Mitt. Mus. Nat.kd . Berl., Geowiss. Reihe 2 (1999) 121-134 19.10.1999

On a Pterosaur Jaw from the Upper Jurassic of Tendaguru (Tanzania)

David M. Unwint & Wolf-Dieter Heinrich'

With 3 figures

Abstract

A short section of a mandibular symphysis is the first cranial fossil of a pterosaur to be reported from the Upper Jurassic of Tendaguru, Tanzania. It is made the holotype of a new dsungaripteroid pterosaur, Tendaguripterus recki n. gen. n. sp . Al l previously named pterosaur taxa from Tendaguru are shown to be nomina dubia. The pterosaur assemblage from Tendaguru contains a `rhamphorhynchoid', as well as the dsungaripteroid, and is similar in its systematic composition to other Late Jurassic pterosaur assemblages from Laurasia. The diversity and broad distribution of dsungaripteroids in the Late Jurassic suggests that the group was already well established by this time.

Key words: Tendaguru, Tanzania, Upper Jurassic, pterosaur, Pterodactyloidea, Dsungaripteroidea.

Zusammenfassung

Der erste Schädelrest eines Flugsauriers aus dem Oberjura von Tendaguru in Tansania wird beschrieben. Bei dem Fundstück handelt es sich um ein bezahntes Unterkieferbruchstück aus der Symphysenregion . Der Fund gehört zu einem neuen Taxon, das als Tendaguripterus recki n. gen. n. sp. bezeichnet und zur Überfamilie Dsungaripteroidea gestellt wird. Alle zuvor aus den Tendaguru-Schichten beschriebenen Taxa werden als nomina dubia angesehen. In Tendaguru sind Verteter der ,Rham- phorhynchoidea` und Dsungaripteroidea nachgewiesen . Diese systematische Zusammensetzung ist derjenigen anderer Flug- saurier-Vergesellschaftungen der späten Jura-Zeit ähnlich . Die Vielfalt und die weite Verbreitung der Dsungaripteroidea in Laurasia läßt darauf schließen, daß sich diese Flugsauriergruppe bereits in der späten Jura-Zeit erfolgreich durchgesetzt hatte.

Schlüsselwörter: Tendaguru, Tansania, Oberjura, Flugsaurier, Pterodactyloidea, Dsungaripteroidea.

Introduction and it was supposed, on the basis of size differ- ences, that they represented two different taxa African pterosaur remains are exceptionally rare. (Stremme cited in Janensch, 1914a). Parkinson A few isolated bones have been found at various (1930: 145) also mentioned pterosaur bones in Cretaceous localities (Swinton 1948, Monteillet his account of the British expeditions to Tenda- et al. 1982, Kellner and Mader 1996, Mader and guru. Reck (1931) published a short account of Kellner 1997, 1999, Sereno et al. 1998, Martill the material in the MfNHUB collections in pers comm. 1999), but only one location, Tenda- which he described a few of the more com- guru, has yielded Jurassic material. plete and better preserved bones and designated Most of the pterosaur fossil remains from some of them as holotypes for four new species Tendaguru were collected from the Upper Sau- of pterosaur : `Rhamphorhynchus tendagurensis', rian beds by the German led expeditions of 1909 `Pterodactylus maximus', `Pterodactylus brancai', to 1913, and are currently housed in the Mu- and `Pterodactylus arningi'. Subsequently, Galton seum für Naturkunde of the Humboldt Univer- (1980) published descriptions of tibiotarsi from sity, Berlin (MfNHUB) . The British expeditions Tendaguru and assigned `Pterodactylus brancai' of 1924-1931 also collected a few pterosaur Reck, to Dsungaripterus on the basis of similari- bones, now housed in the Natural History Mu- ties between the African and Chinese material. seum, London. The first mention of pterosaur At present, all Tendaguru pterosaur taxa are bones was by Janensch (1914a: 82) in his report based on isolated elements from the fore and on the vertebrate fauna of the Tendaguru Beds, hind limbs. It is rare, however, for such material

1 Museum für Naturkunde, Institut für Paläontologie, Invalidenstraße 43, D-10115 Berlin, Germany. Received January 1999, accepted March 1999 122 Unwin, D. M. & W-D. Heinrich, Pterosaur Jaw of Tendaguru to exhibit characters that are diagnostic at low isolated and not so abundant as in the Upper taxonomic levels and Wellnhofer (1978) has al- Saurian Bed. The Middle and Upper Saurian ready raised doubts regarding the validity of beds together with the intervening Transitional some of the Tendaguru taxa. By contrast, skull Sands are generally considered to be Kimmerid- remains and, in particular, the jaws and denti- gian to Tithonian (e.g., Aitken 1961, Russell tion, often exhibit distinctive characters that are et al. 1980). Pterosaurs do not appear to have critical for diagnosing species and genera (Welln- been recovered from the Lower Saurian Bed, or hofer 1978, 1991) . Recently, during preparation from the three marine units, but this may be of a block of matrix from the Middle Saurian due, at least in part, to collecting bias. bed, as part of a programme of microvertebrate The pterosaur localities : Pterosaurs have small sampling, one of us (W.-D. H.) found a been found at a number of sites in the Tenda- M. How- bone that was preliminarily identified by guru area (Fig. 1B). These are as follows: gate, London (unpublished note of May 2, 1986, Catalogue of Fossil Reptiles, Institute of Paleon- S ite J g (W j) : located 2.3 km north-north-west tology, MfNHUB) as a lower jaw fragment of a of Tendaguru hill, excavations began at this site pterodactyloid pterosaur bearing four teeth. The in 1910 and continued in 1911 and 1912. The site block was treated with acetic acid to free the was most intensively collected in 1912, under the fossil from the matrix and preparation was com- direction of H. Reek, and it seems likely that the pleted by H.-H. Krueger (MfNHUB) who em- sediment that yielded the pterosaur material de- ployed mechanical preparation to remove re- scribed below was collected during this period. maining sediment adhering to the jaw. Site 69a : Reek (1931) does not give the exact Here we describe this specimen and comment location of this site, but it was situated south of on its significance for the systematics of ptero- Tendaguru Hill, approximately at the same posi- saur remains from Tendaguru. tion as the Fraas camp in 1908. The locality, which is quoted in the GTE-field catalogue (p. 133) as 'Graben mit Vogelknochen' (trench Abbreviations with bird bones), produced fossil material from a horizon high in the Upper Saurian Bed (Reek BSP, Bayerische Staatssammlung für Paläontolo- 1931 :324) and is the type locality for `Rham- gie, München; GTE, German Tendaguru Expedi- phorhynchus tendagurensis'. tion; MB, Museum für Naturkunde, Berlin; MGUH, Geological Museum, University of Co- Site B : This locality, the type locality of `Ptero- 328-330) was penhagen; LAMING, National Museum of Ire- dactylus maximus' (Reek 1931 : land, Geological Section. situated about 1 km south-south-east of Tenda- guru Hill and produced remains from the Upper Saurian Bed. Geology, localities and taphonomy Site MD : This locality, indicated by Janensch in his site map published in 1925, was situated Geology : The Tendaguru Beds, well known for about 2 km north-east of Tendaguru Hill. Here, their extensive dinosaur assemblage, outcrop in in 1912, sediments of the Transitional Sands at south-eastern Tanzania (Fig. 1) . Janensch (1914b) the base of the Upper Saurian Bed produced the gave a detailed description of the sedimentary type material of `Pterodactylus brancai' and succession, dividing it into Lower, Middle, and many other bones and teeth of pterosaurs which Upper Saurian Beds, which were separated by were found relatively close to a well preserved three marine sandstone units: the Nerinea Bed, vertebral column of Barosaurus africanus (Reek the Trigonia smeei Bed, and the Trigonia schwarzi 1931). Bed (Fig. 1D). Transitional deposits between the Site IV : This excavation, in sediments of the Saurian beds and the marine units were formally Upper Saurian Bed, was situated about 1 km recognised as the Transitional Sands (Übergangs- south-south-east of Tendaguru Hill, close to Site schichten) (see Heinrich this volume, a) . B, and is the type locality of `Pterodactylus arnin- The majority of pterosaur remains appear to gi' (Reek 1931). have been recovered from the Upper Saurian Bed and the Transitional Sands at the base of Site T: This pterosaur locality is indicated on the Upper Saurian Bed (Reek 1931) . They were the site map by Janensch (1925), but was not also found in the Middle Saurian Bed, but were mentioned by Reek (1931).

Mitt . Mus. Nat.kd. Berl ., Geowiss. Reihe 2 (1999) 123

D Trigonia schwarzi Bed

Upper

Saurian

Bed

TS TS Trigonia

smeei

Bed

_TS TS

Saurian

Bed _TS TS

Nerinea

Bed

TS TS

E USB L~~- TSMB E 01 \ " \ \ MSB l f J l l I J ! J f f l f f J J l f f ~_ NB f f f I f f f f f f f f f f f f f f f f f f ! f I ! f ! f f f f f f I f ! f f f f I f f f f f f ! ! f f f f f f f f f f f f f f J f ! f f f ! f I f ! f U) _H I l f ! ! ! ! ! f f ! ! ! ! ! ! ! f f f f f f f i ! I I I I ! ! I I f I J / C:)-

Fig . 1. Locality map for pterosaur remains found at Tendaguru. A, Africa, showing the location of Tanzania. B, Geology of the Tendaguru region indicating the relationships of the Saurian Beds. Inset map shows the location of Tendaguru within Tanzania. C, simplified stratigraphic cross-section through Tendaguru showing the topographic relationships of sediments that have yielded pterosaur remains. D, stratigraphic column showing, in diagrammatic form, the relationships of the main Upper Jurassic horizons at Tendaguru. Pterosaur localities : IV, B, Jg (Wj), MD and T. Abbreviations: G, Gneiss; LSB, Lower Saurian Bed; MSB, Middle Saurian Bed ; NB, Nerinea Bed ; TS Transitional sands; TSMB, Trigonia smeei Bed; TSB, Trigonia schwarzi Bed; USB, Upper Saurian Beds

124 Unwin, D. M. & W-D. Heinrich, Pterosaur Jaw of Tendaguru

Site M2 : This British expedition locality (Par- Family Germanodactylidae kinson 1930: 145) was at Nguruwe, on the way Remarks : The Germanodactylidae currently from Tendaguru Hill to Namunda. However, contains two named genera: Germanodactylus nothing is known about the material from this Young, 1964 and Normannognathus Buffetaut, site (Reek 1931: 325) . Lepage & Lepage, 1998. It seems likely that Other Tendaguru sites : Reek (1931: 325) Germanodactylidae is paraphyletic since some of mentioned that localities situated along the steep the included taxa seem to be closer to the Dsun- scarp slope extending from Nguruwe in the garipteridae than they are to other germanodac- south to the ornithopod site at Kindope in the tylids. However, this has yet to be formally de- north had produced single pterosaur bones, but monstrated, so, for the present, we continue to did not give the precise locations of these sites. use the family name Germanodactylidae. Reek noted (1931: 325) that with regard to the importance, quality and number of finds, that Tendaguripterus n. gen. sites 69a and MD were the most important Type and only known species : Tendaguri- pterosaur localities. He also stated (1931 : 325) pterus recki n. gen. n. sp. that all levels of the Upper Saurian Beds yielded single bones, that a small number of bones were Derivation o f name : Combination of Tenda- discovered, by chance, in the dinosaur trenches guru, the type locality in Tanzania, and `ptero' and that the latter was also true for the Middle (Gr.) meaning wing. Saurian Beds. Ta p ho n o my : Most of the pterosaur material recovered from the Tendaguru beds consists of isolated bones and according to Reek (1931) ar- ticulated remains were not found. However, ac- cumulations of irregularly jumbled bones were discovered at Site 69a, where they were thought to represent one individual, and at Site MD, where they were thought to represent a `swarm' of pterosaurs (Reek 1931) . The MfNHUB collec- tions are dominated by postcranial remains, mainly from the forelimbs, though hind limb ele- ments and some incomplete vertebrae are also present. A few teeth were recovered, but skull remains are extremely rare and seem to be con- fined to the specimen described here and a poorly preserved bone fragment of what appears to be the caudal end of a mandible (MB. unum- bered) . Generally, the limb bones are well pre- served, and, in contrast to material from other Late Jurassic localities such as Solnhofen, they are usually uncrushed and undistorted.

Systematic palaeontology

Order Pterosauria Kaup, 1834 Suborder Pterodactyloidea Plieninger, 1901 Superfamily Dsungaripteroidea Young, 1964 Remarks: The Dsungaripteroidea is diagnosed by a series of cranial characters (see Unwin 1995, Unwin & Lii 1997) and is currently com- Fig . 2. Tendaguripterus recki n. gen. n. sp., holotype, posed of the families Germanodactylidae and MB.R.1290, section of the mandibular symphysis in (A) left Dsungaripteridae. lateral, (B) right lateral and (C) right dorsolateral view

Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 2 (1999) 125

Diagnosis : Posterior section of mandibular Table 1 symphysis with deeply concave ventral profile. Dimensions of the holotype of Tendaguripterus recki n. gen. Teeth in the posterior section of the mandibular n. sp. (in mm) symphysis inclined posterodorsally. Length' 25.9 Maximum depth anterior end' 6.0 Maximum Tendaguripterus recki n. gen. n. sp. depth posterior end' 9.9 Crown height fourth tooth, right side 6.0 Figs 2, 3 Maximum basal tooth diameter2 Derivation of name : The specific name Left side, tooth position # 1 11.6 honours Hans Reck, who was the first to de- # 2 21.5 scribe pterosaurs from Tendaguru. # 3 31 .8 # 4 41.8 Ho 1 o ty p e : An incomplete section of the man- Right side, tooth position # 1 11.6 dibular symphysis, MB.R.1290. # 2 21.7 # 3 31.8 Type locality : Excavation site Jg (Wj), Ten- # 4 41.75 daguru, Tanzania (Fig. 1B) . Gap length between tooth positions3 Type horizon : Middle Saurian Beds, Tenda- Left side positions # 1-2 3.6 # 2-3 4.4 guru (the exact stratigraphic position is un- # 3-4 4.5 known). Right side positions # 1-2 3.2 # 2-3 4.0 Age : Upper Jurassic, Kimmeridgian-Tithonian. # 3-4 4.15

Dimensions : See Table l. Relative size of inter-tooth spacing4 Left side positions # 1-2 2.25 # 2-3 2.94 # 3-4 2.5 Right side positions # 1-2 2.0 # 2-3 2.35 # 3-4 2.37

Associated fauna : Site Jg (Wj) was one of the most productive Tendaguru localites yielding dinosaurs (Brachiosaurus, Elaphrosaurus, Dryo- saurus and Kentrosaurus), a paramacellodid liz- ard (Broschinski, this volume) and mammals (Tendagurodon, Tendagurutherium and a hara- miyid: Heinrich this volume, b) . Diagnosis : As for the genus.

Description

B Preservation : The holotype material consists of a short section (25 .9 mm long, as preserved) of the lower jaw symphysis. The anterior termi- nation of the symphysis is missing. Comparison with related taxa such as Germanodactylus (Wellnhofer 1970) suggests that the missing ante- rior portion was at least 15 mm long. Judging by the apparent widening of the symphysis posteri- C

1 As preserved. Fig. 3. Tendaguripterus recki n. gen. n. sp., holotype, 2 Measured from anterior edge to posterior edge. MBR.1290, section of the mandibular symphysis in (A) left 3 Gap length is measured from the posterior edge of the lateral, (B) right dorsolateral and (C) palatal view. Scale bar leading tooth to the anterior edge of the following tooth. = 5 mm 4 Ratio of gap length to basal diameter of leading tooth. 126 Unwin, D. M. & W-D. Heinrich, Pterosaur Jaw of Tendaguru orly it seems likely that this bone represents the edge. In lateral profile this edge has a distinctive posterior part of the symphysis immediately concave outline reflecting the deepening of the anterior to the point at which the left and right symphysis posteriorly. The palatal surface of the mandible diverged. symphysis is flat posteriorly, but, even taking The fragment retains some of its three dimen- compression and distortion into account, the sional shape (as with many other pterosaur re- anterior two thirds of this surface appears origi- mains from Tendaguru), but it has been some- nally to have been somewhat depressed along what compressed, so that the palatal surface now the mid-line, forming a shallow groove whose slants down towards the right side and the jaw sides sloped gently towards the dental margins has a somewhat asymmetric outline in anterior of the symphysis. The presence of a midline or posterior aspect. The specimen is transected groove confirms that the fossil represents the by three prominent fractures. The bone surfaces lower jaw since, in pterosaurs, the palatal surface are reasonably well preserved, though eroded in of the symphysis often bears a midline groove, places, and somewhat checked and fractured, a or is flat, whereas the palatal surface of the common feature of the relatively thin compacta upper jaw has a midline ridge, or is flat. of pterosaur bones. A distinctive feature of the specimen is the The left dental margin bears four alveoli. The expansion of the margins of each dental alveolus anterior two alveoli contain teeth lacking the tips to form a low, rounded tumescence, from which of the crowns, but the teeth in the posterior two the tooth erupts. This is most clearly exhibited alveoli are broken off at their base, though in on the left side of the symphysis (Figs 2A, C, each case the tooth root remains. The right den- 3A). In lateral view each alveolus is surrounded tal margin bears four complete alveoli and the by a low boss of bone, while the dental margin remains of a fifth adjacent to the posterior break. between these bosses is gently concave : as a re- Each of the first four alveoli contains a tooth: the sult, the dorsal profile of the symphysis has a first tooth lacks its tip, the second and third lack distinctive scalloped appearance in lateral view. the distal half of the crown, but the fourth ap- The teeth are relatively slender (the one com- pears to be complete. Only a small portion of plete tooth has a crown/maximum base width the medial margin of the fifth alveolus remains. ratio of 3.0), sharp-pointed and gently recurved, It is not possible to determine the absolute posi- each exhibiting about the same degree of recur- tion of these teeth with respect to the anterior or vature. The outer, labial surface is gently convex, posterior end of the tooth row, hence the first while the inner, lingual surface is flatter. The preserved pair are nominally referred to here as labial and lingual surfaces meet in rounded an- pair one, the second as pair two and so on. terior and posterior margins and carinae are not developed. Viewed from anterior the teeth ap- Anatomical description : The preserved re- pear to have stood approximately vertically in mains indicate that this pterosaur had a rela- the jaw. However, when viewed from either the tively long symphysis that bore at least five pairs left or the right side, the teeth are clearly in- of dental alveoli. Anteriorly, the symphysis ap- clined toward the rear at a steep angle, hence, in pears to have been relatively narrow, with a tri- this part of the jaw at least, the dentition was angular cross-section that is somewhat deeper raked backward. This does not appear to be an than broad. The depth of the symphysis at the artefact of preservation since the same type of anterior break corresponds to the total height of orientation is evident in all reasonably well pre- the complete tooth crown in the fourth right al- served teeth, irrespective of the distortion ex- veolus. Proceeding posteriorly there is a rapid perienced by different parts of the jaw and the increase in the depth of the symphysis and at the dental margins. posterior break it is more than 1 .5 times deeper All preserved teeth appear to have been fully than at the anterior break. This increase in erupted and no replacement teeth are visible, depth is offset by widening of the symphysis, though in the case of the third and fourth posi- which begins just prior to the posterior break so tion on the right side of the jaw the point at that, when restored to its original shape, the which replacement teeth should appear (in the cross-section at this point was a tall, narrow tri- posteromedial margin of the alveolus) is ob- angle of similar proportions to the profile of the scured by sediment. The teeth show a small but anterior break. consistent increase in basal diameter proceeding The lateral surfaces of the symphysis were flat posteriorly (see Tab. 1), the fourth pair being or slightly convex and met ventrally in a sharp 1 .125x the size of the first pair. In addition, the Mitt . Mus. Nat.kd. Berl ., Geowiss. Reihe 2 (1999) 127 teeth are relatively widely spaced: generally the taxa, such as Tropeognathus (Wellnhofer 1987) inter-tooth spacing (measured from the posterior and Pteranodon (Eaton 1910), the ventral mar- edge of the leading tooth to the anterior edge of gin is also concave in this region, but this is asso- the trailing tooth) is equivalent to at least twice ciated with anterior deepening of the lower jaw, the basal diameter of the leading tooth (Table 1). not posterior deepening, as in the case of dsun- In addition, the posteriorward increase in tooth garipteroids. The ventral margin of the sym- size is accompanied by a small but consistent in- physis in Tendaguripterus n. gen. has a lateral crease in inter-tooth spacing (see Table 1) . profile that is more concave than in any known The jaw fragment of Tendaguripterus recki dsungaripteroid, though this shape may have n. gen. n. sp. is slightly smaller than the corre- been overemphasised by postmortal deformation sponding region of the lower jaw of Germano- of the specimen . dactylus rhamphastinus (BSP AS I 745) . As- (2) The margins of the dental alveoli are ex- suming that Tendaguripterus n. gen. broadly panded to form low, rounded turnescences, from resembled Germanodactylus (see below) this which the teeth erupt: thus, when viewed later- suggests an original skull length of somewhat ally, the dental margin has a scalloped profile. less than 200 mm and a wingspan approaching This feature is particularly well developed in 1000 mm. Dsungaripterus (Young 1964, Fig. 2) and in the Tatal dsungaripterid, and is present, though only weakly developed, in Germanodactylus cristatus Comparison (BSP 1892 IV 1) and G. rhamphastinus (BSP AS I 745 ; BSP 1977 XIX 1). Inflated alveolar rims Various features indicate that the find is ptero- also occur in the Cretaceous pterosaur `Ptero- saurian: the smooth outer surface of the bone dactylus' giganteus (Owen 1851, pl. 38, Figs 4-6) cortex, the thin bone walls of the jaw fragment, and `Pterodactylus' sagittirostris (Owen 1874, which is otherwise hollow, and the tooth mor- p1.2, Figs 1-8). In these cases, however, the phology, relatively long, simple, sharp-pointed expansion takes the form of a pedestal-like teeth lacking any serrations or carmae. The structure, rather than the rounded tumescence elongate mandibular symphysis shows that the seen in dsungaripteroids . The dental alveoli of specimen is pterodactyloid, since `rhamphorhyn- Tendaguripterus n. gen. are tumescent, as in choids' either have a very short mandibular sym- dsungaripteroids, and more inflated than in physis, or no symphysis at all. Germanodactylus, but less so than in dsungari- Recent phylogenetic analyses have identified pterids. four major clades of pterodactyloids (Unwin & (3) The largest teeth occur toward, or at, the Lü 1997): Ornithocheiroidea, Seeley, 1901, Cte- posterior end of the tooth row. In Dsungari- nochasmatoidea Kuhn, 1967, Dsungaripteroidea pterus (Young 1964, Fig. 2) and in the Tatal dsun- Young, 1964, and Azhdarchoidea Unwin, 1992. garipterid (Wellnhofer 1991, p. 120) the smallest The jaw described here lacks any diagnostic fea- teeth occur at the anterior end of the tooth row tures of Ornithocheiroidea, Ctenochasmatoidea, and there is a steady increase in size posteriorly, or Azhdarchoidea (see Unwin 1995 and Unwin the largest teeth occurring at the caudal end of & Lü 1997 for apomorphies of these taxa) but the tooth row. In Germanodactylus the largest exhibits three characters only found in dsunga- teeth occur in the middle, or just posterior to ripteroids. These are as follows: the middle of the tooth row (Wellnhofer 1970, (1) A relatively deep lower jaw which, when Figs 12, 13) . In other pterodactyloids the largest viewed laterally, has a ventral margin that is dis- teeth usually occur at or toward the anterior end tinctly concave toward the posterior end of the of the tooth row. The complete tooth row is not symphysis. This is evident in Dsungaripterus preserved in Tendaguripterus n. gen., but the (Young, 1964, Fig. 2) and the dsungaripterid steady increase in size, caudally, and the location from Tatal, Mongolia (Bakhurina pers comm.), of the largest preserved tooth adjacent to the and is also visible, though less pronounced, in posterior end of the symphysis corresponds to Germanodactylus cristatus (Wellnhofer 1970, the situation in Germanodactylus where the lar- Fig. 12) though not in G. rhamphastinus (Welln- gest tooth pair is usually located in the region of hofer 1970, Fig. 13). By contrast, in other ptero- the posterior end of the symphysis. This suggests dactyloids, the ventral margin of the posterior that the largest teeth of Tendaguripterus n. gen. symphysial region usually has a flat or a convex may also have been located in the posterior half profile (Wellnhofer 1978, Figs 4, 5). In some of the tooth row, as in dsungaripteroids. 128 Unwin, D. M. & W-D. Heinrich, Pterosaur Jaw of Tendaguru

Tendaguripterus n. gen. is closely comparable Dsungaripteridae . The Jurassic dsungaripteroids with dsungaripteroids in other respects, though including Germanodactylus, Normannognathus these features may not be unique to this clade. and Tendaguripterus n. gen. lack apomorphies of The rapid broadening of the lower jaw at the the Dsungaripteridae, but their relationships to posterior end of the symphysis is also evident in each other and to the Dsungaripteridae are dif- Dsungaripterus (Young 1964, Fig. 2) and the ficult to resolve, principally because Normanno- tooth morphology is almost identical to that gnathus and Tendaguripterus n. gen. are so of Germanodactylus rhamphastinus (Wellnhofer poorly known. Moreover, the distribution of 1970, Fig. 13) though this tooth type occurs else- characters, at least as determinable from pre- where in pterodactyloids. More importantly, in served remains, exhibits some incongruency. Tendaguripterus n. gen. the teeth are separated Thus, as Buffetaut et al. (1998) have pointed out, by relatively large inter-tooth spaces (Table 1), Normannognathus has a cranial crest whose mor- that are always more than twice the size of the phology is similar to that of Dsungaripterus and basal diameter of the leading tooth. Relatively is clearly more derived than that of Germano- large inter-tooth spacing also occurs in all known dactylus, yet the dentition of Normannognathus dsungaripteroids (Young, 1964, Fig. 2; Wellnhofer is less derived than that of G. cristatus. The latter 1970, Figs 12, 13, Wellnhofer 1991, p. 120; Buffe- has toothless jaw tips and short, rather broad taut et al. 1998, Fig. 1), though it seems to be teeth whereas the dentition extends to the tip of somewhat less in the corresponding region of the the jaw in Normannognathus and the teeth are dentition in these taxa. By contrast, in most relatively slender - the typical condition for Jurassic pterodactyloids, the inter-tooth spacing early pterodactyloids. A similar character incon- is usually little more than the basal diameter of gruency occurs in Tendaguripterus n. gen. In this the leading tooth (e.g. Pterodactylus), and some- genus the rims of the dental alveoli show a times even less than a single tooth diameter (e.g. marked expansion, but the teeth are relatively Ctenochasma) . slender and sharp-pointed, like those of other A number of features distinguish Tendaguri- early pterodactyloids. Germanodactylus cristatus, pterus n. gen. from other dsungaripteroids. The by contrast, has only very weakly developed al- concavity of the ventral margin, when seen in veolar rims, but a tooth morphology that is more lateral view, is deeper than in any other dsunga- like that of dsungaripterids than that of any of ripteroid and the teeth are raked backwards . In other Jurassic dsungaripteroid. It is doubtful that Germanodactylus the teeth stand vertically or the interrelationships of Tendaguripterus n. gen ., are procumbent in the anterior end of the denti- Germanodactylus and Normannognathus can be tion and the mandibles are only gently sigmoidal resolved until more complete material is avail- (Wellnhofer 1970) . In Normannognathus the able, thus, for the present, we assign these taxa teeth also stood vertically (Buffetaut et al. 1998) to the Germanodactylidae, while noting that and appear to have been more closely spaced ultimately this taxon may be shown to be para- than in Tendaguripterus n. gen. In later dsunga- phyletic. ripteroids (the Mongolian form, Dsungaripterus and Noripterus) the mandibular teeth are verti- cally oriented and the teeth are much more ro- A reassessment of other pterosaur taxa bust (shorter and more broadly based) than in from Tendaguru Tendaguripterus n. gen. (cf. Wellnhofer 1991) . In addition, these forms also have a relatively deep Reck named four species of pterosaur from mandible. Tendaguru, a `rhamphorhynchoid', `Rhampho- rhynchus tendagurensis', and three pterodacty- loids, `Pterodactylus arningi', `Pterodactylus maxi- Relationships of Tendaguripterus n. gen. mus' and `Pterodactylus brancai'. to other dsungaripteroids `Rhamphorhynchus tendagurensis' (MB.R.2845) was based on the associated, and well preserved, The interrelationships of dsungaripteroids are but incomplete distal end of a right radius and ulna still only poorly understood. Dsungaripterus and (Reck 1931, Fig. 1) . This material has a number the Tatal dsungaripterid share a number of de- of characters that are found in `rhamphorhyn- rived features (such as the occurrence of the lar- choids', but are absent in pterodactyloids: gest teeth at the caudal end of the tooth row) (1) The distal end of the radius is of similar that unites them in a monophyletic taxon, the height (measured dorsoventrally) to the ulna.

Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 2 (1999) 129

This is typical of `rhamphorhynchoids' (Welln- teristic in pterosaurs. The distal end of the ulna hofer 1978, Fig. 11), but not pterodactyloids, is expanded and bears a well developed articular where the distal end of the ulna tends to be surface consisting of a large dorsal facet, a small more robust and a little deeper to much deeper tubercle (now badly eroded) at mid-height and a than the radius (Wellnhofer 1978, Fig. 11). small fovea located immediately below the tuber- (2) The articular surface on the distal end of cle. These features show that the specimen repre- the ulna is of relatively simple construction, as in sents a pterodactyloid. Pterodactyloids have two `rhamphorhynchoids' (e.g. Dorygnathus, MB distinct ulna morphologies. In ornithocheiroids, 1977.21), and consists of a single, large, roller- the dorsal facet occupies only one half, or less, like structure . In pterodactyloids, the articular of the entire distal articular surface (Wellnhofer surface is more complex with a large, convex, 1985, Fig. 9). By contrast, in dsungaripteroids dorsal facet, a tuberculum at mid-height and a (e.g. Young 1964, Fig. 4a), azhdarchoids (e.g. shallow fovea ventral to the tuberculum (e.g. Nessov 1997, pl. 15, Fig. 16), and possibly cteno- Wellnhofer 1985, Figs 9, 25) . These features are chasmatoids, the dorsal facet occupies consider- clearly absent in MB.R.2845 (though present in ably more than half the articular surface and the some other ulnae from Tendaguru - see below) tuberculum and fovea are relatively small struc- suggesting that these remains are not pterodacty- tures. The Tendaguru specimen MB.R.2844, with loid. its prominent dorsal facet clearly belongs to the (3) Immediately adjacent to its distal termina- second type, but it does not bear any features tion, the anterodorsal margin of the ulna shaft that might help to further resolve its taxonomic bears a large, well-marked tendon groove, identity. Moreover, it does not have any cha- bounded dorsally by a prominent 'lip-like' pro- racters that distinguish it from other pterodacty- cess. This feature is evident in `rhamphorhynch- loids (other than ornithocheiroids) thus we con- oids' (e.g. Rhamphorhynchus : BMNH R42380 sider `Pterodactylus maximum' to be a nomen and Eudimorphodon ranzii, Wild 1978, Fig. 17) dubium. but is absent, or only weakly developed in ptero- `Pterodactylus brancai' was erected on the dactyloids. basis of a left tibiotarsus (MB.R.2842) and a left In other respects, such as their general mor- wing-phalanx one (MB.R.2848), though the asso- phology and relative dimensions, these bones ciation of these remains is unclear: they were also compare well with those of `rhampho- found at the same locality, but it is not known if rhynchoids' such as Rhamphorhynchus (Welln- they came from the same individual. The tibio- hofer 1970), Dorygnathus (MB 1977 .21) and tarsus (Reck 1931, Galton 1980) is almost cer- Eudimorphodon (Wild 1978, Fig. 17). Reck iden- tainly pterodactyloid, since the fibula is relatively tified these remains as a new species of Rham- short, and fuses with the shaft of the tibia just phorhynchus, but, comparison with the corre- beyond the mid length point of the latter - a sponding region in well preserved specimens diagnostic feature of this clade. This specimen of this pterosaur (MGUH VP 2304; BMNH also compares closely in other respects to the R42380) reveals at least one clear difference. In tibiotarsi of pterodactyloids and, in particular, Rhamphorhynchus, the distal articular surface of shows a strong resemblance to the corresponding the ulna is divided into two facets, the lower of element in the dsungaripteroids Germanodacty- which is saddle-shaped. This is quite unlike lus (BSP 1892 IV 1, BSP AS 1 745, NMING MB.R.2845 which has a single, continuous facet, 15005), Dsungaripterus (Young 1964, Fig. 7), the ventral half of which forms a tapering roller- Noripterus (Young 1973, Fig. 5) and the Tatal like structure . Further work on pterosaur phylo- dsungaripterid (Bakhurina 1982, Fig. 1). The geny might ultimately allow MB.R.2845 to be main distinction between the tibiotarsi of these assigned to a particular clade within `Rhampho- taxa and that of `Pterodactylus brancai' is size, rhynchoidea', but, until this can be established but otherwise they are remarkably similar. The the specimen is referred to `Rhamphorhynchoi- Tendaguru specimen does not exhibit any char- dea' incertae sedis. acters that might enable it to be assigned to any `Pterodactylus maximus' is based on a large particular genus of dsungaripteroid, nor does it right ulna (MB.R.2844), 198 mm long. The bone have any features that distinguish it from these is uncrushed, but each articular end is somewhat taxa, thus, for the present, we identify it as eroded. Reck diagnosed this new species on the Dsungaripteroidea gen. et sp. indet. basis of its large size compared to other Tenda- The wing phalanx one assigned to `Pterodac- guru remains, but size is not a diagnostic charac- tylus brancai' (MB.R.2848) is complete, except 13 0 Unwin, D. M. & W.-D . Heinrich, Pterosaur Jaw of Tendaguru that the proximal tip of the proximal extensor Discussion process is missing (Reck 1931, Fig.4) . The pha- lanx has a very distinctive and marked forward The evolutionary history of dsungaripteroid ptero- curvature, an important character of dsungari- saurs: Until recently, little was known of the pteroid pterosaurs (Unwin et al. 1996), and has fossil record of dsungaripteroids and they were two further characters of this taxon: the bone represented only by Germanodactylus from the walls appear to have been relatively thick, with Upper Jurassic of Bavaria and Dsungaripterus only a narrow lumen (Unwin 1988, Bennett and Noripterus from the Early Cretaceous of 1989), and the distal articular end is markedly China (Young 1964, 1973, Wellnhofer 1970, expanded (Unwin et al. 1996). Assignment to 1978) . In the last two decades a series of impor- Dsungaripteroidea is clearly warranted. How- tant new finds have been made (Table 2) and ever, within this group the morphology of the these have considerably expanded our knowl- wing phalanx one shows little variation, other edge of this group. The most important of these than size, and the Tendaguru specimen cannot discoveries includes material from Mongolia be safely assigned to any of the known genera, (Bakhurina 1982, 1986), the recognition of dsun- nor does it exhibit any characters that enable it garipteroids in the Tendaguru pterosaur assem- to be distinguished from these taxa. For the pre- blage (Galton 1980), a new genus from France sent, therefore, we assign it to Dsungaripteroidea (Buffetaut et al. 1998) and the recognition of gen. et sp. indet. dsungaripteroids in South America (Galton 1980, In conclusion, since neither of the specimens Unwin 1996, Martill and Frey pers comm. 1999) . upon which `Pterodactylus brancai' were based This compilation of data on the fossil record exhibits any features by which this taxon may be of dsungaripteroids shows, however, that the diagnosed we consider this name, and the syno- known stratigraphic range of the clade, Upper nym `Dsungaripterus brancai', to be a nomen du- Jurassic (Kimmeridgian) through to the mid bium. However, as Galton (1980) first noted, the Lower Cretaceous (the exact age depends on the holotype material shows some similarity to the dating of the Wuerho pterosaurs which is uncer- corresponding elements of Dsungaripterus, and tain [Doug 1992]) has hardly changed since the this is discussed further below. first recognition of the group by Young in 1964. `Pterodactylus arningi' was erected on the ba- Interestingly, dsungaripteroids have not been sis of a single, almost complete, left wing pha- found in any of the main late Early or early lanx one (MB.R.2843). The anterior tip of the Late Cretaceous pterosaur localities of Europe extensor process is missing and the posterior (Cambridge Greensand and Lower Chalk of margin of the proximal end is badly eroded England), Africa (Cenomanian of Morocco), (Reck 1931, Fig. 5) . The phalanx is almost as Asia (Upper Cretaceous of Uzbekistan) or the long as MB.R.2848, but, as Reck (1931) noted, Americas (Santana and Crato Formations of the shaft has a much more slender build. In ad- Brazil, Niobrara Chalk of Kansas) . This suggests dition, the bone is not reflexed forwards and the that dsungaripteroids may have become extinct bone walls appear to be relatively much thinner sometime in the late Early Cretaceous. than in MB.R.2848. This specimen cannot there- In the Late Jurassic dsungaripteroids appear to fore be assigned to the Dsungaripteroidea. How- have been rather diverse and widely distributed, ever, it is so singularly bereft of any distinctive occurring in Laurasia (Europe and ?North Amer- features that, for the present, it cannot be as- ica) and in Gondwana (Africa and ?South Amer- signed to any taxon other than Pterosauria in- ica) . This suggests that they were already well det., and `Pterodactylus arningi' must be consid- established by this time and that the group origi- ered a nomen dubium. nated somewhat earlier in the Jurassic. Herbsto- In conclusion, none of the previously pro- saurus (Casamiquela 1975), which is late Middle posed Tendaguru taxa appear to be valid, but Jurassic (Callovian), might provide direct evidence this is not surprising since single, isolated, post- for this, but its assignment to the Dsungaripteroi- cranial elements rarely bear features sufficient dea has yet to be established (Unwin 1996) . upon which to base the diagnosis of new genera The Tendaguru pterosaur assemblage: The or species. A corollary of this conclusion is that excavations at Tendaguru yielded more than neither Pterodactylus nor Rhamphorhynchus are 200 remains of pterosaurs. Many of these con- present in the Tendaguru deposits, confirming sist of incomplete bone shafts lacking articular earlier doubts (Wellnhofer 1978) regarding the ends and can only be identified as Pterosauria taxonomic assignments made by Reek (1931). incertae sedis. A small proportion of the re- Mitt . Mus. Nat.kd. Berl ., Geowiss. Reihe 2 (1999) mains are more complete and, in many cases, P Wellnhofer, Munich, E. Frey, Karlsruhe, Z. Dong and J. Lii, can be identified with a little more precision. Beijing, and S. Etches, Kimmeridge, for giving access to and assistance with material in their care . We are especially Most of these bones appear to be pterodacty- grateful to N. Bakhurina, M. Benton, D. Martill and P Welln- loid and often compare well with corresponding hofer for their valuable comments on earlier versions of the elements of dsungaripteroids (e.g. MB.R.2842 MS. and 2848, discussed above). These remains cor- respond broadly in size to the holotype of Ten- References daguripterus recki n. gen. n. sp. and probably belong to this taxon, but, unless more complete Aitken, W. G. 1961 . Geology and Palaeontology of the Juras- material is found, this cannot be demonstrated sic and Cretaceous of Southern Tanganyika. - Bulletin of with certainty. A few of the identifiable remains the Geological Survey of Tanganyika 31: 1-144. Bakhurina, N. N. 1982 . [A pterodactyl from appear to represent `rhamphorhynchoids' the Lower Cre- (e.g. taceous of Mongolia] . - Palaeontologicheskii Zhurnal MB.R.2845), but further study is needed to es- (1982) 4: 104-108 [In Russian] . tablish their identity more precisely. - 1986. [Flying reptiles] . - Priroda (1986) 7: 27-36 [In Russian] . Thus, at a general level, the taxonomic com- Bakhurina, N. N. & Unwin, D. M. 1995. A survey of ptero- position of the Tendaguru pterosaur assemblage saurs from the Jurassic and Cretaceous of the former resembles that from other Upper Jurassic sites, Soviet Union and Mongolia . - Historical Biology 10: 197-245 . in that both `rhamphorhynchoids' and pterodac- Bennett, S. C. 1989. A pteranodontid pterosaur from the tyloids are present, and most of the individuals Early Cretaceous of Peru, with comments on the relation- appear to have been small to medium sized, the ships of Cretaceous pterosaurs. - Journal of Paleontology largest reaching 63: 669-676 . an estimated 2-2.5 m in wing- - 1994. Taxonomy and systematics of the Late Cretaceous span. However, when considered in more detail, pterosaur Pteranodon (Pterosauria, Pterodactyloidea) . - the composition of the Tendaguru assemblage Occasional Papers of the Natural History Museum . The exhibits University of Kansas, Lawrence, Kansas 169: 1-70 . some distinct differences from that of Benton, M. J., Cook, E., Grigorescu, D., Popa, E. & Tallodi, other Upper Jurassic pterosaur localities. `Rham- E. 1997. Dinosaurs and other tetrapods in an Early Cre- phorhynchoids' are exceptionally rare in the taceous bauxite-filled fissure, northwestern Romania . - Palaeogeography, Palaeoclimatology and Palaeoecology Tendaguru collections, which is in sharp contrast 130: 275-292 . to the Solnhofen Limestone, Kimmeridge Clay Bonaparte, J. F 1978. El Mesozoico de America del Sur y and Karatau assemblages, wherein `rhampho- sus Tetrapodos. - Opera Lilloana 26. Tucuman, Argen- tina . rhynchoids' represent between 50 and 100 per Bonaparte, J. E & Sanchez, T M. 1975 . Restos de un ptero- cent of the total number of identifiable indi- saurio, Puntanipterus globosus de la Formacion La Cruz, viduals (e.g. Wellnhofer 1991). Similarly, the ap- Provincia de San Luis, Argentina . - Acta primero Con- parent greso Argentino de Paleontologia y Bioestratigrafia 2: preponderance of dsungaripteroids at 105-113 . Tendaguru also contrasts with the situation in Broschinski, A. 1999. Ein Lacertilier (Scincomorpha, Para- the other Jurassic assemblages mentioned above, macellodidae) aus dem Oberen Jura von Tendaguru/Tan- sania. - Mitteilungen aus wherein dsungaripteroids are rare or dem Museum ftir Naturkunde absent (see Berlin, Geowissenschaftliche Reihe 2: 155-158 . Wellnhofer 1991). In this respect, the taxonomic Buffetaut, E., Lepage, J.-J. & Lepage, G. 1998. A new ptero- composition of Tendaguru is most closely com- dactyloid pterosaur from the Kimmeridgian of the Cap de la Heve (Normandy, France). - Geological Magazine parable with the Early Cretaceous dsungaripte- 135: 719-722. roid dominated pterosaur assemblages of Tatal, Casamiquela, R. M. 1975. Herbstosaurus pigmaeus (Coeluria, Mongolia (Bakhurina 1986) and Xinjiang, China Compsognathidae) n. gen. n. sp. del Jurasico medio del (Dong Neuqu6n (Patagonia septentrional) . Uno de los mäs pe- 1992) . The contrasts and similarities quenos dinosaurios conocidos. - Acta primero Congresso among these various Upper Jurassic and Lower Argentino Paleontologia et Bioestratigrafia 2: 87-102 . Cretaceous localities are so marked that they Casamiquela, R. M. & Diaz, G. C. 1978. La presencia de Pterodaustro Bonaparte cannot be entirely attributed to (Pterodactyloidea), del Neo- taphonomic bias- jurassico (?) de la Argentina, en Los Andes del norte de ing and may perhaps reflect ecological or palaeo- Chile. - Actas del Segundo Congresso Argentino de Pa- biogeographical differences between these ptero- leontologia y Bioestratigrafia y Primer congresso Latin- americano saur assemblages. de Paleontologia 1: 201-209. Dong, Z. 1992. Dinosaurian faunas of China. 188 pp. China Ocean Press. - 1993. The field activities of the Sino-Canadian Dinosaur Acknowledgements Project in China, 1987-1990 . - Canadian Journal of Earth Science 30: 1997-2001 . Eaton, G. E 1910. Osteology of Pteranodon. - Memoirs of We are very grateful to H.-H . Krueger (MfN) for his assist- the Connecticut Academy of Arts and Sciences 2: 1-38 . ance with the preparation of the specimen and to W Harre, Galton, P M. 1980. Avian-like tibiotarsi of pterodactyloids E. Siebert, V. Heinrich and T. Bergmann for their help with (Reptilia: Pterosauria) from the Upper Jurassic of East preparation of the figures . We thank N. Bakhurina, Bristol, Africa . - Paläontologische Zeitschrift 54: 331-342 .

132 Unwin, D. M. & W.-D . Heinrich, Pterosaur Jaw of Tendaguru

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F & G. Witherby, London . Young, C. C. 1964. On a new pterosaurian from Sinkiang, Plieninger, E 1901. Beiträge zur Kenntniss der Flugsaurier . - China. - Vertebrata Palasiatica 8: 221-255 . Palaeontographica 48: 65-88. - 1973. [Wuerho pterosaurs] . - Special Publication of the Reck, H. 1931 . Die deutschostafrikanischen Flugsaurier . - Institute of Vertebrate Palaeontology and Palaeoanthro- Centralblatt für Mineralogie, Geologie und Paläontologie pology, Academica Sinica 11: 18-34 [In Chinese]. 1931: 321-336 . Table 2: The fossil record of dsungsaripteroid pterosaurs

G Taxon Horizon and locality Age Material Author Notes z Dsungaripteroidea nx. Germanodactylidae td Germanodactylus Solnhofen Plattenkalk, Late Jurassic Five skeletons of varying Wellnhofer 1970 rhamphastinus Mornsheim Schichten, completeness (and refs therein) . G Bavaria, Germany 0 Germanodactylus cristatus Solnhofen Plattenkalk, Late Jurassic A single almost complete Wellnhofer 1970 Bavaria, Germany skeleton (and refs therein) Germanodactylus sp. Kimmeridge Clay, Kimmer- Late Jurassic Seven bone remains, Unwin 1988 idge Bay, Dorset, England some incomplete N Tendaguripterus recki n. gen. Tendaguru, Tanzania Late Jurassic Incomplete lower jaw this paper Other dsungaripteroid remains n. sp. from Tendaguru may belong to this taxon Normannognathus wellnhoferi Octeville, Seine-Maritime, Late Jurassic associated anterior portions Buffetaut et al. 1998 Haute-Normandie, France of upper and lower jaws

Dsungaripteridae `Dsungaripterus parvus' Tsagan-Tsab Svita, Tatal, Early Cretaceous Extensive series of cranial Bakhurina 1982, 1986, `Phobetor' Bakhurina, 1986 Western Mongolia and postcranial remains Bakhurina & Unwin 1995 is preoccupied Noripterus complicidens Tugulu Group, Wuerho, Early Cretaceous Jaws plus a complete left Young 1973, Sun et al. 1992 Xinjiang China. fore and hind limb Dsungaripterus weil Tugulu Group, Wuerho, Early Cretaceous Skulls and an extensive Young 1964, 1973 Xinjiang China. series ofpostcranial remains Noripterus sp. Xinminbao Formation, Early Cretaceous Remains of a small Niu 1987 Xinminbao, China pterosaur New genus and species Quebrada de la Carreta, Early Cretaceous skull fragments Martill and Frey Previously identified of dsungaripterid Cordillera de Domeyko, pers comm. 1999 as Pterodaustro by Province of Antofagasta, Casamiquela & Diaz 1978 Chile. Dsungaripteridae gen. Ookurodani Formation, Early Cretaceous wing phalanx Unwin et al. 1996 et sp. indet. Shoukawa Village, Gifu Prefecture Japan Dsungaripteridae gen. Pingfengshan locality, Late Jurassic-Early numerous fragmentary Dong 1993 et sp. indet. Wucaiwan region, Cretaceous remains Junggar Basin, Xinjiang, China w w

Table 2: Aw Continued

Taxon Horizon and locality Age Material Author Notes

Dsungaripteridae gen. Qingshan Formation, Early Cretaceous isolated bones Young 1964 et sp. indet. Laiyang, Shandong Province, China Dsungaripteridae gen. Mengyin Formation, Late Jurassic-Early wing phalanx Young 1964 et sp. indet. Mengyin, China Cretaceous

Putative dsungaripteroids

Kepodactylus insperatus Morrison Formation, Late Jurassic Fragmentary remains Harris & Carpenter 1996 The humerus (Harris and Garden Park, Colorado, of a single individual Carpenter 1996, fig. 2) is USA remarkably similar to that of dsungaripterids Puntanipterus globosus La Cruz Formation, Early Cretaceous A complete tibiotarsus Bonaparte & Sanchez 1975, The tibiotarsus is similar San Luis, Argentina Galton 1980 to that of dsungaripteroids (Galton 1980), but this assign- ment requires confirmation Herbstosaurus pigmaeus Lotena Formation, Middle Jurassic Pelvis and parts Casamiquela 1975, Similarities with dsungaripter- Arroyo Picdn Leufd, of associated femora Bonaparte, 1978, oids were noted by Unwin Province of Neuqu6n, Galton 1980, (1996): this material requires Argentina Unwin 1996 further study Dsungaripterus sp . bauxite deposits, Cornet, Early Cretaceous fragmentary jaw see refs in Benton et al. Oradea, Romania 1997 Pterodactyloidea incertae sedis Morrison Formation, Late Jurassic wing phalanx one Jensen & Ostrom 1977 The wing phalanx one is Dry Mesa quarry, recurved forwards as in Montrose County, dsungaripteroids Colorado, USA

Non dsungaripteroid b `Santandactylus spixi' Santana Formation, Early Cretaceous Incomplete forelimb Wellnhofer 1985, The carpal morphology of Araripe, Bennett 1989, 1994, this form is widely distributed Chapada do öN Unwin and Lü 1997 in pterodactyloids and cannot Brazil Gm be used to support assignment to Dsungaripteroidea m F (see Unwin & Lii 1997) 0 M

a

-S G