Panorama of the Flying Reptiles Study in Brazil and South America * (Pterosauria Pterodactyloidea|Anhangueridae) DIOGENES DE ALMEIDA CAMPOS and ALEXANDER WILHELM ARMIN KELLNER Segio de Paleontologia, Divisio de Geologia e Mineralogia, Departamento Nacional da Produc Mineral, 22290) Rio de Janeiro, RJ (With 3 plates and 1 text-figure) INTRODUCTION The study of the flying reptiles in South America has developed very much in the last two decades. This is due to the fact that more and more remains of these animals have been found in this continent, especially in Brazil and Argentina. Therefore, a lot of publications on this subject have been made recently. Unfor- tunately, the South American pterosaurs are not yet widely known as the greatest part of the material found is incomplete and sometimes very fragmentary. Up to now, only one com- plete specimen has been found. The principal objective of this study is the presentation of the results of the various papers published regarding the South American pterosaurs. A preliminary description and diagnosis on a new flying reptile has also been made in order to contribute to the study of this very important and interesting group of animals which in former times habitated several areas of the Southern American Continent. THE FLYING REPTILES IN BRAZIL The first reference to flying reptiles in the Southern Hemisphere was made in the last cen- tury. This was based on two quadrate bones discovered by Joseph Mawson in rocks of the “Bahia, group”, that. is in Pedra Furada, 604. -Aecepted for publication July 18, 1985; presented by IoNacio MacHADO BRITO. 44 (j3 icons Salvador, in the State of Bahia, Brazil. Wood- ward (1891) studied these bones and concluded they looked like the ones attributed to the Rhamphorhynchus manseli, of the Kimmeridge Clay, in the British Islands. As this material was insufficient, he did not make a_ generic classification. Some years later, Mawson discovered in the same place a quadrate bone, approximately three times larger than the first one. Woodward (1896) classified it as belonging to the greatest pterosaur attributed to the sub-order Rham- phorhynchoidea. Times later, while examining. more com- plete material of the “Recéncavo Baiano”, Woodward was able to determine that the quadrate bone — once classified by him as be- ing part of a gigantic pterosaur —~ in reality belonged to a celacantidas fish (Mawsonia gigas Woodward, in Mawson & Woodward, 1907). In the same publication, this autor maintains the existence of the pterosaurian quadrate bones described by him in 1891 and added the discovery of a typical pterosaurian tooth made later by Mawson in the same region. In the English edition of Zittel (Text-Book of Paleontology, v. 2), which was revised by Woodward in 1932, is again mentioned the ex- istence of a small pterosaurian quadrate bone from the Lower Cretaceous of Bahia. This time, this presumed. pterosaurian - rémain’ was classified inthe Pterodactyloidea Plieninger, 1901.21 ai laity Soaugudl BOECIENCIAS fos cox at.) siT siT sir 518 519 520 523 524 524 524 525 526 526 526 oct 30 1985 LIBRARIES ANAIS DA Vol. 57 CIENCIAS 1985 ISSN 0001 — 8765 ACADEMIA BRASILEIRA Ne 1Roxo (1936) called attention to the correc- tion made by Woodward in 1907, but didn’t mention the note made by the later on the oc- currence of the pterosaurian tooth. This author also considered improbably the existence of the Tittle quadrate bone mentioned by Woodward in the English edition of Zittel.. Price (1953b) considered the material refer- red by Woodward as being pterosauria was im- proper and probably not from’ flying reptiles: Alll the above mentioned material is exposed in the fosssil collection of the British Museum of Natural History. Another mention made of Brazilian flying reptiles was made by Barbosa Rodrigues (1892). ‘As Price (1953b), we think that this reference has no cientifical support. The first doubtless evidence of the presence of flying’ reptiles’in the Southern ‘American Continent was given by Price (1953b). It was due to a left humerus which was discovered in the “Fazenda do Congo”, situated on the right margin of the Gramame river, in the sedimen- tary rocks of Upper Cretaceous age. This material ‘pertains to. a great - collection originating from that region and that was put together by colonel Jo’o Domingués dos Santos and was studied by Maury (1930). Two osseous fraginents and three teeth were found there. One of the bone fragments and the teeth were classified by Professor R. S. Lull and by Dr. M. R. Thorpe, both of the, Department of Vertebrate Paleontology of the Yale University as being crocodilian remains. They assured that all three teeth definitely belonged to Goniopho- Jis hartti (Marsh).The bone fragment, ‘which is illustrated in Maury, (1930 — plate II, fig. 1), was said to, be.correlated to crocodiles, ap- parently being the posterior region of a skull d classified in this species. ‘The second: osseous fragment did not call much attention and Price (1953b),only found a little note from Maury saying that it-was not a cephalopod, Ved castt ‘<4 When Price:(1953a) made.a short revision ofthe vertebrate fauna of the Gramamie Forma- tion, he disagreed that the first osseous. frag- 1D. A. CAMPOS and A. W. A. KELLNER ment belonged to a crocodilian skull and con- sidered it as indeterminable. _ Times later, Price (1953b) noted that the two osseous fragments belonged to the same bone and formed a left humerus of a flying rep- tile. This material is at present deposited in the fossil collection of the Paleontological Section of the National Department of Mineral Produc- tion, DNPM, under the number DGM 238-R. Due to the close resemblance of the humerus to various specimens ‘attributed’ to the genus Nyctosaurus (Marsh, 1876), of the family Pteranodontidae, Price classifified this material in this genus. Expecting a more complete material collection in order to be able to make a more exact classification with — perhaps — the creation of a new genus,, this author denominated this new form as N. Jamegoi. The creation of a new species was based on the geographical location of the material and the very high position where it was found in the cretaceous column (Maastrichtiano). Further to this, Price observed that the humerus of N. Jamegoi was about one third larger than the largest species recorded regarding this genus (N. gracilis Marsh, 1876). The wing-spread of the Brazilian form was estimated around 4m. The second record of flying reptiles in Brazil comes from Lower Cretaceous sediments, the Santana Formation, in the Chapada do Araripe, Ceara. This material was found in a calcareous nodule and consists of parts from the right anterior member. It is deposited in the collections of the Paleon- tological Section of the National Department of Mineral Production, under the. number DGM 529-R, and formed by the distal parts of the radius and ulna; proximal and lateral carpal; pteroid and metacarpal, proximal part of the first alar phalanx and four phalanxes ‘of the digits I to III. The classification and description of this material was again made by Price (1971), who ‘determinated the formas Araripesaurus castilhoi, “and ‘included it ‘in ‘the ‘family Or- nithocheiridae. For this classification, some in- ‘formation obtained from two: bony. fragments was also used by Price.*These fragments con-PANORAMA OF THE FLYING REPTILES STUDY IN SOUTH AMERICA sists of the distal portion of an ulna an the prox- imal part of a metacarpal IV, and belong to the private collection of Dr. Umberto Silveira Con- sentino, from Piracicaba, Sdo Paulo. They are registered under the numbers RG-147 and RG-212. Plastic moulds were made of both fragments (DGM 549-R; DGM 550-R) and were deposited together with the holotype. The estimated wing-spread of this ‘example is about two meters. Some years later, the first falange of the 4th finger from a right anterior member of a flying reptile was discovered. This material came again from the Santana Formation and is deposited in the collection of the Baviera State Collection of Paleontology and Historical Geology, Muenchen (Nr. 1975 1 166). Based on this material, Wellnhofer. (1977) proposed a new flying reptile in that region, which he called Araripedactylus dehmi. The author emphasized the enormous thickness of the bony walls, which reaches 3 to Smm. However, considering this material insufficient, Wellnhofer did not make a family classification of this form. The wing-spread of this pterosaur was estimated at about 5 meters. , Posteriorly Buisonjé (1980) described an ankylosed dextral scapula-coracoid and the dex- tral humerus and determinated a new flying rep- tile from the Santana Formation: Santanadac- tylus brasilensis. This pterosaur was classified in the family Criorhynchidae, and the material can be found in the collection of the Geological Institute of the University of Amsterdam, Holland, under the number M. 4894. In the same ,lecality were also found two cervical vertebrae, which brought Buisonjé to the con- clusion that S. brasilensis had a long neck and was extremely adapted to gliding rather than to flapping flight. The wing-spread of this pterosaur was estimated around 6 meters. One of the most fascinating and important discoveries regarding flying reptiles was made again in the Santana Formation.’ This was a comiplete. skull (Campos, 1983) which is being studied by the National Department of Mineral © Production.’ It ‘is; the’ first’ registration ‘ofa 455 pterosaur. witha conspicuous sagittal ‘crest situated on the most distal part of the skull. Besides, it also Has a little sagittal crest on the posterior’ part of the skull. This paper informs on the results of the preliminary study on this material, and a new family (Anhangueridae), genus and species (Anhanguera blittersdorffi) is proposed. This skull belongs to the private col- lection of Mr. Rainer A. V. Blittersdorff (Collection Desirée — Rio de Janeiro) and some moillds can be found in the Paleontological Sec- tion of the DNPM. A second skull with mandible coming from the same region as the first one was described by Leonardi & Borgomanero (1983). These authors suggested the creation of the form Cearadac- tylus atrox having made no classification regarding the family. Contrary to the skull, the mandible is complete. This material is deposited in the private collection of Borgomanero (Curitiba, Pr). On the mandible and maxillar a natural notch can be observed forming a cavity for storage of food, probably fish. In this region, the alveolar borders of the premaxillae do not oéclude with the jaw, leaving a wide gap. This spatulated rostral end is one of the most important distinctive characteristics of this pterosaur, whose wing-spread should be about 4 meters. Another form of pterosaur from the San- tana Formation was proposed by Kellner (1984a), based on a mandibular symphysis. It was named Brasileodactylus araripensis, pro- visory classified in the family Ornithocheiridae. The arched form and the enlargement of the distal patt of the mandible, with the formation of a plane surface are some of the characteristics that individualize this flying rep- tile. It should have been of a considerable size and its alimentation probably consisted of fish. More material of the manbible of Brasileodac- tylus was again described by Kellner (1984b). Wellnhofer. et al. (1983) describe a vertebral column with the notarium, that was again found in. the:Santana Formation. This material is available in the collection of Dr. Paul Gigase, ‘in Antwerpen, ‘under, the number In: Acad: brasil: Ciéne.; (1985) 57 (8)456 ‘V-201. According to the authors, this material; normally could not be classified into the.forms ‘A: castilhoi, A. dehmi and S. brasilensis. Still, in order to avoid the creation of a new name for the flying reptiles of the Santana Formation, they: classified this ‘material as. S.: brasilensis (family indeterminated). Besides, the size of the two: cervical vertebrae of the S. ‘brasilensis described by Buisonjé (1981) was about ithe: same as the. last cervical vertebrae of the material studied by Wellnhofer et al., which was the only reason for their classification. Another very interesting remain of a pterosaur also found in the Santana Formation was described by Campos et al. (1984). It con- sists of the limb bones of a flying reptile, which are preserved in connection, formed of a por- tion of the right femur, tibia, fibula, metatar- sals, phalanxes of a right foot, as well as the elements of a folded wing in a Z form with radius, ulna, carpals, pteroid, the metacarpal IV and the phalanxes of the fourth finger. One of the most important facts of this occurrence is the perfect preservation of the wing membrane, this apparently being the first record of this nature in the Southern American Continent. _ Besides these publications, it is worth to mention the reference made by Campos (1983), of a typical pterosaurian tooth, found in former collection in Bahia, made by Orville A. Derby; this tooth is deposited in the National Museum in Rio de Janeiro. So far, these are the most important studies made on the Brazilian pterosaurs. We know of the “existence of other pterosaurian remains, probably coming from the Santana Formation, being kept in the National Department for Mineral Production, West Germany and in private collections. ” THE FLYING REPTILES, IN ARGENTINA ‘The first record of pterosaurs in Argentina comes from the region of Hualtarn, in the oriental’ part! of ‘the'Serra de“Las Quijadas, ‘situated’ at’ NW, ‘in’ the ‘Lagarcito''Formation (Upper Jurassic/Liower! Cretaceous); ::This material’ can‘be‘ found jin’ the collection) of the jad. Brasil: Clene., (1985) 57.(8) suD.'As CAMPOS: ‘and, A’ W. APKELLNER> js Fig. 1 — South America with all known pterosaur localities; taken from Kellner (1984a), complemented. BRASIL — 1) Gramame Formation, Parafba: Nyctosaurus lamegoi Price, 1953; 2) Santana Formation, chapada do ‘Araripe, Cear&: Araripesaurus.castilhoi Price, 1971; Ataripedactylus dehmi Wellnhofer, 1977; Santanadactylus brasiliensis Buisonjé, 1980; Cearadactylus atrox Leonardi & Borgomanero, 1983; Brasileodactylus araripensis Kellner, 1984; Anhanguera blittersdorfti Campos & Kellner, 1985; 3) Bahia Supergroup, Recéncavo, Bahia: Pterosaurian tooth (2) Campos, 1983. ARGENTINA — 4)'Lagarcito Formation, San Lui Pterodaustro guinazui Bonaparte, 1970; 5) Santa Cruz For- mation, San Luis: Puntanipterus globosus Bonaparte & Sanchez, 1975. CHILE’ — 6) Neojurassic (2) sediments, Cordillera de Domeyko, Antofagasta: Pterodaustro sp. Casamiquela & Diaz, 1980. Institute and Foundation" Miguel Lillo, Na- tional University of Tucumén. It is composed of anearly complete right humerus (PVL 2571); one, cervical vertebra: (PVL. 2574); a cranium fragment (PVL 2586); a scapula coracoid (PVL 2585); proximal part f a left tibia-fibula (PVL 3403); two, dorsal vertebras (PVL 2575);:an in-PANORAMA OF THE FLYING REPTILES STUDY IN SOUTH AMERICA complete femur (PVL 2572); and an incomplete Phalanx of the 4 digit (PVL 2573). These pterosaurian remains were studied by Bonapar- te (1970) who proposed the form Pterodaustro guinazui, initially classified in the family Pterodactyloidea. A year later, Bonaparte (1971) studied a skull with mandible and a fragment of the anterior part of another skull depositated in the collection of the Municipal Museum of Natural Sciences of Mar del Plata (MMP 1018 and MMP 1035). He proposed the creation of a new family Pterodaustridae (Pterodactyloidea) with the type-species P. guinazui. The principal characteristic of this family is the dentition with filiform teeth and the singular shape of the skull. In the following years other skulls of P. guinazui were found, especially two which can be found in the collection of the Municipal Museum of Natural Sciences of Mar del Plata under the numbers MMP 1087 and MMP 1086. Additionally an almost complete skeleton was also found and deposited in the collection of the Institute and Foundation Miguel Lillo of Na- tional University of Tucuman (PVL 3860). This material was studied by Sanchez (1973) who redescribed the form P. guinazui. In the same region of Hualtaran, remains of another pterosaur coming from the La Cruz Formation (Upper Jurassic/Lower Cretaceous) were found. This material can be seen in the col- lection of the Miguel Lillo Institute and Funda- ton. It consists of a tibia-fibula (PVL 3869), an alar phalanx of a posterior member and a dorsal vertebra. Based on this material, Bonaparte & Sanchez (1975) created the form Puntanipterus globosus. These are all the publications on flying rep- tiles coming from Argentina that we know. The region of Hualtaran, San Luis, proved to be very rich in remains of pterosaurs; they should be studied more thoroughly. THE FLYING REPTILES IN CHILE »Up to present date, we only know one oc- currence of ,pterosaurian. remains. -in., Chile, 457 which come from Neojurassic sediments of the cordillera of Domeyko in the Antofagasta pro- vince." This’ material,° which was studied by Casamiquela & Diaz (1980), includes diverse bone fragments, that are kept in the “Oficina Regional Antofagasta” of the Geological In- vestigation Institute of Chile (nr. 250973, 1-39). The cranial bones also examinated by these authors were not numerated. Based on parts of the rostrum (part of the maxilas provided with teeth) a mandibular rama also dentated and a humerus, the authors determinated that this material belonged to a pterosaur of the genus Pterodaustro (Bonaparte, 1970). Because of the fragmentary condition of this material, Casami- quela & Diaz did not propose a new designation. As for now, those were all the publication about the flying reptiles in South America. At the end of this paper, we present a map showing all the pterosaurian localities in the Southern American Continent (Plate 4), and Table I with the summary of the remains from all different Pterosaurs found in this continent. A NEW FLYING REPTILE FROM BRAZIL The pterosaurian skull object of this study was found in a calcareous nodule from the San- tana Formation, chapada do Araripe, Ceara. It is part of a private collection which belongs to Rainer Alexander von Blittersdorff (Desirée Collection), Rio de Janeiro, who permitted the study of this material in the Paleontology Sec- tion of the 9th District of the National Depart- ment of Mineral Production. The material was given to us entirely covered on two thirds of its area by the calcareous mass, and was prepared manually (i. e. without using acid) by Otavio da Silva San- tos, who has made a fabulous job in-spite of its fragility. In general the skull is very well preserved, being complete on the right side and only miss- ing a small part of the maxilla/premaxilla in the more distal region of the left side..The bones “have an average, thickness of 2mm. We can point out. the existence of a small, parietal crest ‘An Acad, brasil, Clénc, (1985) 57 (4)458 D. A. CAMPOS and A. W. A. KELLNER TABLE 1 Summary of All the Pterosaurian Remains Which Were Found in South America Axiat, | ANTERIOR PosTERton} Forms SeuLt_ [Manornte Toor ‘OnseRvaTion SkELETON| Litas} Limes Nyctosaurus lamegoi - - - | part of a humerus Araripesaurus castilhoi - - ~ | radius, ulna and carpus Araripedactylus dehmi - - - ~ | first wing phalanx Santanadactylus parts of a scapula brasilensis - - x - | coracoid, humerus, vertebrae Cearadactylus part of a skull with atrox x x - x — | mandible Brasileodactylus araripensis - . - - x | incomplete mandible Anhanguera blittersdorffi x 7 - - x | complete skull Tooth in the National Museum | ~ - - - x Wing in study preserve the impression in France - - - x = | of the wing membrane Pterodaustro guinazui x x x x x | complete materials Puntanipterus tibia and fibula, phalanxes slobosus - - x x ~ | and dorsal vertebrae Prerodaustro sp. x x - x | very fragmented material on the posterior part of the skull and a thin and long sagittal crest on the anterior part of the skull, never before mentioned in this position until this work, REGION OF ORIGIN OF THE MATERIAL Unfortunately the exact site where the bones were found is unknown. All we know, however, is that this material came from calcareous sediments of the Chapada do ‘An. Acad. brasil. Ciéne., (1985) 57 (4) Araripe, probably of a flank in the state of Ceara, NE Brazil. ‘The stratigraphy of the Chapada do Araripe has been studied by many researchers, and we can point out the works of Small (1913), Beurlen (1963, 1971), Braun (1966), Mabe- soone & Tinoco (1973) and Campos & Wenz (1982). A summary of the stratigraphical sequence is as follows:PANORAMA OF THE FLYING REPTILES STUDY IN SOUTH AMERICA — Exu Formation: formed by sand- stones, supposed to be of fluvial origin and of Albian age; — Santana Formation: divided in three members: Crato (limestone of lacustrine origin), [pubi (gypsum layer, that indicates the opening of the lake to the sea) and Romualdo (formed by calcareous beds and marts, very rich in fossils, represents the return to lacustrine conditions). This stratigraphic unit is con- sidered of Lower Cretaceous age (Aptian- Albian); —~ Missdo Velha Formation: formed by intercalations of sandstones and clays, with locally some schistes pyrobituminous and of Neocomian age; — Brejo Santo Formation: represented by alternations of pelite and red-whitish clays, thought to be of continental origine and of Superior Jurassic age; — Cariri Formation: _ situated on the base of the sequence, it is formed by conglomerates and conglomeratic sandstones, dated as Devonian. As one of the richest and important fossile locality in the world, the Santana Formation is the most important strata in Chapada do Araripe region. There has been found many kinds of fossils, some of them very well preserv- ed, like echinoderms, lamellibranches, gastropods, conchostracans, ostracodes, plant remains, pollens, crustaceans, insects, arachnids, fishes and reptiles. The material studied in this paper is also thought to be from that stratigraphic unit, specifically from the Romualdo Member. The reptilian fauna known in the Santana Formation is represented by a crocodilian (Araripesuchus gomesii Price, 1959), a turtle (Araripemys barretoi Price, 1973) and dinosau- rian remains (Leonardi & Borgomanero, 1981; Campos, 1985). There are also some occur- rences of flying reptiles, as we have seen in the first part of this paper. With the present work we are adding the description of a new pterosaur, Anhanguera 459 blittersdorffi and introduce a new family, Anhangueridae. SYSTEMATICS Sub-ordo: PTERODACTYLOIDEA Plieninger, 1901 Familia: $ANHANGUERIDAE nov. fam. Genus typicus: Amhanguera nov. gen. Type-species: Amhanguera blittersdorffi, type by monotypy. DIAGNOSIS: Pterosaurs with a large sagittal crest on the anterior part of the skull, situated on the premaxillas, which ends almost at the beginning of the external naris; presence of a small parietal sagittal crest on the posterior part of the skull; dentition from distal part of the skull on to the middle region of the naris preor- bital opening, corresponding to the beginning of the internal naris; presence of an enlarge- ment of the distal part of the skull, where the premaxillar teeth, the biggest teeth of all, are found. Pterossauros com uma grande crista sagital na~ parte anterior do cranio, situada nas pré-maxilas, que termina pouco antes do inicio da abertura naso-preorbital; e também uma pe- quena crista sagital parietal situada na parte posterior do cranio; denticdo presente desde a parte mais distal do cranio até o meio da fenestra naso-preorbital, regidio correspondente ao inicio da nariria interna; presenca de um alargamento na parte distal do cranio, onde se encontram os dentes pré-maxilares, que séo bem maiores do que os demais. Pterosaurier mit einer grossen Sagittalcrista, auf dem praemaxillaren Schaedelteil, die vor dem Nasopracorbitalfenster endet; auch eine kleine Sagittalcrista am hinteren Schaedelteil, auf den Parietalen; Bezahnung von der Schnauzenspitze bis ungefaehr der Mitte von der Nasopraeorbitaloeffnung, die dem Anfang der internen Naris entspricht; Verbreitung an dem distalen Bereich des Schaedels, wo sich die praemaxillaren Zaehne befinden, die viel groesser als alle anderen Zaehne sind. An. Acad, brasil. Ciéne., (1985) 57 (4)casas oacaatamaaasum 460 D.A.CAMPOS and A.W. A. KELLNER | Derivatio nominis: anhanga: ‘‘devil’’ + nera: “what was, old”. Anhanguera is a malignant genius in the Tupi culture, indigenous tribe of Brazil. A. blittersdorffi nov. sp. Holotype: Complete skull without mandible which belongs to the private Collection Desirée, from Mr. Rainer Alexander von Blittersdorff. Stratum typicum: This material was found in a typical calcareous nodule of the Santana for- mation, probably coming from the Romualdo Member, Chapada do Araripe, Brazil, Lower Cretaceous, Aptian — Albian. Diagnosis: ‘The same as for the genus. Derivatio nominis: in honour to Rainer Ale- xander von Blittersdorff, that has lend the skull for study. TABLE It Dimension of the Skull of Amhanguera blittersdorffi (én mm) Maximum length of the skull 499 Maximum height of the skull between the quadrate and the skull-roof 84 Maximum height of the skull at the region of the sagittal crest 6 Thickness between the quadrate, preserved part n Thickness between the quadrate, estimated a.80 Minimal thigkness estimated for the maxilla, (between the 9 and 10 alveoli) cal} Maximum thickness of the maxilla at the distal part of the skull (between the 3" and 4'” alveoli) : cad Length of the anterior sagittal crest a.192 Maximum length of the naris-preorbital fenestra (left side) 60 Maximum height of the naris-preorbital fenestra (left side) 48 Maximum height of the orbita 56 ‘An, Acad, brasil. Ciénc., (1985) $7 (4) DESCRIPTION As in the most pterosaur, the skull of Anhanguera is lightly built, with many open- ings, It is long, Slender and comparatively low. The bones are normally fused together, so that the interpretation of the pattern is become dif- ficult. Many of the sutures have not been easily interpreted. The greatest opening in the skull is the naris-preorbital fenestra; situated between the maxilla and the premaxilla. It has an enlongated form, becoming thinner in the distal part of the skull. In the posterior region, it’s border is formed by the jugal and the nasal. We could not observe if the lacrimal also takes part of this fenestra. The orbit has an ovaled form, and is form- ed by the parietal, frontal, lacrimal, postorbital and the jugal. After this, we can find the upper temporal opening, which seems to have a more rounded form then the orbita. Under the last is situated the lower temporal opening, which is more elongated. On the dorsal part of the skull we can observe the internal naris, which has an ovaled form. It is limitated by the ectopterygoid, parietal and laterally by the maxilla. After that, there is a subtemporal fenestra, formed by the ectopterygoid, pterygoid, quadrate and pro- bably by the jugal. The premaxillas are very long and fused together on the dorsal part of the skull. The suture which separates them from the maxillas can not be easily observed. It begins above the end of the naris-preorbital opening and get lost in the distal part of the crane. Where it can be observed, this suture is parallel to the alveolar margin of the maxilla. The separation with the frontal, lacrimal and nasal are also not well defined. The most interesting observation that could be made on the premaxilla is the presence of a large and very thin sagittal crest. It begins on the anterior part of the skull and ends almost at the beginning of the naris-preorbital fenestra. ‘That is the first time that a crest like that occurs in that position on a skull of flying reptiles,PANORAMA OF THE FLYING REPTILES STUDY IN SOUTH AMERICA 461 PLATEL ‘The skull of Anhanguera blittersdorffi, Fig. 1 — left side of the skull; Fig. 2 — right side of the skull. PLATE II “pom fs The skull of Anhanguera blittersdorffi, Fig. 1 — dorsal view of the skull; Fig. 2 — ventral view of the skull Abbreviations used in Plates I and II: al — alveoli; bo — basioccipitale; bs ~ basisphenoid; co — condylus occipitalis; ec — ectopterygoid; eo — exoceipital; f — frontale; j —"jugale; | —lacrimale; mx — maxillare; n — nasale; p — parietale; pl — palatinum; pmx — premaxillare; po — postorbitale; ps — parasphenoid; q — quadratum; aj — quadratojugale; scp — sagittal crest on the parietale; scpmx — sagittal crest on the premaxillare; so — supraoccipitale; sq — squamosal; t ~ tooth. An. Acad. brasil, Ciénc., (1985) $7 (8)D, A. CAMPOS and A. W. A. KELLNER ‘The skull of Anhanguera blittersdorffi, photographs. An, Acad, brasil. Ciéne., (1985) $7 (8)PANORAMA OF THE FLYING REPTILES STUDY IN SOUTH AMERICA which is one of the most characteristics of this form. important The maxillas are also very long and form practically the whole inferior border of the naris-preorbital opening. The limits with the jugal, quadrate and quadratejugal, as with the premaxilla could not satisfactorily be deter- minated. On the dorsal region the suture be- tween the maxillas and the palatal can be well observed, especially on the posterior half of the skull. The part which carries the alveoli is enlarged and stands out from the palatal region. The alveolis have a rounded or ovaled form. On the right side of the skull we could observe 26 alveolis, which let us conclude that Anhanguera has 52 superior teeth. The size of the alveolis become smaller in the posterior part of the skull. There were only 2 teeth préserved — the I" and the 10”. The nasal and the lacrimal could not be properly separated. Only a little part of the suture between these bones could be observed. The lacrimal has a little processus lacrimalis which penetrates into the orbita. The suture between the frontal and the parietal is also very weak. We could just observe that the frontal forms a great part of the superior border from the orbita and that the parietal delimitate part of the upper temporal opening. The limits of the parietal with the squamosal and with the supraoccipital can be better observed. There is also a little sagittal crest on the parietal, at the posterior part of the skull. The postorbital take part of the margins of the orbita, upper and lower temporal openings. It is been formed with three extremities, which are in contact with the parietal/frontal (?), jugal and squamosal. ‘The jugal separates the inferior parts of the orbita and narispreorbital fenestra. The exten- sion of this bone could not be observed, due to its fragmentary state on both sides of the skull. ‘The limits of the jugal with the quadratojugal and the quadrate could not be established; the sutures between this bones are absent. 463 The quadrate of the left side of the skull is better preserved then the one on the right side. It has an inclined position and forms part of the lower temporal fenestra’s margin. The separa- tion from the squamosal is made by a weakly suture. With the rest of the bones it could not be satisfactorily individualized. ‘The palatal and the occipital region of this skull is also preserved. We could identify some of the bones (Plate II, Fig. 2), but the sutures are so unclear that a more detailed study of these parts of the skull could not be made until now. SYSTEMATIC DISCUSSION According to the skull characteristics, Anhanguera can easily be classified in the sub- order Pterodactyloidea. However, when trying to classify this new form in any of the known families, we observed many differences, which are described below. — Pterodautridae Bonaparte, 1971: the only genus of this family, Prerodaustro Bonaparte, 1970, exhibits a singular skull and filiform den- tition, completely different from Anhanguera. —Ctenochasmatidae Nopesa, 1928: Creno- chasma Meyer, 1852 is a pterosaur that present a large quantity of teeth (200-360), that is one of the distinguishing features from Anhanguera. Also the dimensions of the skull-openings are very different from our specimen. Gnathosaurus Meyer, 1834, too differs from Anhanguera due io the presence of a low and elongated sagittal crest situated on the middle part of the crane on the first one. In Gnathosaurus we noticed also the absence of the little sagittal crest on the posterior part of the crane, that is present in Anhanguera, which is just one more difference between both forms. — Pteranodontidae Marsh, 1876: This family envolves the more specialized pterosaurs from the Superior Cretaceous. Pteranodon Marsh, 1876 and Nyctosaurus Marsh, 1876 are toothless forms. In Pteranodon we can see a ‘An. Acad, brasil. Ciéme., (1988) 57 (4)cmmnnat spenenentmropeen 464 D. A. CAMPOS and A. W. A. KELLNER huge parietal crest on the posterior part of the skull, which in Anhanguera is much smaller. —Pterodactylidae, Bonaparte, 1838. This family envolves small pterosaurs as Pterodac- tylus Curier, 1809 and Gallodactylus Fabre, 1974. Both have normally their teeth confined in the more distal part of the skull. Pterodac- tylus has a processus nasalis, which divide part of the naris-preorbitalopening; absent in Anhanguera. Galladactylus has great dif- ferences in the skull openings and a posterior extended part of crane, which is formed by the parietals. All the particularities distinguish it from Anhanguera. — Germanodactylidae Young, 1964. The sagit- tal crest of Germanodactylus Young, 1964 is placed in the middle of the skull. The teeth are small and this form exhibits a processus nasalis, like Pterodactylus. This form is also smaller then Anhanguera. — Criorhynchidae Hooley, 1914. The vertical teeth and massive rostrum of Criorhynchus Owen, 1874, are the most important differences to Anhanguera. — Ornithodesmidae Hooley, 1913. One of the distinguishing features of Ornithodesmus Seeley, 1887, is the presence of a small orbit, which is ventrally opened, and goes to an in- fraorbital fenestra. Also the presence of short teeth confined on the distal part of the skull dif- ferentiates it from Anhanguera. — Dsungaripteridae Young, 1964. The slightly arched rostrum, without teeth, of Dsungaripterus Young, 1964, which is the most important genus of this family — distinguished it from Anhanguera. Furthermore, Dsungarip- terus has a well developed median crest above the nasal and preorbital opening, which is com- pletely different from the sagittal crest on the premaxilla of the anterior part of the skull pre- sent in Anhanguera. An, Acad, brasil. Ciénc., (1985) $7 (4) — Ornithocheiridae Seeley, 1870. The classification of Amhanguera in this last known Pterodactyloidea family is also very difficult. Among the most important characteristics of Ornithocheirus Seeley, 1869 —~ the most impor- tant genus of this family — which differentiates this form from Anhanguera. We can point out the following: short and strong teeth, all with the same shape; parallelism between the distal alveoli-carrying parts of the maxilla, small orbit and naris-preorbital opening not much elongated; lacking of a lacrimal process that penetrates in the orbit; the shape of the skull itself, higher as in Anhanguera. Futhermore, we could observe in Anhan- guera an enlargement of the distal part of the skull, where we find the biggest teeth. But the most important characteristics that make us propose a new family for this specimen, is the presence of a sagittal crest above the premaxilla on the anterior part of the skull, which was never before observed in that position on any pterosaurian skull, ‘The comparison between Anhanguera and the known Brazilian pterosaurs is very restricted, since great part of them are defined on fragmented material from anterior members Nyctosaurus lamegoi, Araripesaurus castilhoi, Araripedactylus dehmi, Santanadactylus brasilensis). In relation to Brasileodactylus araripensis, we can notice a similarity of the mandibular symphysis and the anterior end of the skull of Anhanguera blittersdorffi. Both have bigger teeth in the most distal part, where they present an enlargement. Better comparision is at pre- sent not possible. Cearadactylus atrox can be distinguished from this new pterosaur by its natural notch in the mandibula and the maxilla, forming an empty space or gap. Also the form of the naris- preorbital opening in the skull of Cearadactylus is different from the one in Anhanguera. Due to all these differences and the im- possibility of a better comparison between the different forms of pterosaurs, the creation of a new genus and species is justifiable.PANORAMA OF THE FLYING REPTILES STUDY IN SOUTH AMERICA, FINAL CONSIDERATIONS As we have shown in this paper, there have been found several forms of flying reptiles in the Southern American Continent. In Brazil, there are presently known seven different forms of flying reptiles, from which six came from the sediments of the chapada do Araripe region. Many of them are based on material restricted to some parts of the body, and not on more complete individuals. Consequently there is the possibility that one or more forms of pterosaurs, which are until now known under different names, be synonyms. With the progress of the study on flying reptiles from Brazil, we will be able to prove the importance of the chapada do Araripe, especially the Santana Formation. An increas- ing number of remains of these animals is being discovered in that region, confirming that the Chapada do Araripe is one of the most impor- tant pterosaur localities in the world. The study of these remains will help us more and more to understand this facinating group of animals that once inhabited not only different parts of Brazil, but different regions of the world as well. After finishing this paper, we have received the notice of one more pterosaur locality in Argentina, situated at the Neuquen Province, Northern Patagonia. The only specimen found there consists of impressions and fragments of the sacrum, pelvic girdle and both femora. This came from the Lotena Formation (Upper Jurassic) and was first described by Casami- quela (1975) as a small Coelurosauria (Herb- stosaurus pigmaneus). Bonaparte (1978) disagreed with Casamiquela and associated this material to Pterosauria, probably Pterodactyloidea. SUMMARY It presents all the results of the various papers publish- ed regarding the South American pterosaurs. The remains 465 of these animals have been found in Chile (1 form), Argen- tina 2 forms) and Brazil (7 forms). Besides, there is made a preliminary description of a Pterosaurian skull, coming from the chapada do Araripe, Santana Formation, Ceard, N. E. — Brazil. Among the dif- ferent features, it presents a sagittal crest above the premax- illas on the anterior part of the skull, never found in that position before on any known pterosaurian skull. There is proposed a new form, Anhanguera blittersdorffi, and a new family is introduced: Anhangueridae. Until now, Brazil demonstrates a great variety of flying reptiles. From the seven forms found in this country, six came from the Chapada do Araripe, more specifically from the Santana Formation. This region can be considered as ‘one of the most important pterosaur localities in the world. REFERENCES BeuRten, K., (1963), Geologia ¢ estratigratia da chapada do Aratipe. An. 17° Congr. Nac. de Geol., Sudene: 1-47, BeuRLen, K., (1971), As condigdes ecolégicas e facioldgicas da formagio Santana na chapada do Araripe (Nordeste do Brasil). Supl. An. Acad. brasil. Ciénc., 43: 411-415. BONAPARTE, J. 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Bayer. Staatssig. Paldont. hist. Geol., 17: 157-167. WeLLnnoreR, P., (1978), Handbuch der Palioherpetologie — Teit 19 — Pterosauria, Gustav Fischer Verlag, 1-82. WeLinnorer, P., et al., (1983), A pterosaurian notarium from the Lower Cretaceous of Brazil. Paliiont. Zeit., 57, (1/2): 147-157. Wooowaro, A. S., (1932), In: Text-book of Paleontology, vol. 2, by Kari ZitreL, Second Revised English Edi- tion, London. MARIO SERC Inst A bacia médio do rio des de Engen oeste do Esta alongada, de com largura superficie de cidrios e qua de 240 km’, ( para agua sul sedimentos € mente essa e: que os sedime ficie de erosi A bacia embutida enti serra da Mai NW, ca serri altitude dos t cerca de 500: Sao Paulo, T de Resende cx frogénicas co de rifts da Se conjunto de tect6nica dest tal adjacente + Aceito r Trabalho re: Elétricas S.A.