Pterosauria, Pterodactyloidea, Anhangueridae);

Home , Anhangueridae, Brasileodactylus, Crato Formation

CRANIUM 22,1 (2005)

Brasileodactylus (Pterosauria, Pterodactyloidea, Anhangueridae);

an update

André+J. Veldmeijer Marco Signore and Hanneke Meijer

Summary

Brazil and The majority of the teethed pterosaurs from the Araripe Basin in (Santana Crato Formations) have premaxillary sagittal crests and dentary sagittal crests. A few skulls, however, lack such crests and ever since these

sexual fossils have been published, discussion continueswhether these are independent taxa, ontogenetic variants or

dimorphics of other taxa. The present work presents an update, discussing the dentitionand other morphological features, previously unnoticed, in order to evaluate the systematic position of the taxon.

Samenvatting

Brazilië Crato Het merendeel van de getande pterosauriërs die afkomstig zijn uit het Araripe Basin in (Santana en

kam kam. Er echter die Formaties) hebben een premaxilla sagittale en een dentary sagittale zijn een paar taxa geen kammen hebben deze de schedel vanafhetallereerstemoment dat deze fossielen bekend is op plekken op en waren, of deze of dat deze dieren of sexueel dimorfische varianten er gespeculeerd een geldig taxon zijn ontogenetische zijn

kort van andere taxa. Deze bijdrage presenteert een update waarin het gebit gepresenteerd en besproken wordt, alsmede enkele morfologische kenmerken dieeerder over het hoofdwerden gezien, met als doel de systematische

evalueren. positie van het taxonBrasileodactylus te

Introduction measurements are most of the time not included

(exceptions are Fastnacht (2003), Lee (1994) and first this Ever since the pterosaur was discovered, Veldmeijer (2003b)). Therefore, measurement of of vertebrates has been the extraordinary group teethand alveoli are includedin the present work subject of much debate and discussion. This is and discussed. A few problems, however, occur skeleton partly dueto the fact that the pterosaur is which complicate the taxonomic issue. and there extremely fragile as a consequence, are only relatively few fossils and even fewer well preserved complete specimens which have been Materials and methods adequately prepared. Nowadays, there are a few

sites for of which the major pterosaurs, Araripe The used for following specimens are compa- basin in Brazil is the most important one. rison:

The aim of the work is to present present an of - Skull Anhanguera blittersdorffi, MN-4805-V of the toothedpterodactyloid taxon Brasi- update - Skull and mandible of referred specimen of leodactylus on the basis of the skull, since long the Anhanguera blittersdorffi, n. 40 Pz-DBAV-UERJ most distinctive element in the pterosaur - Skull Anhanguera santanae, AMNH 22555 skeleton. Consequently, only comparable taxa - Mandible 22573 Anhanguera sp, AMNH from Brazil will be discussed of which cranial

- Mandible SAO 200602 Anhanguera sp, Arthurdactylusmaterial is known and therefore - Anhanguerid skull in the Iwaki Museum, Araripedactylus& Martill, 1994, conandoylei Frey IMNH 1053 castilhoi Price, 1971, Araripedactylus dehmi Brasileodactylus- Skull and mandible cf. arari- Wellnhofer, 1977, Santanadactylus brasilensis De pensis, MN-4797-V 1980, Santanadactylus pricei Wellnhofer, Buisonje, - BrasileodactylusAnterior part mandible araripen- 1985 and 1985) Santanadactylus spixi (Wellnhofer, sis, MN-4804-V are left out. - Anterior mandible Brasileodactyluspart MN-6517-V Dentition is sometimes used for systematic ? araripensis,

- Skull and mandible Unwin used the Brasileodactylus sp., AMNH purposes (especially (2001) 24444 relative size, but the material he used is extremely

- Partial skull BSP19911 27 fragmentated and no reliable picture of the entire Brasileodactylus sp,

- Anterior skull dentition can be obtained), although the part Coloborhynchus clavirostris, BMNH 1822

45 Brasileodactylus (Pterosauria, Pterodactyloidea, Anhangueridae);

an update

= Staatliches Museum fur - MandibleColoborhynchus robustus, BSP1987147 Karlsruhe; SMNS

- Skull and mandible Coloborhynchus piscator, Naturkunde, Stuttgart.

NSM-PV19892 All measurementsof alveoli and diastemae have skull and mandible - Anterior parts Coloborhyn- been taken laterally except for the anterior and chus robustus, SMNK 2302 PAL anterolateral alveoli. These are measured

- Skull Coloborhynchus sp, MN-4735-V lateral-lateral and anterolateral-ventrolateral SAO 16494 - Skull Coloborhynchus sp, respectively. The patterns are visualised in a

- Skull and mandible ? Coloborhynchus sp., graph (graph 1 & 2). MN-6687-V

? MN-6503-V - Skull Coloborhynchus sp,

- Mandible ? Coloborhynchus sp., MN-6687-V and The known - Skull mandible Coloborhynchus spielbergi, specimens

RGM 401880 Type specimen ofBrasileodactylus andholotype of

- Skull and mandible Criorhynchus mesembrinus, Brasileodactylus araripensis (Fig. 1). BSP1987146 Order Pterosauria Kaup, 1834 - Mandible cf. Criorhynchus mesembrinus, SMNS Suborder 1901 56994 Pterodactyloidea Plieninger, & Kellner, 1985 Undescribed skeleton of crestless Family Anhangueridae Campos - a pterosaur in GenusBrasileodactylus Kellner, 1984 the Kitakyushu museum

and arari- Institutional abbreviations: Type species specimen: Brasileodactylus pensis, anterior part of mandible, MN 4804-V,

BMNH = British Museum of Natural History, Museu Nacional, Rio de Janeiro, Brazil. London; BSP = Bayerische Staatsammlung fur Brasileodactylus as diagnosed by Palaontologie und historische Geologie, Munich; Diagnosis: Kellner (1984:580): "Pterosaurier mitUnterkiefer IMNH = Iwaki Museumof Coal Mining & Fossils,

aus einer am Ende = gebildet langlichen abgerun- Iwaki; MN Museu Nacional, Rio de Janeiro; detenSymphyse, leicht nachoben gebogen, trian- NSM = National Science Museum, Tokyo; gularem Querschnitt, Schmalerung ab dem Pz-DBAV-UERJ = Geological Museum Univer- Teil, wobei ene an = proximalen Verbreiterung sity of Rio de Janeiro; RGM Nationaal Natuur- dem distalen Bereich ab der dritten Alveole exis- historisch Museum (Naturalis), former tiert, die eine flache Oberflache bildet. Vorhan- Rijksmuseum voor Geologie en Mineralogie, denseineiner medialenFurche an derDorsalseite Leiden; SAO = Sammlung Oberli, St. Gallen; der sehr ab dem SMNK = Staatliches Museum fur Naturkunde, Symphyse, ausgepragt Beginn

Fig 1 Type specimen ofBrasileodactylus and holotype of Brasileodactylus araripensis, MN 4804-V. Courtesy of Museu

Nacional, Rio de Janeiro, Brasil. Photograph by E. Endenburg/A.J. Veldmeijer

MN 4804-V. Met dank Natio- Type specimen van Brasileodactylus en holotype vanBrasileodactylus araripensis, aan naal Museum, Rio de Janeiro, Brazilië. Foto door E. Endenburg / A.J. Veldmeijer

46 CRANIUM 22,1 (2005)

Fig 2 The Crato specimen, MN 4797-V. Courtesy of Museu Nacional, Rio de Janeiro, Brasil. Photograph by E. En- denburg / A.J. Veldmeijer

Het Crato stuk, MN 4797-V. Met dank aan het NationaalMuseum, Rio de Janeiro, Brazilië. Foto door E. Enden- burg / A.J. Veldmeijer

in des Unterkiefers (distaler Teil), die sich proxi- ting on the rostral tip and widening caudally." maler Richtung verbreitet. Alveolen mit elipti- have to be regarded as apomorphies of Brasi- scher und rundlicher Form, Zahnabstande leodactylus.” They regard the degree of expansion ibidem Kellner vergrossern sich in proximaler Richtung. as apomorph ( : 103). (1984) regards

Bezahnung bis an die Unterkieferspitze, Zahne Brasileodactylus as Ornithocheirid. Therostral end schmal und spitz, nach vorne stehend." (For starts to expand between the third and fourth

English translation, see Kellner & Tomida (2000: alveoli, which is between the fourth and fifth 102). alveoli in Anhanguera and Coloborhynchus.

However, the expansion in SMNS 55414 starts Emended diagnosis: Combination of first pair of betweenthe fourth and fifth alveolus as well. The the the alveoli positioned at anterior aspect; expansion inBrasileodactylus is small but distinct, second pair of alveoli positioned anterolateral similar to the in very situation Anhanguera, and is and the third pair of alveoli lateral. The dentary not consistent with the robust expansion in Colob- has small sagittal groove exten- orhynchus. Thefirst threepairs of alveoli are at the ding sub-grooves. anterior, anterolateral and lateral aspect respecti-

Discussion of diagnosis: Kellner & Tomida (2000: vely (these are positioned anterodorsally and

103) evaluated Brasileodactylus and came to the laterodorsally in Anhanguera and Coloborhynchus; conclusion that "4) rostral end expand from the both second and third pairs are orientated antero-

3rd alveoli, forming a flat surface. 5) medial dorsally. The remaining alveoli are placed the dorsal of the relative for groove on part symphysis, star- strongly laterodorsally, to, instance,

47 Brasileodactylus (Pterosauria, Pterodactyloidea, Anhangueridae);

an update

Fig 3 The New York specimen, AMNH 24444. Courtesy of AMNH, New York. Photograph by E. Endenburg/A.J Veldmeijer

Het stuk uit New York, AMNH 24444. Met dankaan AMNH, New York. Foto door E. Endenburg/A.J Veldmeijer

comparable alveoli in Coloborhynchus as prelimi- The third alveolus is the largest of the alveoli

in in nary results of study progress suggests. found the distal expansion; the following alveoli decrease in size continuously but the size In general, thealveoli are small. The first tooth is increases again steadily from the sixth alveolus the smallest, after which the tooth size increases.

48 CRANIUM 22,1 (2005)

onwards. This results in the ninth alveolus being The Crato specimen (MN 4797-V; Fig. 2)

as large as the third one (remarkable is Kellner's Various fossils have been assigned to Brasileodac- statement on the teeth, as they are not preserved tylus. The fossil in Figure 2, originating from the in the specimen). Crato Formation of the Araripe plateau (in the

The small sub-grooves of the dentary sagittal collection of the Museu Nacional, Rio de Janeiro;

in MN has been to the grooveare only seen Brasileodactylus (holotype 4797-V), assigned genus by

& Kellner as and SMNS 55414) and are regarded as Sayao (2000) Brasileodactylus sp. apomorphy. Finally, the symphysis is extremely mainly on the basis of the absence of crests and

the of a mandibular However, a elongated relative to other Brazilian pterosaurs presence groove. of the like Anhanguera (for instance, Veldmeijer et al, in mandibular groove starting at the tip

Veld- can be found in other as well press a), Criorhynchus (Wellnhofer, 1987; dentary pterosaurs meijer, 2002; Veldmeijer & Hense, 2004) and (for instance, Coloborhynchus robustus), but the Coloborhynchus (for instance, Kellner & Campos, type specimen of Brasileodactylus as well as the 2000; Veldmeijer, 2003b). mandiblein theStuttgarter collection (see below) clearly shows small anterolateral running sub- Although we think that the absence of features the fact the fossil grooves. Due to that is severely should not be used in a diagnoses, the absence of deformed by compression, it cannot be deter- crests seems to be important in the studied taxa. mined whether these peculiar sub-grooves are Most pterosaurs from this area have crests, present or not.Furthermore, it is difficult to deter- premaxillary as well as dentary. Though recently mine the exact position of the alveoli, due to the a new pterosaur has been published (Frey et al., deformation, although at least the second and 2003) without premaxillary and dentary sagittal third right alveolus seem to be in a laterodorsal crests but with a parieto-occipital crest, discussed position rather than anterolateral and lateral in more detail below. So far, Cearadactylus and respectively. According to the authors, the most Brasileodactylus are the only genera without the anterior portion is expanded, containing the premaxillary and dentary crests.

Fig 4 The Stuttgarter mandible, SMNS 55414. Courtesy of SMN, Stuttgart. Photograph by R. Harling

De onderkaak uit Stuttgart, SMNS 55414. Met dank aan SMN, Stuttgart. Foto door R. Harling

49 Brasileodactylus (Pterosauria, Pterodactyloidea, Anhangueridae);

an update

the lack of and largest teeth, although it is not stated which ones most important ones are maxillary the teeth. in exactly. However, the graph shows that the dentary sagittal crests and However, table second to fourth alveoli are the biggest together due course the published (Veldmeijer, details with the seventh alveolus (see below). 2003a: 11) should be revised because more of the can be added after complete preparation the The measurements are comparable to type fossil. Thecomplete preparation of this specimen, specimen of Brasileodactylus and holotype of which at this moment is being undertaken in The Brasileodactylus araripensis (MN 4804-V), Netherlands, is important, because relevant although slight differences can be noticed (for characteristic features, such as the exact position instance, the first and sixth alveolus). A major of the first three alveoli, a possible expansion of of the difference is seen in the measurements the jaws and the sub-grooves, are obscured by which eighth, tenth and eleventh alveolus, are and has matrix. Recently, a new genus species decreasing in size relative to the previous alveoli. been published, which originates from the Crato This contrasts with the holotype (MN 4804-V), member (Fig. 4), Ludodactylus sibbicki Frey et al. which shows an increase in alveolar diameter, 2003 (in the collection of the Staatliches Museum relative to the preceding ones. fiir Naturkunde Karlsruhe, Germany) that looks

dentition of the skull shows similar to The a comparable remarkably Brasileodactylus sp. (teeth,

for the but with crestless and but seems pattern as seen mandible, two dentary premaxilla) to

differences. The third alveolus differ the of a distinct is clearly in presence parieto-occipital crest, the biggest; the differences in measurement of seemingly lacking in the AMNH24444specimen. of the this and the other alveoli are distinctly bigger However, the dorsal view cranium (Veld-

the mandible. Furthermore, the 2003a: 3 and shows a than seen in meijer, figures 4) possible eleventh alveolus is the smallest in the mandible, start of a parieto-occipital crest (referred to as fifth The but in the skull, the smallest alveoli are the frontoparietal crest in Veldmeijer, 2003a). and sixth; the size of the eleventhalveolus equals crest might have been broken off, so the exact

that of the tenth and of size the and its extend be determined. are comparable to shape cannot fifth and sixth. Detailed research of Ludodactylus sibbicki,

will shed currently in progress, possibly light on The New York specimen (AMNH 24444; Fig. 3) the attachmentof the crest to theskull. Only after

The fossil in the collection of the American the full preparation of AMNH 24444 a detailed

Museum of Natural History (AMNH 24444), comparison can be done and conclusions be preliminary described by Veldmeijer (2003a) as drawn. The first alveolus of the mandible is

has been to this the other Brasileodactylus sp. assigned comparable to the size of this alveolus in which the genus for several characters, among discussed specimens, but the second, third and

Fig 5 The Munichspecimen, BSP 1991 I 27. Courtesy of BSP, Munich. Photograph by A. ’t Hooft

dank Foto A. ’t Hooft Het stuk uit München, BSP 1991 I 27. Met aan BSP, München. door

50 CRANIUM 22,1 (2005)

the fifth The Munich fourth alveoli are substantially smaller; specimen (BSP 1991 I 27; Fig. 5) alveolus is bigger and equals the size of the first. The material housed in the Bayerische Staats- The position of the alveoli could not be estab- sammlung für Palaontologie und historische lished due to the unprepared state; the mandible Geologie, Munich, Germany is tentatively classi- faces lateroventrally, limiting necessary visibility fied as Brasileodactylus by Veldmeijer et al. (in to describe alveolar position as well as reliable review). The material consist largely of post-cra- measurements. nial material and includes only a small portion of

Theright side of the skull is exposed, allowing the the crestless maxilla; a direct comparison between this and the is measurement of 26 alveoli. The alveoli are small specimen type specimen because the Munich materials do with bigger measurements for the second, third, notpossible, not include the mandible. Nevertheless, in fourth, ninth and sixteenth alveoli. These are assigning the thebasis of the substantially bigger than the corresponding specimen to Brasileodactylus on dentition is distinct alveoli in the mandible; most others are smaller, comparable (which clearly

from the dentition in and the crest- except for the eleventh alveolus. Although the Cearadactylus ) less maxilla in this well ninthand sixteenth alveolus are bigger relative to (only occurring taxon as

the and still as in the authors the previous following ones, they are Cearadactylus), presented material thus smaller relative to the second, third and fourth first fully prepared post-cranial

the establishment of another taxon. alveoli. The biggest alveoli of the skull are posi- avoiding

tioned opposite to the smallest alveoli of the We interpret the first alveolus visible as the mandible; a pattern, if confirmed after prepara- fourth one on the left side. If however, we tion, that is not seen in the other discussed the of this with compare measurements specimen specimen that includes cranial alveoli (MN those of MN 4797-V the following observations 4797-V). Differences with the alveolar pattern in can be made. The first four measurementsof BSP

the skull of MN 4797-V are clear. MN 4797-Vhas a 1991 I 27 are comparable to those in MN 4797-V. distinct larger third alveolus where AMNH 24444 After that, the differences are slightly bigger. This has a larger ninthalveolus, lacking in MN 4797-V. questions the conclusions by Veldmeijer et al. (in whetherthe fourthalveolus is indeed the The Stuttgarter mandible (SMNS 55414; Fig. 4) review),

fourth and not the third because then the curves This small, partly prepared but largely complete would be more or less equal in pattern. We also mandible in the Staatliches Museum fur Natur- have to take into account that the data of the kunde, Stuttgart (SMNS 55414, Veldmeijer et al., Munich maxilla is based on average measure- in review), has been assigned to Brasileodactylus ments of both sides, in contrast to MN 4797-V as well, as it clearly shows diagnostic features for Differences which is only based on the right side. this The extremely elongated mandible genus. the with AMNH 24444 is basically same as clearly shows the dorsoventrally compressed described for MN 4797-V. anterior part with the expansion, the particular position of the first three alveoli and the mandibular with This groove sub-grooves. specimen Diastemae differs from Brasileodactylus araripensis in its smaller size and in its distal which is expansion Thediastemae of the measuredmandibles show a

andsmoother, theauthors do not longer although remarkably uniform picture, i.e. a strong increase think that the differences justify the erection may in size posteriorly, despite some small fluctua- of a new species. tions. As with the alveoli, the specimen AMNH

differs this the alveolus 24444 on point (for explanation, SMNS 55414 shows a small first (the see the description of the alveolar pattern). smallest of all known specimens) and a large second alveolus (the biggest of all known speci- of The pattern of the sizes the diastemae of the this with MN 4804-V which has mens); contrast skull in MN 4797-Vis remarkable similar, despite the third alveolus its The as biggest. following small differences, to those of the mandible. alveoli to the follows the and (up seventh) pattern Again, a difference can be noted for AMNH approximate size as seen in the holotype of Brasi- 24444. Here, a steady increase in diastemae size the ninth and tenth leodactylus araripensis; eigth, can be seen until the diastema 7-8. The following alveolus show the and are opposite pattern diastemae decrease in size at least until diastema smaller. slightly 9-10 after which the diastemae increase in size; it

remains uncertain whether this starts with

51 Brasileodactylus (Pterosauria, Pterodactyloidea, Anhangueridae);

an update

1990; for diastema 10-11 or with the following. Another species for Ornithocheiridae (Kellner,

an Kellner & Tomida, As strong decrease in size occurs with diastemae overview, see 2000).

14-15 and 15-16 after which again a strong pointed out elsewhere (Veldmeijer, 2003b),

last decrease is as a valid and increase in size occurs. A a sharp can Anhangueridae regarded taxon

after different from Ornithocheiridae; we with be seen with diastemae 19-20 and 20-21, agree

be the of which for the last time an increase in size can designation Ornithocheirus compressirostis

less relative the of Ornithocheiridae Kellner & noted, but far to foregoing ones. as type species (see Tomida, The of the O. 2000). acceptance compres- This means that, when the fluctuations are sirostris as the type species for Ornithocheiridae included, the size of diastemae increases first forces to include Brasileodactylus in Anhangue- posteriorly, but decreases from diastema 10-11 or ridae, as the laterally compressed jaws that 11-12. Full preparation of the specimens, allo- strongly decrease in width in anterior direction left needed wing the measurement of the side, is resulting in a sharp pointed beak in order confirm this The in to deviating pattern. Ornithocheiridae, contrast distinctly from the pattern of diastemae in BSP 1991 I 27 is, despite expanded and dorsoventrally compressed jaws with the small fluctuations, in line pattern seen in in Brasileodactylus. for the decreasein which MN 4797-V, except size, have starts later relative to the others (with diastema Various authors suggested Brasileodactylus

differ with 11-12 instead of 10-11 for MN 4797-V), but being synonymous Anhanguera (Unwin, markedly with AMNH 24444. These differences 2001 although in a later paper, Unwin (2002) still less the with the refers it are more or same as explained to as Brasileodactylus araripensis) or even comparison of the New York specimen with the Coloborhynchus (Frey et al., 2003). As remarked by of Crato specimen. Veldmeijer & Signore (2004) "The explanation

the supposed difference as theresult of ontogeny,

sexual dimorphism or variation cannot be

mainly due to the scanty fossil record Discussion proven, (most of the species are represented by only one in cases teethare not preserved and Firstly, many (published) specimen, often consisting only of

the alveolar diameter serves as indicator of tooth (parts of) the skull); the fossils should thereforebe size, though there need not necessarily be a posi- treated as different species unless proven (and tive relation between alveolar size and tooth size; not suggested) otherwise", hence the designation diameter the could a large of alveolus mean a is here as Brasileodactylus. Brasileodactylus long, perhaps fanglike tooth, but a short, regarded as a valid taxon, on the basis of lack of a bulb-like tooth could have been well. possible as premaxillary sagittal crest and a dentary sagittal On the other the teeth of the taxa discussed hand, crest and the of the presence unique configura-

here all have a dentition inferred comparable tion of dentition and morphology of the dorsal from few of teeth and the the (remnants) assump- above. The aspect of thelower jaw, as pointed out tion that the the alveolus, the the bigger longer hitherto most complete cranial dentition cannot tooth, is reasonable. The second is that problem be assigned to Brasileodactylus with full confi- of the and measurements teeth/alveoli of dence yet (Veldmeijer, 2003a), but the features almost have been diastemae never published of the and lower the dentition upper jaw in MN Veld- (exceptions are Fastnacht, 2001; Lee, 1994; 4797-V show of the a more erratic pattern skull, 2003b), rendering scientific evaluation meijer, with larger differences in diameter of alveoli: the impossible. Therefore, the data of the herein second, third and fourth alveoli are the biggest, have been described specimens presented, followed smaller after which by two ones, again

described and discussed as well as shortly impor- but less than the second until bigger ones, big tant morphological peculiarities. fourth alveoli, follow. Comparison with other taxa(mainly Anhanguera andColoborhynchus) will The situation on family (in a narrow sense) level

have to shed on the conformities and diffe- is complex. The Ornithocheiridae are primarily light

rences in dentition of these related taxa composed of taxa from the Cambridge closely and whether dentition be used to Greensands; the Anhagueridae mainly from taxa patterns can differentiate certain taxonomic level. from the Araripe Basin. The two taxonomic units on a

have been synonymised by various authors (for Furthermore, it can be suggested that if AMNH instance Unwin, 2001) but this is based on the 24444 can be classified as Brasileodactylu s as difference in view of the assignment of the type suggested in the preliminary description and if it

52 CRANIUM 22,1 (2005)

The dentition pattern of the various specimens showing the size distribution of the alveoli as described in the text;

number of alveolus Y-axis size in mm, X-axis

de verschillende de de alveolen zoals Hetdentitiepatroon van specimens met grootte-verdeling van (tandkassen)

de alveole beschreven in de tekst; Y-aslgrootte in mm, X-as nummer van

distribution of the diastemae described the The dentitionpattern of the various specimens showing the size as in Y-axis text; size in mm, X-axis number of diastema

verschillende de de diastamen Hetdentitiepatroon van de specimens met grootte-verdeling van (tandenloze ruimtes) zoals beschreven in de tekst; Y-as grootte in mm, X-as nummer van het diasteem

53 Brasileodactylus (Pterosauria, Pterodactyloidea, Anhangueridae);

an update

be established that AMNH 24444 and Marco can Signore della Ludodactylus sibbicki are the same (which seems to Dipartimento di Scienze Terra

the basis of the observations Universita Studi di "Federico II" be so on preliminary degli Napoli the first S. Marcellino 10 on the half preparated holotype by L.go

author in 1998), Ludodactylus sibbicki should be 80132 Napoli Brasileodac- re-named as Brasileodactylus (either Italy

tylus araripensis or Brasileodactyus sibbicki). [email protected]

Finally, the generally larger alveoli in the skull of H.J.M. Meijer AMNH 24444 relative to MN 4797-V might be Faculty of Human MovementSciences explained by the fact that AMNH24444 is ontoge- Free University of Amsterdam

relative to 4797-V. This Van der Boechorststraat 9 netically younger MN however, cannot be the explanation for the large 1081 BT Amsterdam difference in size between AMNH 24444and BSP [email protected]

1991 I 27, as the latter is regarded as not fully

well. grown as

Acknowledgments

the AMNH 24444 was shipped to Netherlands of and is currently being prepared with courtesy the American Museum of Natural History, New York, USA (Mark Norell and Carl Mehling), and which has kindly been made possible by the

Natuurmuseum Rotterdam, the Netherlands.

AJV is grateful to M. Dorling, E. Frey, A.W.A.

Kellner, H. Mayr, A.C. Milner, S. Nabana, M.A.

Norell, U. Oberli, Y. Okazaki, M. Oshima, Y.

Takakuwa, Y. Tomida, P. Wellnhofer, R. Wild for

kindly allowing access to the collections under

their care. Material in Berlin remained inacces-

sible, unfortunately. The study of various collections has been made possible with financial

support to AJV by the Jan Joost ter Pelkwijkfonds, Stichting Molengraaff Fonds, Mej. A.M Buiten- dijkfonds and Mr. & Mrs. Endenburg; the study

of the material in various collections in Japan was

made possibly by the Netherlands Organization

for Scientific Research (NWO). Due to the grant of the Egypt Exploration Society for studying

archaeological material in Cambridge, AJV was of the from the able to study some type specimens Cambridge Greensands. We thank E. Endenburg for his all round help.

Adresses of the authors

Andre J. Veldmeijer (corresponding author)

Natuurmuseum Rotterdam

PalArch Foundation

Mezquitalaan 23

1064 NS Amsterdam

The Netherlands

[email protected]

54 CRANIUM 22,1 (2005)

Literature

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