A Pterosaurian Notarium from the Lower Cretaceous of Brazil

Pationt. Z. 147-157 Stuttgart, Juni 1983

A pterosaurian notarium from the Lower Cretaceous of Brazil

PETER WELLNHOFER, Miinchen, ERIC BUFFETAUT, Paris and PAUL GIGASE, Antwerpen'"

With 4 figures in the text K urzfassung: Aus derSantana-Formation (Apt) derChapada do Araripe in Nordost-Brasilien wird ein zusammenhangendes Stuck einer Pterosaurier-Wirbelsaule beschrieben und vorlaufig als Santanadaety Ius brasilensis DE BurSONJE bestimmt. Es besteht aus dem letzten Cervicalwirbel, den funf ersten, zu einem Notarium verwachsenen Dorsalwirbeln und den folgenden vier freien Dorsalwirbeln. Es handelt sich urn das besterhaltene Flugsaurier-Notarium, das bisher bekannt ist. Die funktionelle Bedeutung des Pterosauri er-Notariums wird diskutiert. A bs t r act: A pterosaur vertebral column consisting of the last cervical vertebra, the first five dorsals fused to a notarium, and the following four dorsal vertebrae, is figured and described from the Aptian San tana Formation of the Chapada do Araripe in northeastern Brazil. The specimen is tentatively referred to Santanadactylus brasilensis DE BurSONJE. It is the best preserved and most complete pterosaurian notarium known. Its functional significance is discussed.

Introduction

The Santana Formation (Aptian) of the Chapada do Araripe in northeastern Brazil has been famous for its fish fossils occurring in concretions since early in the last century (WELLNHOFER 1977). It was PRICE (1971) who first described pterosaur remains from the Santana Formation consi sting of parts of the wing skeleton (ulna, radius, carpals, metacarpals, first wing phalanx). He established the name Araripesaurus castilhoi and referred it to the family Ornithocheiridae. WELLNHOFER (1977) described the isolated first wing phalanx of a larger individual enclosed in a nodule of the same origin. A new name,Araripedactylus dehmi, was given to that specimen. It was included in an undetermined family of the suborder Pterodactyloidea. The wingspan of this individual must have been about five meters. In 1980 DE BurSONJE described two more pterosaur specimens from the Santana Formation ofthe Chapada do Araripe, a partial ankylosed scapulacoracoid in articulation with the proximal end of the humerus, and two cervical vertebrae in original contact. This material has been prepa red by acetic acid by DE BUISONJE and could be freed from the matrix. The bones turned out to be completely uncrushed, a condition hardly found with the thin walled pterosaur bones from other localities. DE BUISONJE considered the specimens to represent a new genus and a new spe cies, Santanadactylus brasilensis, belonging to the family Criorhynchidae.

". Address of the authors: Dr. P. WELLNHOFER, Bayerische Staatssammlung fur Palaontologie und histori sche Geologie, Richard-Wagner-StraGe 10,8000 Miinchen 2; Dr. E. BUFFETAUT, Universite Paris VI, Labo ratoire de Paleontologie des Vertebres et de Paleontologie Humaine, 4, place Jussieu, F-75230 Paris Cedex 05; Dr. P. GIGASE, Instituut voor Tropische Geneeskunde "Prins Leopold", Nationalestraat 155, B-2000 Antwerpen.

0031-0220/83/0057-0147 $ 2.75

This pterosaur material from the Santana Formation hitherto published consists of more or less isolated bones from different parts of the skeleton. Although it is questionable if these few specimens from the same horizon are systematically distinct, the distinction is justified until more complete material is available. Direct comparisons between these Brazilian specimens and the evaluation of their relationship is impossible. The same is true for the specimen dealt with in this paper. For the first time pterosaurian dor sal vertebrae and a notarium from the Santana Formation are being described. The specimen dif fers from all otherpterosaurs in which a notarium is known, and these are only a few. In fact, it is the best preserved and most complete notarium of a pterosaur known and contributes a great deal to our understanding of this special structure only found in Cretaceous pterodactyloids.

Pterosaur notarium material known The first record ofa pterosaurian notarium was given by OWEN (1859: 12, pI. 4, fig. 6-8) who described a symmetrical, compressed bone which had been found in the Upper Greensand of Cambridge (Cenomanian; Sedgwick Museum Cambridge Nr. B 54.408). OWEN referred to it as "Pterodactylus" (= Ornithocheirus) sedgwicki orfittoni, and characterized the specimen as fol lows: "The sides presenta smooth concave plate of bone, as ifthe piece had been nipped between a finger and thumb." OWEN took this fragmentary bone as a median longitudinal expansion of the frontal, the concave facets being interpreted as the inner walls of the orbits. Later the same specimen was redescribed by SEELEY (1870: 88, pI. 12, Fig. 15, 16). He located it as a bone from the underside of the skull such as the vomer and took the smooth oval depres sions of the compressed bone for the inner walls of the nares. It was MARSH (1881: 343; 1882: 254) who first reported that in the Cretaceous genus Ptera nodon of the Kansas Niobrara Chalk the pectoral arch was strengthened by the ankylosis of se veral vertebrae and by the robust scapulae articulating on opposite sides of the common neural spine of these vertebrae. SEELEY (1891: 438) referring to these observations by MARSH gave a new interpretation of the bone from the Cambridge Greensand, taken by OWEN (1859) for a frontal and by himself (1870) for a vomer. Now he recognized it as a portion of separated neural arches of three consecutive neural spines blended together. Based on some more similar bone fragments SEELEY concluded that this structure consisted of three dorsal vertebrae fused by a common supra-neural ossifica tion. The lateral oval facets then would correspond to the articular surfaces of the free ends of the scapulae. In his "Osteology ofPteranodon" EATON (1910: 17) mentioned "several more orless perfect examples of the anterior dorsal or notarial vertebrae ofPteranodon" in the MARSH Collection in New Haven (USA). He presented a restoration of the notarium based on the most complete though compressed specimen (YPM No. 2692). According to EATON the notarium of Ptera nodon consists of the eight anterior dorsal vertebrae firmly co-ossified. The dorsal spines are bound together and surmounted by a thin median ossification, the supraneural plate, where on each side it presents an oval facet for the articulation with the upper ends of the scapulae. These eightvertebrae are also bound together by paired ossifications that extend the length of the nota rium, fusing with the ends of the transverse processes. According to HOOLEY (1913: 382) the notarium of Ornithodesmus latidens SEELEY from the Wealden of the Isle of Wight consists of six ankylosed vertebrae. The neural spines were fused into one bearing lateral facets between the third and the fourth dorsal for the articulation with the scapulae. It differs from the notarium of Pteranodon in the absence of the fusing of the ex tremities of the transverse processes by a band-like ossification. Pterosaurian notarium 149

Recent investigation of this specimen (British Museum London Nr. RJ3877) revealed a different position of the lateral facets. The structure identified by HOOLEY (1913: pI. 38, fig. 3) as the "supposed articular facet for the scapula" is in fact an artificial depression due to compres sion. The actual facet is clearly developed laterally on the neural plate between the fifth and the sixth notarial vertebra and, as found with the notarium described here, on the widest expansion of the neural plate. YOUNG (1964: 243) has shown that in Dsungaripterus weii from the Lower Cretaceous of Sinkiang (China) a notarium was also developed. Itis composed of"atleast eightvertebrae", but only the last three are preserved, followed by three free dorsals. Therefore we know only the posterior part of the notarium in Dsungaripterus. The dorsal spines are fused at their tops and the centra are more or less co-ossified. The lateral facets for the scapulae must have been situated at the anterior portion of the fused neural spines which has been broken away.

Geological occurrence The pterosaurian remain described here comes from the Santana Formation of the Chapada do Araripe, a vast plateau of northeastern Brazil, stretching for 180 km from East to West and for about 40 km from North to South, on the border between the states of Ceara and Pernambu co, and reaching the state of Piaui in the West. More accurate data about the exact geographical origin of the fossil are not available (fossiliferous localities are known all around the Chapada do Araripe: see CAMPOS & WENZ 1982). A synthetic section of the sedimentary series of the Chapada do Araripe has recently been published by CAMPOS & WENZ (1982). The Santana Formation itself is divided into three mem bers (BEURLEN 1971): Crato (lower), Ipubi (middle) and Romualdo (upper). The famous fos sil-bearing nodules from the Chapada do Araripe come from a marly layer at the base of the Ro mualdo member; above this basal layer come limestone beds with, at the top, marine fossils. The nodules from the Santana Formation are well known for the large numbers of well pre served fishes they have yielded (SILVA SANTOS & VALENCA 1968), including rare selachians, coe lacanths (CAMPOS & WENZ 1982), and abundant actinopterygians. Other vertebrates also occur in the nodules, although they are much less frequent than the fishes: chelonians, crocodilians and pterosaurs are represented. The invertebrate fauna of the Santana Formation as a whole, as listed in detail by LIMA (1979), is varied, and includes foraminifera, bivalves, conchostracans, ostracods, copepods, malacostracans, insects and echinoids. The microflora and macroscopic plant remains, also listed by LIMA (1979), are abundant. Various geological ages have been suggested for the Santana Formation. A late Cretaceous age has long been admitted. In 1962, for instance, BEURLEN referred the Santana Formation to the Turonian. More recent studies, however, indicate an early Cretaceous age: BEURLEN (1966), on the basis of echinoids and SILVA SANTOS & VALENCA (1968), on the basis of fishes, favoured an Aptian age, a conclusion also accepted by MABESOONE & TINOCO (1973). Other recent pa laeontological studies would suggest a slightly later age: LIMA (1979), on the basis of the micro flora, concludes that the Santana Formation is Albian. CAMPOS & WENZ (1982) refer the Crato and Ipubi members to the Aptian, and consider that the marine beds at the top of the Santana Formation (in the Romualdo member) correspond to the first Albian marine transgressions. The fossil-bearing nodules at the base of the Romualdo member would then come from near the Ap tian/Albian boundary. Transatlantic correlations based on fossil vertebrates can also be of some use in dating the no dule layer of the Santana Formation. The notosuchian crocodilianAraripesuchus gomesii PRICE, 1959, from the Santana Formation, is very closely related to the species Araripesuchus wegeneri BUFFETAUT, 1981, from the Elrhaz Formation of Gadoufaoua, in Niger (BUFFETAUT & TAQUET 150 P. Wellnhofer, E. Buffetaut and P. Gigase

1979; BUFFETAuT 1981). This close relationship is evidence that at this time South America and Africa had not yet been completely separated by a sea barrier, and it suggests that the Santana Formation and the Elrhaz Formation are of approximately the same age. The probable occur rence of the turtle Araripemys, first described from the Santana Formation, at Gadoufaoua (DE BROIN 1980) also supports this view. On the basis of stratigraphic and palaeontological evi dence, the Elrhaz Formation is referred by TAQuET (1976) to the Aptian. A late Aptian age is thus suggested for the vertebrate-bearing nodules from the Santana For mation, and thus for the pterosaurian remains described here. The depositional environment of the Santana Formation has been discussed by many au thors. It appears that ecological conditions changed during the deposition of the formation, from lacustrine to lagoonal with marine influence to lacustrine again (MABEsooNE & TINOCO 1973; LIMA 1979). According to CAMPOS & WENZ (1982), a marine episode occurs at the top of the Santana Formation. It is generally admitted that the vertebrate-bearing nodule layer at the base of the Romualdo member was formed in a freshwater environment, although some authors admit connections with the sea (SILVA SANTOS & VALENCA 1968, suggest estuarine conditions).

Preparation of the specimen The specimen was obtained in Belgium in December 1980 from the owner of a large collec tion offossil fishes from the Santana Formation. It consisted of an unsplit spindle-shaped nodule about 210 mm long, made up of homogeneous light brown matrix. One face showed the section of a series of procoelous vertebrae while parts of the neural and transverse apophyses were also apparent, corresponding to the worn parts of Fig. 2. The structure of the bone was extremely light and brittle. Mechanical preparation was excluded. The apparent bony structures were im pregnated with butyl metacrylate dissolved in acetone and the specimen was immersed in 10 % acetic acid saturated with calcium hydroxytriphosphate. The matrix appeared to dissolve very slowly, exposing more and more bone. The specimen was repeatedly washed, dried and treated again with plastic. The whole process took about five months. The impregnation was in fact not imperative due to the silicification of the bone, but it helped in the preservation of some delicate structures. The specimen was eventually immersed in acetone to clear it from excess metacryla teo The dissolved matrix left only a limited residue of argillaceous nature. The presence of numerous ostracods was noted in the matrix during the preparation.

Description of the specimen

Ordo Pterosauria KAUP, 1834 Subordo Pterodactyloidea PLIENINGER, 1901 Familia indet. Santanadaetylus brasilensis DE BUISONJE, 1980 Figs. 1-4

The specimen is housed in the collection ofDr. PAUL GIGASE, Antwerpen, under the number V-201. It consists of a series of ten vertebrae in natural contact. They are uncrushed and not compressed. The ventral parts of most of the centra are eroded away, exposing the spongy internal struc ture of the bone. The vertebrae are procoelous, the articular surfaces forming a transverse kid ney-shaped ball and socket articulation. Only the transverse processes of the last five vertebrae are more or less complete, those of the first five vertebrae are broken, and their distal extremities are lacking. There is a relatively wide neural canal, triangular in outline in the first vertebra and more oval and higher than broad in the last one. Pterosaurian notarium 151

Fig. 1. Santanadactylus brasilensis DE BursONJE, Santana Formation (Aptian), Chapada do Araripe, Brazil. Vertebral column from the last cervical up to the tenth dorsal, including the notarium (i. e. the first five dor sals). A. in lateral view from the left, B. in top view, C. in ventral view, showing the eroded surface of the centra, and D. in lateral view from the right. P. GIGASE collection, nr. V-201. (Photo by F. HOCK, Munich) 152 P. Wellnhofer, E. Buffetaut and P. Gigase

The top of the neural spine of the first vertebra is broken away. At its base above the post zygapophyses it is 10 millimeters wide. The width of the neural spine of the following vertebra measures at the same level only 6.5 millimeters. Therefore the neural spine of the first vertebra appears to have been rather stout. Neitherthe prezygapophyses nor the transverse processes of this vertebra are preserved. The postzygapophyses are overhanging the centrum which is true for all vertebrae ofthe series. They are still in close contact with the prezygapophyses of the following vertebra. The plane of the zygapophyseal articulation is rather steep, almost vertical, converging slightly to the centrum and in top view to the rear. There seems to be no firm co-ossification between the postzygapo physes ofthe first vertebra and the prezygapophyses of the second. But in some places where the internal bone structure of the centra becomes visible due to erosion a partial fusion of the first tWQ vertebrae can be observed. Since the first vertebra is so much different from the following vertebrae and since it closely resembles the last cervical ofPteranodon described by EATON (1910) it seems likely that this ver tebra represents the last cervical rather than the first dorsal still in close contact with the dorsal series and showing a tendency of fusion with the first dorsal. This conclusion is in agreement with the fact that the articular facets for the scapulae on the fused neural spines of the notarium are located between the third and the fourth dorsal in both Pteranodon and Ornithodesmus. In the Santana specimen these facets are above the fourth dorsal if we take the first vertebra as the last cervical. The notarium proper, to follow, is composed of the first five dorsals. They are co-ossified in different ways. As is shown on the eroded ventral part ofthe first three vertebrae, the centra have confluent spongy bone structures, but only the first and the second dorsals have fused neural spines. There is a clear separation between the neural spines of the first two dorsals and the fol lowing three. On the other hand the second and the third vertebrae are co-ossified by their cen tra. The dorsals three, four and five form a unit where the neural spines are fused with their upper extremities to form a bony plate. In contrast to the neural spines the centra and the zygapo physes of these vertebrae are not co-ossified and their sutures can be clearly distinguished. The posterior block of fused spines is more solid and stout than the anterior one composed of the spines of the first two dorsals. It expands rapidly above the fourth dorsal and becomes re markably bulbous. The facets for the scapular articulations are in the middle of the neural plate above and slightly posterior to the fourth dorsal. They are oval in outline, 22 millimeters long and 18 millimeters high. The articular surfaces are slightly depressed and saddle-shaped. Although in firm contact, the sixth dorsal is not co-ossified with the fifth and the posterior series of the dorsals six to nine is not fused by co-ossification either. Central and zygapophyseal contacts are clearly marked by sutures, bone structures are not confluent. So, these four verte brae represent free dorsals and cannot be included with the notarium. Still in contact with the ninth dorsal the anterior portion of the neural arch with the prezygapophyses of the tenth dorsal is preserved. The transverse processes of the last cervical are missing. On the first notarial vertebra these processes are lacking their distal extremities. They are rather stout, originating on the centrum from a broad base and from the anterior half of the neural arch. The cross section at the broken end is square. On the right side, ventraly to the base of the transverse process and situated ante rolaterally on the centrum a downward facing capitular rib facet is present. This part of the cen trum is not preserved in the last cervical nor in the second or third dorsal. A capitular facet is de finitely absent from the centrum of the fourth dorsal. There, both the capitular and tubercular facets are located at the extremity of the rather broad transverse processes. The capitular facet is situated on a short expansion of the transverse process directed anteroventrally. Evidently the fifth dorsal still supported double-headed ribs, a feature which cannot be demonstrated for the Pterosaurian notarium 153

cervical ~ 'I------notarium------,r-----tree dorsals------, sct i

\ \ \ ----J------'------"------_L i 1. 2. 3 4. 5. 6. 7. 8. 9. 10.

o 3 5cm L'_--'-_--'-_--',_---..J_---',

Fig. 2. Santanadactylus brasilensis DE BmsoNJE, Santana Formation (Aptian), Chapada do Arari pe, Brazil. Vertebral column including the nota rium, specimen of Fig. 1. A. Complete specimen as preserved in lateral view. B. Top view of the specimen. C. Last cervical vertebra in anterior view. D. Posterior view of the specimen showing the ninth and fragments of the tenth dorsal verte bra. Abbreviations: cf capitular rib facet, nc neu ral canal, nsp neural spine, poz postzygapophy sis, scf articular facet for the scapula, tf tubercular rib facet, trp transverse process.

following vertebrae. The transverse processes of the third dorsal differ from all others in that they are directed posteriorly. The free dorsals six to nine are still in close contact with each other but not fused. They are more slender than the notarial vertebrae with slender transverse processes projecting out hori zontally at right angles. Their neural spines are angular in outline with flattened tops. They are only three to four millimeters thick, much less than the fused neural spines of the notarium. All vertebrae are lightly built. There are numerous foramina especially in the ventral surfaces of the transverse processes of the notarium and in the surfaces transitional between the trans verse processes and the neural spines. The internal structure of all vertebrae consists entirely of a trabecular framework of bone tissue. 154 P. Wellnhofer, E. Buffetaut and P. Gigase

Measurements (in millimeters): Length of vertebral column as preserved 205 Length of centra Last cervical 21 Dorsal one to three (fused) 66 Dorsal four 22 D~alfift II Total length of the notarium 110 Dorsal six 17 Dorsal seven 17 Dorsal eight 17 Dorsal nine 17

Discussion and comparison The pterosaur notarium described here cannot be referred to any of the notaria known. The notarium of Ornithocheirus from the Cambridge Greensand (SEELEY 1891) is much too frag mentary to be compared with the Brazilian specimen. The notarium ofPteranodon is clearly di stinct. It consists of eight co-ossified dorsal vertebrae and the fused dorsal spines are surmoun ted by a thin median ossification, the supraneural plate. These eight vertebrae are also bound to gether by the fusion of the extremities of the transverse processes. Moreover the transverse and capitular processes of the first three notarial vertebrae are completely fused with the ribs which they support (EATON 1910). In the notarium of Ornithodesmus from the Wealden of the Isle of Wight the transverse pro cesses are not fused as in the Santana specimen. But it consists ofsix co-ossified vertebrae and the neural spines are fused into one strong ridge (HOOLEY 1913). The notarium of the Lower Cretaceous Dsungaripterus from China described by YOUNG (1964) is too fragmentary to be compared with the specimen described here. We know neither the number ofvertebrae it consists of nor the articular facets for the scapulae on the fused spines. In general, there is some resemblance with the notarium of the English genus Ornithodes mus, however, although the degree of fusion seems to have proceeded further in that form. There is no evidence for the systematic position of the notarium described here. As pointed out already, inclusion in one of the three Santana pterosaur generaAraripesaurus, Araripedaety Ius andSantanadactylus is not possible. On the other hand the establishing ofa new name should be avoided. The only reason for the determination of this specimen as Santanadaety Ius brasilen sis is the size. The two cervical vertebrae from the Santana Formation described by DE BUISONJE (1981) are about the same size as the last cervical of our specimen. More complete material should clarify this relationship. The tentative reconstruction of the notarium ofSantanadactylus brasilensis given by DE BUISONJE (1981: 43, fig. 9) should be modified according to the evidence presented here.

Functional significance The development of a notarium in pterosaurs is only known within the short-tailed suborder Pterodactyloidea and here only in the Cretaceous genera Ornithocheirus, Ornithodesmus, Dsungaripterus, Pteranodon, Nyctosaurus andSantanadactylus. All these pterosaurs are large or very large flying vertebrates which are thought to have performed gliding rather than active flight (WELLNHOFER 1978). The extreme condition is realized inPteranodon, where not only the first eight dorsal verte brae are fused, forming a rigid mass of bone, but where also the anterior ribs are immovably coa- Pterosaurian notarium 155

scI

prz ------paz

/ / 2. et' 1. o 2 3 4 5 em L.'_----'-__,'---_-"-,_---J'c------',

Fig. 3. Santanadactylus brasilensis DE BUISONJE. Reconstruction of the notarium in lateral view from the left. Abbreviations as in Fig. 2. lesced with the notarium and fixed to the sternum, thus building a rigid structure consisting of the notarium, the rib cage and the sternum. The shoulder girdle of most pterosaurs is composed of the scapula and the coracoid ankylo sed into one hook-like bone which articulates medioventrally with the sternum. The scapulae usually overlie the ribs, converging with their free ends towards the dorsal vertebral column. Only where a notarium is developed, are the scapulae anchored to the fused neural spines of the anterior dorsals by articulation. 50 the scapulacoracoid can swing back and forth with the cora coid articulation on the sternum and the scapula articulation on the notarium functioning as hin ges. From this results a forward and backward shifting of the glenoid articulation for the hume rus. The consolidation of dorsal vertebrae is met within many birds, for example Pelecanus, Fal cinellus, Phoenicopterus, Falco, Microhierax, Polyborus, Podiceps and most ofthe Galli. They all have an extended os dorsale or notarium (5TRESEMANN 1932: 517). This is generally regarded as directly correlated with increased power offligh t in order to provide the trunk with the greatest possible rigidity for the attachment of flight musculature. The coalescence of dorsal vertebrae in the avian skeleton sometimes can involve the entire se ries of dorsal vertebrae. In pterodactyls with a notarium at least three or four dorsals anterior to the pelvis are freely movable. As is shown in Santanadactylus (Fig. 4) the articular surface of the

Fig. 4. Santanadactylus brasilensis DE BUISONJiL Reconstruction of the ninth dorsal vertebra in lateral view A B (A) and in posterior view (B). c cen trum. Other abbreviations as in o 2 3 4 5cm

Fig. 2. <-'-----'-----',---'-,----',----', 156 P. WeIlnhofer, E. Buffetaut and P. Gigase

centra are broader than high and the zygapophyseal facets are rather steep. Thus the flexibility of these free dorsal vertebrae was possible only in a more sagittal plane rather than in a lateral movement. In the avian notarium a supraneural plate bearing articular facets for the scapulae is never de veloped. This feature is a peculiarity of the pterosaur condition. It is related to the much longer wings in pterosaurs which have to be anchored effectively to the vertebral column. The nota rium of the pterodactyls is a rigid structure to which during flight the centre of gravity was su spended. The fused neural bony plate not only gave support to the scapulae but also served as an area for the origin of strong brachialis muscles, especially of the latissimus dorsi group which was responsible for pulling the humerus upwards and backwards (FURBRINGER 1902). Powerful latissimus dorsi muscles were necessary for holding the long wings up and thus for sustained gliding.

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Eingang des Manuskripts bei der Schriftleitung am 15. 12. 1982.