation (Krei-dezeit) von Piaui, Brasilien, repräsentiert das erste Pterosau-rierelement aus dieser Formation. Es könnte sogar das erste Makrofossil aus diesem mächtigen Quarzsandsteinhorizont sein. Das Fragment wurde vorläufi g Lophocratia UNWIN, 2003 zugeordnet. Die Erhaltung des Fossils ist schlecht, da es aus der für ihren Pterosaurierreichtum bekannte Santana Formation, dem Liegenden der Exu Formation in der Region, aufgearbeitet und umgelagert wurde. Schlüsselwörter: Kreide, Pterosauria, Exu Formation, Brasilien1. IntroductionThe Araripe Basin of north east Brazil contains a largely Cretaceous sedimentary infi ll in which two fossil Konservat Lagerstätten occur, the Aptian Nova Olinda Member of the Crato Formation (MARTILL & FREY 1998; UNWIN & MARTILL2007) and the ?Albian Romualdo Member of the Santana Formation (MARTILL 1993). Both of these horizons are well known for the exceptional preservation of fossil vertebrates (MARTILL 1988, 1997; WENZ & BRITO 1990). The sequence is capped by the massively bedded Exu Formation sandstone with conglomerates that form an extensive plateau known as the Chapada do Araripe. The precise age of the Exu Formation is unknown, but LIMA(1978) suggests a mid Albian age on the basis of palynomorphs. However, recent revisions of the age of the underlying Santana Formation based on palynomorphs suggests that the Exu For-mation is probably somewhat younger (U. HEIMHOFER, pers. com. 2007). Although the formation is an important aquifer and supports a productive agriculture, its stratigraphy and sedimen-tology remain largely unstudied and the internal stratigraphy of the Exu Formation remains to be established. There appears to be a gradational coarsening upward from mudstones of the Santana Formation to the sandstones of the Exu Formation in the east of the region, but widespread slumping and jungle growth at the contact means that the basal boundary of the Exu Formation is rarely exposed. The situation is somewhat diffe-rent in the west, where a basal unit of highly oxidised laminated mudstones, siltstones and fi ne sandstones termed the Simões Member by MARTILL & WILBY (1993) occurs widely, before passing up into massive sandstones with conglomerates. This formation has largely been regarded as unfossiliferous but, during fi eld work in 2005, a fragment of a pterosaurian humerus and plant remains were collected from exposures of the Exu Formation in the western Araripe Basin. The humerus, described here, apparently represents the fi rst vertebrate fossil from this formation.2. LocalityThe pterosaur humerus was collected from slumped exposu-res of the Exu Formation on the western fl anks of the Chapada do Araripe between Marcolandia in Pernambuco and Simões in Piauí. This is close to locality 23 of MARTILL (1993: 129). These exposures are slumped masses of light brown Exu Formation Zitteliana B28 243 - 248 4 Textfi gs, 1Pl. München, 31.12.2008 ISSN 1612 - 4138First pterosaur remains from the Exu Formation (Cretaceous) of the Araripe Basin, BrazilByDavid M. Martill*Palaeobiology Research Group, School of Earth and Environmental Sciences, University of Portsmouth, Portsmouth, PO1 3QL, United KingdomManuscript received November 9, 2007; revised manuscript accepted February 28, 2008.*E-mail: David.Martill@port.ac.uk244sandstones that have overridden the mudstones of the Santana Formation. The locality is easily accessed by scrambling down the bank at the side of the asphalted road (Textfi g. 1). A small stream dissects the slumped masses of sandstones and a net-work of small paths provide access through the thorny scrub. Exposure is excellent, but determining the precise stratigraphic relationships of each slumped mass is diffi cult.3. StratigraphyDue to the extensive landslip of the western fl ank of the Chapada do Araripe details of the stratigraphy are diffi cult to elucidate. Nevertheless, it is possible to identify the concreti-on-bearing Romualdo Member of the Santana Formation at the locality on account of its abundant laminated carbonate concretions with fossil fi sh and, as everywhere, the bright pink sandstones of the Exu Formation make up the escarpment face of the Chapada. Field brash of silicifi ed laminated limestone may represent the Crato Formation, but no in situ material was found. A simplifi ed stratigraphy for the Cretaceous part of the Araripe Basin in this region typically comprises black shales and laminated limestone that can be referred to the Crato Formation, which is overlain by bedded gypsum of the Ipubi Formation. A disconformity usually marks boundary between the Ipubi Formation and the overlying Santana Formation. There then follows a unit of fi nely laminated silts and fi ne sands, the Simões Member that near Araripina may reach 5 m in thickness. These laminated silts and sands can be identifi ed at the locality, but their relationship to the underlying Santana Formation cannot be observed.  Above the Simões Member are 5 or more metres of well bedded buff coloured sandstones with small (2–10 cm in diameter) dark brown ironstone concretions. These are especially common on weathered ground where they accumulate as a lag (Plate 1) and it is in these concretions that fossils have now been located in the Exu Formation. The buff coloured sands are overlain by several metres of bright pink sandstones and conglomerates that comprise the remainder of the Exu Formation. Although a complete stratigraphic log cannot be constructed for the locality without considerable effort, a simplifi ed stratigraphy is presented in Textfi gure 2.4. Palaeontology4.1 Systematic palaeontologyPterosauria KAUP, 1834Pterodactyloidea PLIENINGER, 1901Lophocratia UNWIN, 2003Lophocratia Indet.Specimen: Fragment of proximal right humerus lacking the humeral head. Number SMNK PAL 6414 in the collec-tion of the Staatliches Museum für Naturkunde Karlsruhe, Germany.Locality: Natural exposure on road approximately 8km east of Simões, Piauí, north east Brazil. Map reference SB.24-Textfi gure 1: Sketch map of the new locality where slumped masses of the Exu Formation sandstones yield rare macrofossils. The cross in a circle marks the site where the partial pterosaur humerus was collected.245Plate 1: New fossil locality for the Exu Formation (Cretaceous) of the Araripe Basin, Brazil. (1) View of the western escarpment of the Chapada do Araripe (background) with landslipped mass of buff coloured sands (foreground) that yield the fi rst Exu Formation vertebrate fossils; (2) Weathered face of the buff sandstones with ironstone concretions and harder cemented patches weathering out.246Y-C-VI  308 9160.Horizon: Unnamed member of the Exu Formation, above the Simões Member of MARTILL & WILBY (1993).Age: Upper Cretaceous, most probably younger than Albian.4.2 DescriptionThe humeral fragment measures 33 mm in length, with a maximum proximal diameter of 47 mm and 18 mm distally. The bone is preserved as an iron oxide (presumed to be goethite on account of its dark brown colour) cemented fi ne sandstone infi ll of the void space of the humerus. All cortical bone has fl aked away to reveal an internal mould of the bone with trabe-culae intact. There is no crushing and the missing termini have clean fractures suggesting that more of the bone was originally preserved. Although the cortical bone is missing, there is no evidence of wear due to abrasion, but the trabeculae are incised perhaps due to some slight chemical weathering. The exposed trabeculae are widely spaced, forming irregular polygonal cells that become denser proximally (Textfi g. 4a,f) with individual trabeculae approximately 0.75 mm thick. The cross-sectional shape at the distal termination is a rounded asymmetrical triangle (Textfi g. 4b), whereas the cross section is a fl attened ‘D’ shape proximally (Textfi g. 4e). The palmar surface expands proximally equating to the base of the deltopectoral crest (Textfi g. 4c,d). Fossil plant material was also found at this locality and is enclosed within ironstone concretions, but the bone lacks any ironstone overgrowths.5. DiscussionThe internal structure of this bone fragment is typical of the proximal and distal parts of the larger long bones of pterodac-Textfigure 2: Simplified stratigraphic log for the slopes of the Chapada do Araripe between Marcolandia and Simões. Bed thicknesses are approximate as widespread slumping prevents accurate measurements.Textfi gure 3: Pterosaur humeri. A, ornithocheirid type; B, lophocratan humerus, Tapejara wellnhoferi with the Exu Formation fragment shaded in dark grey. A, based on WELNHOFFER (1991); B, based of FREY et al. (2003: fig. 4B’ SMNK PAL 1137). 247Textfi gure 4: Partial pterosaur proximal humerus from the lower units of the Exu Formation between Marcolandia and Simões, Piauí, Brazil. Specimen no. SMNK 6414. a: detail of trabeculae, b: proximal view, c: ventral fascia, d: dorsal fascia, e: distal view, f, cranial fascia.tyloids (DE RICQLÈS 2000). The gentle proximal expansion of the diaphysis into the deltopectoral crest is reminiscent of the condition found in the humeri of the lophocratian pterosaurs Tapejara (Textfi g. 4b) and Lonchodectes (UNWIN 2003: fi g. 17j), rather than the near 90 degree defl ection seen in orni-thocheirids. Although referral here to Lophocratia UNWIN, 2003 is speculative given the highly fragmentary nature of the specimen, it is worth noting that the lophocratians TupuxuaraTapejara, Tupandactylus and possibly Cearadactylus occur in the underlying Santana and Crato Formations (UNWIN & MAR-TILL 2007). Regardless, despite the lack of taxonomic precision, the discovery of this pterosaur fragment is signifi cant in two respects. Firstly, it demonstrates that body fossils, although clearly rare, do occur in the Exu Formation. Perhaps this locality represents an unusual facies of this dominantly sandy unit, and further work should be undertaken to determine the geographical distribution of this fossiliferous horizon. Secondly, the preservation, although far from ideal, is of an uncrushed bone that is not enclosed within a calcareous concretion. Although the possibility exists that originally the 248fragment was indeed enclosed in a concretion, and subsequent weathering released the fossil, perhaps with the bone compacta remaining on the concretion, it seems more parsimonious that the bone was preserved in three dimensions in uncompacted sandstone. If this latter scenario is correct, then it is most likely that the pterosaur bone is at least penecontemporaneous with the sediment and does not represent an element reworked from the underlying Santana Formation. If this is the case, then this specimen represents another occurrence of a pterosaur in a non-marine horizon.AcknowledgementsI am indebted to Artur ANDRADE of the DNPM, Crato for assistance in Brazil and to Robert LOVERIDGE for photogra-phing the specimen. Thanks to Mark WITTON for helpful dis-cussion and Dan BONTER, Sam HEADS, Natalie MITCHELL and Judith WOHLRABE for help in the fi eld. Thanks to Dave HONEfor assisting with the refereeing and editorial process.6 ReferencesFREY, E., BUCHY, M.-C. & MARTILL, D. M. (2003): Middle- and bot-tom-decker Cretaceous pterosaurs: unique designs in active fl ying vertebrates. – In: E. BUFFETAUT & J.-M. MAZIN (Eds), Evolution and Palaeobiology of Pterosaurs. – Geological Society of London Special Publication, 217: 267–274.KAUP, J. J. (1834): Versuch einer Eintheilung der Saugethiere in 6 Stäm-me und der Amphibien in 6 Ordnungen. – Isis 3: 311–315.LIMA, M. R. (1978): Microfosseis da Formação Exu. Cretáceo do Nordeste do Brazil. – Annais XXX Congresso Brasileira Geo-logia, 2: 965–969.MARTILL, D. M. (1988): The preservation of fossil fishes in concretions from the Cretaceous of Brazil. – Palaeontology, 30: 1–18.MARTILL, D. M. (1993): Fossils of the Santana and Crato Formations, Brazil. –  The Palaeontological Association Field Guides to Fos-sils, 5: 1–159.MARTILL, D. M. (1997): Fish oblique to bedding. – Palaeontology 40:1011–1026.MARTILL, D. M., BECHLY, G. & LOVERIDGE, R. F. [Eds] (2007): The Crato Fossil Beds of Brazil: window into an ancient world; Cam-bridge (Cambridge University Press), 624 pp.MARTILL, D. M. & FREY, E. (1998): A new pterosaur Lagerstätte in N. E. Brazil (Crato Formation: Aptian, Lower Cretaceous): preliminary observations. – Oryctos 1: 79–85.MARTILL, D. M., & WILBY, P. R. (1993): Stratigraphy. – In: D. M. MARTILL (Ed.), Fossils of the Santana and Crato Formations, Brazil. – The Palaeontological Association Field Guides to Fos-sils, 5: 1–159.PLIENINGER, F. (1901): Beiträge zur Kenntnis der Flugsaurier. – Palae-ontographica 48: 65–90.RICQLÈS, A. J. DE, PADIAN, K., HORNER, J. R. & FRANCILLON-VIELLOT, H. (2000): Paleohistology of the bones of pterosaurs (Reptilia: Archosauria): anatomy, ontogeny, and biomechanical implications. – Zoological Journal of the Linnean Society, 129: 349–385.UNWIN, D. M. (2003): On the phylogeny and evolutionary history of pterosaurs. – In: E. BUFFETAUT & J.-M. MAZIN (Eds), Evolution and Palaeobiology of Pterosaurs. – Geological Society Special Publication, 217: 139–190.UNWIN, D. M. & MARTILL, D. M. (2007): Pterosaurs from the Crato Formation. – In: D. M. MARTILL, G. BECHLY & R. F. LOVERIDGE(Eds), The Crato Fossil Beds of Brazil: Window into an Ancient World; Cambridge (Cambridge University Press), 475–524.WENZ, S. & BRITO, P. M. (1990): L’ichthyofaune des nodules fossili-fères de la Chapada do Araripe. – In: D. DE A. CAMPOS, M. S. S. VIANA, P. M. BRITO & G. BEURLEN (Eds), Atas do simpósio sobre a Bacia do Araripe e Bacias Interiores do Nordeste, Crato (14–16 de Junho de 1990), 337–349.WELLNHOFER, P. (1991): Weitere Pterosaurierfunde aus der Santana-Formation (Apt) der Chapada do Araripe, Brasilien. – Palaeon-tographica A, 215: 43–101.

(1834): Versuch einer Eintheilung der Saugethiere

A new pterosaur Lagerstätte in N. E. Brazil (Crato Formation: Aptian, Lower Cretaceous): preliminary observations.

Atas do simpósio sobre a Bacia do Araripe e Bacias Interiores do Nordeste,

Beiträge zur Kenntnis der Flugsaurier.

Fish oblique to bedding.

Fossils of the Santana and Crato Formations,

L’ichthyofaune des nodules fossilifères de la Chapada do Araripe. – In:

Microfosseis da Formação Exu. Cretáceo do Nordeste do Brazil.

Middle- and bottom-decker Cretaceous pterosaurs: unique designs in active ﬂ ying vertebrates.

On the phylogeny and evolutionary history of pterosaurs.

Paleohistology of the bones of pterosaurs (Reptilia: Archosauria): anatomy, ontogeny, and biomechanical implications.

Pterosaurs from the Crato Formation. – In:

The Crato Fossil Beds of Brazil: window into an ancient world; Cambridge (Cambridge

The preservation of fossil fishes in concretions from the Cretaceous of Brazil.

Weitere Pterosaurierfunde aus der SantanaFormation (Apt) der Chapada do

First pterosaur remains from the Exu Formation (Cretaceous) of the Araripe Basin, Brazil

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Martill, David

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First pterosaur remains from the Exu Formation (Cretaceous) of the Araripe Basin, Brazil

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