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Home , Batrachognathus, Cearadactylus, Coloborhynchus, Ctenochasma, Dsungaripteroidea, Eudimorphodon

Mitt. Mus. Nat.kd. Berl.. Geowiss. Reihe 3 (2000) 181-206 10. 11.2000

On the systematic and stratigraphic significance of from the Lower Yixian Formation (Jehol Group) of ,

David M. Unwinl, Junchang Lu2 & Natalie N. Bakhurina3

With 7 figures and 4 tables

Abstract

A reassessment of the systematic relationships of pterosaurs from the Lower Cretaceous Yixian Formation of Liaoning Pro- vince, China, shows that Dendrorhynchoides should be reassigned to the Anurognathidae (“”) and that Eosipterus possibly belongs within Ctenochasmatidae (). These pterosaurs formed an integral part of a di- verse community that inhabited lowland terrestrial environments in the region of northeast China in the . A new compilation of data for the Lower Cretaceous hints at a broad differentiation between pterosaurs that lived in continen- tal habitats (anurognathids, ctenochasmatoids, dsungaripteroids) and those that frequented marine environments (ornitho- cheiroids). Moreover, there is evidence of further differentiation within continental habitats, between pterosaurs living in low- land and coastal regions (anurognathids, ctenochasmatoids) and those living in more inland environments (dsungaripteroids). The temporal and geographical range extensions for high rank taxa that are implied by the Yixian pterosaurs further empha- sise the incompleteness and unevenness of the record and its unreliability for biostratigraphic zonation.

Key words: Pterosaur, Lower Cretaceous, Jehol Group, China, Anurognathidae, Ctenochasmatidae.

Zusammenfassung

Eine Neubewertung der systematischen Stellung der Flugsaurier von der unterkretazischen Yixian-Formation der Provinz Liaoning, China, zeigt, dass Dendrorhynchoides den Anurognathiden (,,Rhamphorhynchoidea“) zugeordnet werden kann und dass Eosipterus vermutlich zu den Ctenochasmatiden (Pterodactyloidea) gehort. Diese beiden Flugsaurier bilden einen inte- gralen Bestandteil einer diversen Fauna, die in der Unteren Kreide ein terrestrisches Flachland-Okosystem im Nordosten Chinas bewohnte. Fasst man die fur die Untere Kreide verfiigbaren Daten zusammen, so zeigt sich eine weitgehende Diffe- renzierung zwischen Flugsauriern, die uberwiegend in kontinentalen Okosystemen lebten (Anurognathidae, Ctenochasmatoi- dea, ) und jenen, die auch oft in marinen Bereichen auftreten (Ornithocheiroidea). Dariiber hinaus gibt es auch Hinweise auf eine Differenzierung innerhalb der kontinentalen Habitate, zwischen Pterosauriern, die sich in den Ebenen und kiistennahen Bereichen aufhielten (Anurognathidae, Ctenochasmatoidea) und den Bewohnern von mehr kiistenfernen Okosystemen (Dsungaripteroidea). Die von den Taxa der Yixian-Formation angezeigte Erweiterung der stratigraphischen und geographischen Reichweite fur Taxa hoheren Ranges unterstreichen die Unvollstandigkeit und Unausgewogenheit des Fossil- berichtes der Flugsaurier und seine Unzulanglichkeit fur biostratigraphische Zonierungen.

Schliisselwortec Pterosaurier, Unter Kreide, Jehol Gruppe, China, Anurognathidae, Ctenochasmatidae.

Introduction covered from just a handful of fossil lagerstatten scattered across a 140 million interval, ran- Pterosaurs, flying reptiles, have a rela- ging from the Late to the end of the tively modest fossil record compared to most Cretaceous (e.g. Unwin 2000). The rest of the other groups (Benton 1993). The re- pterosaur fossil record consists, largely, of iso- cord is also highly uneven: only about 30 , lated and fragmentary remains that, with the ex- from the 100 or so currently recognised (Welln- ception of jaw fragments, are difficult to identify hofer 1991a), are known from relatively com- or assign, except at very general taxonomic le- plete skeletons and most of these have been re- vels. Thus, while much progress has been made

’ Museum fur Naturkunde, Zentralinstitut der Humboldt-Universitat zu Berlin, InvalidenstraBe 43, D-10115 Berlin, Germany. Institute of Vertebrate Palaeontology and Palaeoanthropology, Academica Sinica, 142 Xi-Zhi-Men-Wai Street, P.O. Box 643, Beijing 100044, China. Department of Earth Sciences, University of Bristol, Queen’s Road, Bristol BS8 lRJ, U.K. Received April 2000, accepted May 2000 182 Unwin. D. M.. Lii J. & N. N. Bakhurina. Chinese uterosaurs recently in understanding pterosaur anatomy. lated with lavas and tuffs (see Wang et al. 1998, functional morphology and phylogeny (e.g. Pa- 1999 and refs therein). The lower part of the se- dian 1988, Wellnhofer 1991a. Unwin 1999). the quence has been assigned to the Yixian Forma- evolutionary history of the group remains com- tion while the upper part is referred to the Jiufo- paratively poorly known. tang Formation (Wang et al. 1998, 1999). Some The Lower Cretaceous pterosaur fossil record workers (Ji Qiang et al. 2000) have suggested (Fig. 1. Tab. 1) reflects the general situation. further subdivision of the Jehol Group, assigning although at a somewhat finer scale. The latter the lower part of the Yixian Formation to a se- half ( + ) of this 45 million year parate unit. the Chaomidianzi Formation, a pro- long interval contains three pterosaur lagerstst- posal that we have not adopted here. The Yixian ten, each of which has yielded a number of taxa Formation has produced an exceptionally rich and, in two cases (the and the and well preserved Early Cretaceous terrestrial Santana Formation). exceptionally well pre- biota that includes a diverse palaeoflora, gastro- served remains associated with soft tissues pods, bivalves, , conchostracans, , (Martill & Unwin 1989, Martill et al. 1990. Kell- shrimps, , frogs, , lizards, non-avian ner 1997. Martill & Frey 1998). By contrast. the , , and pterosaurs (see first half of the Lower Cretaceous. the Neoco- Wang et al. [1998, 19991, Barrett [2000] and mian (= Berriasian to ) has a much Chiappe et al. [1999] for principal references on poorer record that is dominated by fragmentary the fauna and flora of this biota). remains (Tab. 1). Until recently. relatively well There has been considerable debate concern- preserved, associated remains had only been re- ing the age of the Yixian Formation, with some ported from the Tsagaantsav Svita of Western workers supporting a Late date while Mongolia (Bakhurina 1982. 1983. 1984. 1986. others argue for an Early Cretaceous age (see 1993), the Tugulu group of . China Barrett [2000] for a detailed review of this prob- (Young 1964, 1973) and the Vectis Formation of lem). Recently, Swisher et al. (1999), have pro- the Isle-of-Wight. England (Hooky 1913). Now. vided some well founded radiometric dates of however, a new Lower Cretaceous pterosaur lo- 124.6 i 0.1 Ma for the Yixian Formation (see cality has been found in western Liaoning Pro- also Wang et al. 1999), which indicate that this vince, northeast China (Ji Shu'an et al. 1999). unit is Barremian. This result is supported by the The Jehol Group of western Liaoning Pro- presence of taxa such as psittacosaurs and an- vince consists of a thick sequence of Lower Cre- giosperms that are currently known only from taceous fluvial and lacustrine sediments. interca- the Lower Cretaceous (Xu & Wang 1998, Taylor

Europe Asia North + Australasia :ambridge Greensand, England 4nhanguera , Ziiunbayan Svita, Khuren-Dukh, Paw Paw Fm., , USA 'oolebuc Fm., Australia !onchodectes, Ornrthocheirus Mongolia Coloborhynchus Inhanguera Ornithostoma' Coloborhynchus Lagarcito Fm., San Luis, Argentina

Santana Fm., Araripe, , flrasfleodacfylus, ', , Tapelara, Crato Fm., Araripe, Brazil Tapejara, Arthurdactylus. Jectis Fm., Isle-of-Wight, England 7Arhdarchidae Ornithodesrnus' Yixian Formation Dendrorhynchoides Eosrpterus Dterodactyloidea a

lastings Beds, Sussex, England ?oloborhynchus. 7 Lonchcdectes

'urbeck Fm., England Cinathcsaurus.Platalecrhynchus Tsagaantsav Svita, Tatal, Mongolia )rnithocheiridae, Purbeckopus 3sungaripteridae

Fig. 1. Stratigraphic and geographic distribution of the main Lower Cretaceous pterosaur localities. Abbreviations: Alb, Al- bian: Apt. Aptian; Bar. Barremian: Ber. Berriasian: Fm. Formation: Hau. Hauterivian; Val. Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 3 (2000) 183

& Hickey 1996). Thus, for the present, we accept Ji Shu’an et al. (1999) appears to support this a late Neocomian (Barremian) age for the assignment, with Dendrorhynchoides occupying a Yixian Formation. position close to, though not within, a cluster of Three pterosaurs have been recovered so far individuals representing , a from the Yixian Formation (Ji & Ji 1997, 1998, Ji well known pterosaur that has Shu’an et al. 1999, Lu & Wang 2000; Tab. 1; been reported from a number of European local- Figs 2, 3). The first pterosaur to be reported, ities (Wellnhofer 1991a). Eosipterus yangi (Ji & Ji 1997), consists of an in- The assignment of Dendrorhynchoides to the complete, but partially articulated postcranial has recently been thrown skeleton (Fig. 3). Initially, this pterosaur was iden- into serious doubt, however, because it has been tified as a pterodactyloid (Ji & Ji 1997), but was reported by Friend (2000) and confirmed by one not assigned to a particular family. More recently, of us (LJ) that the supposedly long tail of this a morphometric comparison led Ji Shu’an et al. pterosaur was not originally associated with the (1999) to suggest that Eosipterus is probably sy- rest of the skeleton and consists of parts of a nonymous with Pterodactylus, a pterosaur that theropod tail that were set into the slab after its has been reported from the Late Jurassic of East initial collection. The remaining caudal vertebrae and a number of localities in Western of Dendrorhynchoides form a rather short tail, a Europe (Wellnhofer 1991a), although it can only character that does not support a close relation- be reliably identified from the Solnhofen Lime- ship with Rhamphorhynchus or other rhampho- stone of Bavaria and Late Jurassic flaggy lime- rhynchids, all of which have a long tail (Wellnhofer stones of Nusplingen in Wurttemburg (Unwin & 1978, 1991a). In addition, the morphometric ana- Heinrich 1999, Unwin unpub. data). There are, lysis suffers from the same problems as those however, two difficulties with the results of Ji noted for Eosipterus. It is a phenetic approach, Shu’an et al.3 morphometric study. First, the as already mentioned, and not only does it omit technique employed (Principal Components Ana- all Cretaceous taxa, it also excludes a number of lysis) does not distinguish between primitive and well known non-pterodactyloids including forms derived characters and is a phenetic, rather than from the (, Peteino- a phylogenetic approach (e.g. Ridley 1986). It is saurus, ), Early Jurassic (Di- possible, therefore, that the apparent similarity morphodon) and even the Late Jurassic (Anur- between Eosipterus and Pterodactylus reflects a ognathus and ). preponderance of comparisons within the analy- A third specimen from the Yixian Formation, sis that are common to many pterodacty- currently under study by one of us (LJ), consists loids (i.e. plesiomorphic within the Pterodactyloi- of a partial skeleton associated with some soft dea). Moreover, these plesiomorphies may be tissue preservation (Lu & Wang 2000). Prelimin- masking synapomorphies that unite Eosipterus ary comparisons indicate that it does not belong to with taxa other than Pterodactylus. A second prob- either of the two taxa named and described so far. lem with the analysis is that it includes only four The problems concerning the systematic rela- pterodactyloids, all from the Late Jurassic of wes- tionships of the Yixian pterosaurs also bear upon tern Europe. It is possible that Eosipterus might the debate concerning the age of the Yixian For- be more closely related to other Jurassic ptero- mation. Ji Shu’an eta]. (1999) argued in favour dactyloids, or even Cretaceous taxa (as we argue of a Late Jurassic date for this unit on the basis below), but these potential relationships are not of supposed similarities between the Liaoning tested in the study by Ji Shu’an et al. (1999). pterosaurs and Late Jurassic forms from Europe. In 1998 a second pterosaur from the Yixian Other have cited their work in support of a Late Formation, represented by a complete and fully Jurassic age (Chiappe et al. 1999), though Bar- articulated skeleton (Fig. 2), was described under rett (2000) has pointed out some difficulties with the name Dendrorhynchus by Ji and Ji, although the assumptions underlying the use of the Yixian the name was subsequently found to be preoccu- pterosaurs for dating. pied and was replaced with Dendrorhynchoides In this paper we present the results of a reas- (Ji Shu’an et al. 1999). This pterosaur was identi- sessment of the phylogenetic relationships of Eo- fied as a “rhamphorhynchoid” (Ji & Ji 1998), sipterus and Dendrorhynchoides, consider the sig- and assigned to the Rhamphorhynchidae, an im- nificance of these results for our understanding portant family that seems to have been present of pterosaur diversity and evolutionary history in throughout much of the Jurassic (Wellnhofer the Early Cretaceous and comment on the use 1991a). A morphometric analysis carried out by of pterosaurs for stratigraphic dating. The term 183 Unnin. D. M.. Lu J. & N. N. Bakhurina. Chinese uterosaurs

“Rhamphorhynchoidea”, and its derivatives. de- 1999) in the Sihetun region, south of Beipiao notes a paraphyletic taxon, which is indicated City. in western Liaoning Province, China (Ji & herein by placing the name in double quotation Ji 1998). The skeleton (Fig. 2) appears to have marks. We prefer the more neutral term non- been complete and almost fully articulated, though pterodactyloid. and restrict use of the term parts of some elements, such as the right femur “Rhamphorhynchoidea” to a historic context. for and right tibia, do not seem to be preserved on example when referring to opinions published in the main slab. The skull is somewhat disarticu- earlier works. Where there is some doubt regard- lated. possibly as a result of displacement of bones ing the validity of a taxonomic name this is during compaction of this three dimensional shown by enclosing it in single quotation marks. structure. Similarly, the bones of the pectoral except in the case of .Oriiithodesmiis‘. which is a girdle also appear to have undergone some dis- valid taxon awaiting a replacement name placement. The preservation of the skeleton, and (Howse & Milner 1993). its current disposition, is remarkably similar to that observed in specimens from other pterosaur lagerstatten, most notably Rhamphorhynchus Institutional abbreviations from the (e.g. Wellnhofer 1975a: pl. 3.5, fig. 1) and from the Kara- BSP. Bayerische Staatssammlung fur Palaontologie und his- bastau Formation (e.g. Sharov 1971: pl. 4). torische Geologic. Munich. German!: CM. Carnegie Mu- seum of Natural History. Pittsburgh. USA: FHSM. Fort Hays A general description of the skeleton, princi- State Museum (= Sternberg Memorial Museum). Fort Hays pal measurements and figures of the specimen State University. Hays. Kansas. USA: GML’. National Geolo- have been given by Ji & Ji (1998). An illustra- gical Museum of China. Beijing. China: GSM. Geological Survey Museum. Keyworth. England: GPIB. Geologisch-Pa- tion of the skeleton, indicating the principal ele- Iaontologisches Institut der Universitat Bonn. German! : ments, including some of the skull bones, is GPIH. Geologisch-Palaontologisches Institut der Universitat shown in Figure 2. Dendrorhynchoides curviden- Heidelberg. Germany: IMCF. In.aki Coal and Fossil Museum. Iwaki, Japan: IVPP. Institute of Vertebrate and tatiis was a relatively small pterosaur with an es- Paleoanthropology. Beijing. China: MB. Museum fur Natur- timated wing span of between 0.4 and 0.5 m. kunde. Berlin. Germany: MCSNB. Museo Civico di Storia A problematic aspect of GMV 2128 is the tail. Naturale. Bergamo. Italy: MFSN. Museo Friulano di Storia Naturale. Udine. Italy: MNHN. Museum National d‘Histoire The basal section, which is clearly preserved in Naturelle, Paris. France: MT. Institut und Museum fur Geolo- contact with the sacrum, consists of six, short, gic und Palaontologie der {JniLersitat Tubingen. Germany: stubby. naturally articulated caudals (Ji & Ji NSM. National Science Museum. Tokyo. Japan: PIN. Pa- laeontological Institute. Russian Academy of Sciences. Mos- 1998). The third to sixth caudals are of similar cow, Russia: PVL. Paleontologia de Vertebrados. Instituto- size. but a little shorter and a little narrower Fundacion Miguel Lillo. Universidad Nacional de Tucuman. than the first and second caudals. This section is Argentina: SMF. Natur-Museum und Forschungsinstitut Senckenberg. Frankfurt am Main. Germany: SMK. Staat- followed by a short gap, then a series of long, liches Museum fur Naturkundc. Karlsruhc. Germany: SMNS. slender caudals that are wrapped in a sheath of Staatliches Museum fur Naturkunde. Stuttgart. Germany: highly elongate zygapophyses and chevrons (Ji & TM, Teyler Museum. Harlem. Holland: TMM. Texas Mem- orial Museum. Austin. Texas. USA: UB. Katedra Geoloeie a Ji 1998: fig. 1). Recently, however, Zhou, quoted Paleontologie. University of Brunn. Czech Republic: UNSM. in Friend (2000), has suggested that the section University of Nebraska State Museum. Lincoln. Nebraska. consisting of the elongate caudals was added USA: ZMNH. Zhejiang Museum of Natural History. Hang- Lhou. Zhejiang Province. China. subsequent to the fossil discovery and does not belong to D. curvidentatus. If this is correct then, originally. D. curvidentatus had a short, slightly Systematic review tapering tail about 1.5-20 mm long and consist- ing of at least six and possibly up to eight small Pterosauria Kaup, 1834 caudals that formed a pygostyle-like structure, as “Rhamphorhynchoidea” Plieninger, 1901 found in anurognathids and pterodactyloids. Anurognathidae Kuhn, 1937 We agree with Zhou for the following reasons: Dendrorhynchoides (= Dendrorhynchus Ji & Ji, (1) in all long-tailed pterosaurs early caudals do 1998 [preoccupied]) Ji, Ji & Padian 1999 not show a decline in size caudally, but are Dendrorhynchoides curvidentatus (Ji & Ji, 1998) either of similar dimensions, or show a slight size Fig. 2 increase (Wellnhofer 1978, 1991a). Moreover, by the fifth or sixth caudal the marked elongation This taxon is represented by a single specimen typical of later vertebrae is clearly evident and (GMV 2128) collected from the Yixian Forma- the sixth caudal is usually at least twice as long tion at the locality of Zhangjiagou (Wang et al. as it is wide. By contrast, in D. curvidentatus the Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 3 (2000) 185

iii ii 186 Unwin. D. M.. Lii J. & N. N. Bakhurina, Chinese pterosaurs early caudals exhibit a decrease in size and the retains two elongate phalanges (Wellnhofer fifth and sixth elements are short and stubby. (2) 1978). An individual involved in the collection and pre- (3) Elongation of the pteroid. Pterodactyloids are paration of GMV 2128 admitted to Wang Xiao- characterised by an elongate, slender pteroid. lin (see Wang et al. 1999) that the elongate sec- Dendrorhynchoides, and other non-pterodac- tion of the tail had been added to the fossil tyloids, exhibit the primitive condition in subsequent to its discovery. As Chiappe et al. which the pteroid is a short, stubby structure. (1999) note, forgery has also been detected in a Ji & Ji (1998) cited the presence of a long tail number of other collected from the Yixian in support of their assignment of Dendrorhynch- Formation. odes to the “Rhamphorhynchoidea”, but, as dis- Dendrorhynchoides is undoubtedly a member cussed above, this character is no longer valid of the Pterosauria as it shows eight unambiguous for this taxon. The presence of a short tail in apomorphies of this taxon: Dendrorhynchoides could be used to argue in (1) coracoid elongate and at least 0.75 x length support of a relationship with the Pterodactyloi- of scapula: dea, because one of the characters that diag- (2) medial carpal-pteroid complex in the wrist: noses the Pterodactyloidea is strong reduction of (3) highly enlarged wing-metacarpal that forms the tail to a relatively short, pygostyle-like struc- part of the wing spar: ture (e.g. Bennett 1994). However, a short tail (4) hyperelongated fourth manual digit: also occurs in the Anurognathidae (Doderlein (5) forelimb at least 2 x length of hind limb: 1923. Riabinin 1948, Wellnhofer 1975a, Bakhuri- (6) metatarsals i-iv highly elongate, slender and na 1988), thus it does not provide unequivocal of subequal length: support for the assignment of Dendrorhynch- (7) pedal digit five composed of two elongate oides to the Pterodactyloidea, a hypothesis of re- phalanges: lationships that, in any case, would directly con- (8) penultimate phalanges of both manus and flict with other character distributions discussed pes elongate. earlier. Dendrorhynchoides appears to lie outside the The position of Dendrorhynchoides within Pterodactyloidea because it exhibits the primitive ”Rhamphorhynchoidea” can also be pinned condition for all three unambiguous apomor- down fairly precisely. Dendrorhynchoides exhi- phies of the Pterodactyloidea that can be scored bits two clear cut apomorphies of the An- for this pterosaur. urognathidae + Campylognathoididae + Rham- (1) Elongation of the wing (fourth) metacarpal. phorhynchidae + Pterodactyloidea (Fig. 4): All pterodactyloids exhibit the derived state (1) Forelimb length more than 2.5 x hind limb for this character wherein the wing-metacar- length. The basal pterosaurs Preondactylus pal always attains at least 0.9 x the length of and the Dimorphodontidae exhibit the primi- the humerus. Dendrorhynchoides and other tive condition in which the forelimb is less non-pterodactyloids exhibit the primitive than 2.5 x the hind limb length. state wherein the wing-metacarpal reaches (2) Ulna longer than the tibia. The basal ptero- less than 0.9 x the length of the humerus (in saurs Preondactyfus and the Dimorphodonti- Dendrorhynchoides the wing-metacarpal is dae exhibit the primitive condition in which only 0.33 x the length of the humerus). the ulna is shorter than the tibia (Tab. 2). (2) Reduction of the fifth pedal digit. Pterodac- By contrast, Dendrorhynchoides exhibits the tyloids are characterised by the derived con- primitive condition for a character that supports dition in which phalange two is lost and pha- the clade Campylognathoididae + Rhamphor- lange one is reduced to a small nubbin of hynchidae + Pterodactyloidea (Fig. 4): bone or is absent. Non-pterodactyloids. in- (1) Skull elongate with a long, low, preorbital cluding Dendrorhynchoides exhibit the primi- region: Dendrorhynchoides exhibits the pri- tive condition, in which the fifth pedal digit mitive condition with a skull that appears to

Fig. 2. Dendr.orh?.nchoides curvidemmrs (Ji & Ji. 1998). holotype (GMV 2128) in dorsal view. Elements located within the shaded rectangle do not appear to form part of the holotypc skeleton (see text for details). Abbreviations: ca, carpus; cv, cervical vertebrae: cav. caudal vertebrae: CO. coracoid: dv. dorsal vertebrae: f. femur: fi. fibula: h, humerus; i, ii, iii, iv, digits one to four: il. ilium: mc. metacarpal: mt. metatarsal: mx. maxilla: n. nasal: pal, palatine; pm, premaxilla; pt, pteroid; pu, pubis; ra. radius: ri. rib: sc. scapula: ti. tibia: ul. ulna: v. pedal digit five. Scale bar = 10 mni Table 1 Compilation of data for the body fossil record and ichnological record of Lower Cretaceous pterosaurs. Data was collected from the literature published up to March 2000. Previous taxonomic assignments were reassessed: the identifications listed here are resolved to the lowest possible taxonomic rank within the limits of current understanding of pterosaur phylogeny. Body fossil record

Locality and horizon Age Depositional Taxon Material Principal references setting

Purbeck Limestone Formation, Dorset, Berriasian marginal marine Gnathosaitrics rnacrurus mandibular symphysis Howse & Milner 1995 Eng 1 and Plataleorhynchus streptophorodon incomplete rostrum mandibular symphysis Pterodactyloidea teeth and isolated post- cranial bones Couches-Rouges, Eastern High Atlas Berriasian deltaic Pterosauria teeth Sigogneau-Russell et al. Mountains, Anoual, Morocco 1998 Cornet, Oradea, Romania Berriasian karst landscape ?Dsungaripteroidea numerous isolated Benton et al. 1997, incomplete bones Jurcsak & Popa 1983 Villar del Rio, Soria, Spain ?Berriasian Pterosauria 3 wing-finger phalanges Fuentes Vidarte & Meijide Calvo 1996 Tsagaantsav Svita, Tatal, Mongolia Berriasian- lacustrine Dsungaripteridae extensive series of cranial Bakhurina 1982, 1983, Valanginian and postcranial remains 1986, 1993 Okurodani Fm., Kobudani Valley, Berriasian- floodplain Dsungaripteridae wing-phalange Unwin et al. 1996, Shokawa, Gifu Prefecture, Japan Valanginian Evans et al. 1998 Pterodactyloidea indet. tooth Evans et al. 1998 (non Dsungaripteridae) Kuwajima Fm., Kuwajima, Shiramine, Berriasian- flood plain-swamp Ornithocheiridae tooth Hiroshige et al. in press Ishikawa Prefecture, Japan Valanginian Gnathosaurinae teeth ?Dsungaripteroidea tooth Hastings Beds, Sussex, England Valanginian fluvio-deltaic Coloborhynchus clavirostris anterior end of rostrum Owen 1874 ? Lonchodectes sagittirostris parts of left and right mandibles Pterodactyloidea 30+ isolated, incomplete Mantell 1827 limb bones Engelbostel, Hannover, Germany Hauterivian marine Ornithocheiridae fragment of mandibular Wild 1990 symphysis, fragments of wing bones Calcaire a Spatangues, Attancourt, Wassy, Hauterivian marine Pterosauria ulna Buffetaut & Wellnhofer Haute-Marne, France 1983 Corrales del Pelejon-2, Calve, Teruel, Spain Hauterivian ?Dsungaripteroidea wing-finger phalange Canudo et al. 1997 La Amarga Formation, Arroyo La Amarga, Hauterivian- coastal plain Pterodactyloidea femur Montanelli 1987 NeuquCn, Argentina Barremian Body fossil record

Locality and horizon Age Depositional Taxon Material Principal references setting

Yixian Formation, Sihetun, Barremian lacustrine complete articulated Ji & Ji 1908. Ji et al. Lieoning Province, China skeleton 1 999 Eosipterit yarigi incomplete postcranial Ji 6i Ji 1997. Ji ct al. skeleton 1 999 Pterodactyloidca genus ii incomplete postcranial skeleton Lu et al. in press Artolcs Formation, Vallipdn, Tcruel, Spain Barremian m;irginal marine Pterodactyloidca tccth Ruiz-Omenaca ct al. I998 Colladico Blanco, Galve. 'Teruel. Spain Barremian Pterosauria isolated tccth, limh hones Krchs 1985, Sanz ct ill. 1990 Ptcrocericn niarls. Scrra Tiago dos Vclhos, Barremian cervical vertebra (ialton I994 Extremadura, Portugal Vcctis Formation. Isle of Wight, England Barremian at least two incomplete Hoolcy 191.'I skeletons, with skull matcrial one incomplete skeleton and this paper various isolated, incomplete bones Crato Formation. Araripc, ('cara. IJra/il Aptian lagoon almost completc postcranial Frcy & Martill I994 skeleton skull with soft tissues Carnpos 6i Kellncr 1997 two skulls with soft tissues Martill & Frey 1998 partial forelimb Martill & Frey 1998, I999 Santana Formation, Araripe, Ccara, Brazil Aptian marginal marine 6+ skulls, some associated Wellnhofer 1985, 1991~ with postcranial remains Atihungiierti blitter.sdorffi skull Campos & Kellner 1985 'Tropeognathus ' rnesrrnhrinus skull + mandibles Wellnhofer 1987 ' Tropeognathus ' robustus incomplete mandibular Wellnhofer 1987 symphysis incomplete mandibular Kellner 1984 symphysis Tupejaru wellnhojeri 4+ skulls, some associated Kellner 1989, Wellnhofer with postcranial remains 1991b, Wellnhofer & Kellner 1991 Tupuxuara longicristatus 3 skulls, some associated Kellner & Campos 1988 with postcranial remains Tupuxuara leonardii jaw fragment Kellner & Campos 1994 Cearadactylus atrox incomplete skull + mandibles Leonardi & Borgomanero 1985 Ornithocheiridae numerous postcranial remains Wellnhofer 1985, 1991b, 1991c Table 1 (continued) Body fossil record

Locality and horizon Age Depositional Taxon Material Principal references setting

Chinshan Formation, Laiyang, Aptian floodplain ?Dsungaripteridae fragmentary limb bones Young 1958, 1964 Shandong Province, China Elrhaz Formation, Gadoufaoua, Aptian fluvial ? Azhdarchidae limb bones Sereno et al. 1998 TenerC Desert, Agadez, Niger Glen Rose Formation, Glen Rose, Aptian-Albian shallow marine ?Azhdarchidae humerus Murry et al. 1991 Somervell County, Texas Otway Group, Cove East, Australia Aptian-Albian fluvial-lacus trine Pterosauria limb bone Rich & Rich 1989 Antlers Formation, Forestburg, Aptian-Albian terrestrial ?Pterosauria bone fragments Zangerl & Denison 1950 Montague County, Texas, USA Hudspeth Formation, Mitchell Quadrangle, Albian marine ? humerus, vertebrae, Gilmore 1928 Wheeler County, Oregon, USA other bone fragment Le Gaty, Troyes, Aube, France Albian marine Ornithocheiroidea cervical vertebra Buffetaut et al. 1989 Upper Greensand, Isle of Wight, England Albian shallow marine jaw fragment Unwin unpublished data Gault, Louppy-le-Chateau, La Meuse, Albian shallow marine Pterosauria vertebra Sauvage 1882 France Toolebuc Formation, Hamilton Hotel, Albian marine Anhanguera ?cuvieri jaw fragment, scapulocoracoid. Molnar & Thulborn Queensland, Australia vertebra, pelvis 1980, Molnar 1987 Lagarcito Formation, Quebrada de Albian lacustrine Pterodaustro guinazui numerous remains including Bonaparte 1970, Sanchez Hualtarh, San Luis, Argentina some complete skeletons 1973, Chiappe et al. 199.5, with skulls 1998a, 1998b Mowry Shale, Teigen, Montana, USA Albian Ornithocheiroidea humerus Bennett 1989 Ziiiinbayan Svita, Khuren-Dukh, Mongolia Albian lacustrine Coloborhynchus skull and most Bakhurina & Unwin 199.5, of postcranial skeleton Unwin & Bakhurina in press Klaudzin Formation, Klaudzin, Fergana, Albian freshwater deposits Pterosauria distal end of femur Nesov 1990 Kirghizia Khodzhakul Formation, Khodzhakul, Albian deltaic-marginal Pterosauria bone fragments Nesov 1990 south west Kizylkum, Uzbekistan marine Tlayda , Tepexi de Rodriguez, Albian closed lagoon Ornithocheiroidea 10 isolated postcranial remains Cabral-Perdomo Puebla, Mexico & Applegate 1994 , Tarrant County Texas, Albian Coloborhynchus jaw fragment Lee 1994 Gault, Folkestone, Kenl, England Albian shallow marine Lonchodectes platysomus incomplete mandibular Unwin unpublished data symph ysis Pterodactyloidea isolated limb bones Body fossil record

Locality ii 11d horizon Age Depositional Taxon Material Principal references setting

Chenini Formation, Tataouine region, Albian fluvial Ornithocheiridae tooth Renton et al. 2000 'Tunisia Chulcc Formation. Hu;~llanca,Huanuco. '1 A 1bia n marine <)rnithocheiroidea incomplete humerus Bennett 1989 Peru Kobiaki, Tambov district. Russia A I bian- marine '? Ptcrosauria r;iinux of lower ,jaw Biwhanov & Ycrcmin C'cnomonian 1977 Seknicncv Formation, Lcbcdinsky Quarry. Albian- shallow mari 11c fragmentary limb bones Ncsov 1990 Bclgorod, Russia ('enonianian ('ambridge Grccnsand. Eiigland reworked fossils shallow niarinc fr:igmcnts of upper Owen 1859, 18hl. Scclcy of Albian age and lowcr .jaws 1869. 1870, 1x71, 1881, 1891. 1901. Hoolcy 1914 incomplctc mandibular symph ysis fragmcnts of upper and lower jaws ~'olrihor1rytri.lrrr.sctrpilo I'ragnicnts of upper and lower jaws Atrhcrrrgr~c~rir('11 viijri fragments of upper and lower jaws Anhnrrgict~rcrfillotri fragments of upper and lower jaws Ornithochcridae gcn. ct sp. iiidel numerous Iragmcntary postcranial remains l.onchocic~ctc~.sconzl~r~.s.~iroslri.s tragmcntary rostrum hnchodectes rnicrociot? fragments of upper and lower jaws Lonchodectes plutysomus fragmentary rostra Lonchodectes machaerorhynchiis incomplete mandibular s ymph ysis Lonchodectes sp. indet. 60+ fragmentary postcranial remains 'Ornithostomn sedgwicki' jaw fragment Quebrada La Carreta, Segunda Region Early estuarine-coastal Domeykodactylus ceciliae part of mandibular symphysis Casamiquela & Antofagasta, Chile Cretaceous environment and fragment of skull Chong Diaz 1978, Martill et al. 2000 Tugulu Group, Urhe, Junggar Basin, Early lacustrine-fluvial Noripterus complicidens incomplete skeleton Young 1973 Xinjiang China Cretaceous Dsiinguripterus weii numerous skulls and postcrani- Young 1964,1973 a1 elements Table 1 (continued) Body fossil record

Locality and horizon Age Depositional Taxon Material Principal references setting

Ejinhoro Formation. Muhuaxiao Village, Early fluvial Pterosauria Dong 1993 Ordos, Inner Mongolia Cretaceous Luohangdong Formation, Zhidan Group, Early terrestrial, ?aeolian Pterosauria bone fragments Brinkman &Peng 1993, Laolonghuoze, Inner Mongolia Cretaceous Russell & Dong 1993 Quebrada Monardes Formation, Early flood plain Pterosauria bone fragments Bell & Padian 1995 Atacama Region, Chile Cretaceous La Cruz Formation, Quebrada Larga, Early fluvial Puntanipterus globosus 1 vertebra and 3 limb bones Bonaparte & Sanchez Hualtaran, San Luis, Argentina Cretaceous 1975

Track fossil record Purbeck Beds, Langton Matravers, Dorset, Berriasian inter-tidal or Purbeckopus prints Ensom 1984, Wright England supra-tidal flats et al. 1997 Cameros Basin, Santa Cruz de Yanguas, Berriasian fluvial-lacustrine cf. Pteraichnus trackway Moratalla 1993, northern Spain Lockley et al. 1995 Villar del Rio, Soria, Spain ?Berriasian pterosaur prints Meijide Calvo & Fuentes Vidarte 1999 La Huergina Limestones Formation, Barremian lacustrine pterosaur tracks Fregenal-Martinez & Las Hoyas, Cuenca, Spain Moratalla 1995, Lockley et al. 1995, Moratalla et al. 1995 Potomac Group, Maryland, USA Aptian flood plain pterosaur tracks Kranz 1998 Lower Glen Rose Limestone, Seco Creek, Albian coastal ?pterosaur irregular scratches made Stricklin & Amsbury Medina County, Texas, USA by three clawed digits 1974, Langston 1974 Rio Limay Formation, Ezequiel Albian flood plain 'Pteraichnus' pterosaur tracks Calvo & Moratalla 1998, Ramos Mexia, NeuquBn, Argentina Calvo 1999 Los Cayos, Spain Early pterosaur isolated prints Moratalla 1993, Cretaceous Lockley et al. 1995 192 UnNin. D. M.. Lii J. & N. N. Bakhurina, Chinese pterosaurs

Table 2 Metric data for the DSV and principal elements of the fore and hind limbs of non-pterodactyloid pterosaurs. * = estimate, + = minimum length. Abbreviations: DSV. dorsal+sacral vertebrae: fem. femur: hum. humerus: mc, metacarpal; mt, metatar- sal: tib. tibia: ul. ulna: wph. wing-phalange.

Taxon Specimen DSV hum ul mc4 wphl wph2 wph3 wph4 fem tib mt3

P reonc1acrjIrr.s hiiffirrinii MFSN 1770 56 32 42 14.3 35.5 39 39 28 32.5 44 16

Dimorphodontidae Pereinosaiiriis zarnbellii MCSNB 3359 82 38.5 48 17 43 43 46.5 34.8 37 49 17.5 Dirnorphodon rnacronys GSM 1546 133.8:* 79 102 34" 84 93 104.6 84 76 105

Anurognathidae Anurognathits amtizoiii BSP 1922 142 46 32 45 11 58 27 39 18 Batrnchognathits volaris PIN 13 46 495+ 30.8 41 21.8 DendrorhwchoideJ ciirride/imrits GMV 2128 38*: 27.8 35.5 9.3 44.5 35.6 19" 26.7 12.1

Campylognathoididae Eudinzorphodorz rnnzii MCSNB 2888 10.3 47 65 29 80+ 41 50+ Eudimorphodon roseri,feltli MFSN 1797 40.5 55 21 64 58.2 63.2 51.5 37 54.2 21 Cnrnpy1ogrznrhoide.s lirrsiciis CM 11421 50.3 60.3 23 93.3 96 84.5 69.6 37.6 47.4 21.6

Rhamphorhynchidae Sordes pilosits PIN 258513 756 -13 1 69 1 16.5 50.5 53 53.8 33.9 36 49.6 14.8 Scaphognnrhiis crassirostris GPIB 1304 110 53.5 94 27 70 75 55 Dorygn arh us ban th ensis MB 1905.15 121.6 62.9 106.2 31.8 79 92.3 93 82.5 53.2 70 29.1 'Rharnphorhynchus lorzgicairdirs' TM 6924 47.5 16.5 26.7 10 37 31.8 28.9 32.4 12.5 15.5 9 'RharnphorhynchiLs inrerniediiis' LIB E55.1 57 21.8 37.5 13.7 53.8 49.4 43.5 47 17 22 12.8 Rhamphorhynchus mzrensrer-i SMFR4128 107 38 66 21 114.5 114 106.5 101 33 48.5 'Rharnphorhynchirs gemrningi' GPIH MYE.13 114 43 68 20 113 108.5 98.5 102 30 44 28.8 'Rhamphorhynclzirs IongicepJ' MT 136 64.5 96 35 149 45.5 64 34

have been short and with a relatively short, condition where the ulna is longer than the high preorbital region as in other basal non- wing-phalange one. pterodactyloids (see below). (2) Wing-phalange two longer than ulna. Again, In addition, Dendrorhynchoides also exhibits most non-pterodactyloids exhibit the primi- the primitive condition for a character that sup- tive condition where the ulna is longer than ports the clade Rhamphorhynchidae + Ptero- wing-phalange two. dactyloidea: However, the derived state of character 1 also (1) Metatarsal iv markedly shorter than metatar- occurs in Eudimorphodon, Carnpylognathoides sals i-iii. Dendrorhynchoides and other ba- and (Tab. 2) and the derived state sal non-pterodactyloids (Preondacphis, Di- of character 2 also occurs in Campylognathoides morphodontidae, Anurognathidae) exhibit (Tab. 2). In view of the homoplasy exhibited by the primitive condition wherein metatarsals both characters they must be considered as pro- i-iv are of almost the same length. viding only weak support for the pairing of Ji & Ji (1998) assigned Dendrorhynchoides to Dendrorhynchoides and Rhamphorhynchus, a re- the Rhamphorhynchidae, but it lacks apomor- lationship that would also require at least seven phies that define this taxon (e.g., deltopectoral reversals to the primitive character state in Den- crest of humerus with constricted base and ton- drorhynchoides. gue-like distal expansion; ulna longer than any In summary, comparison of Dendrorhynchoides other fore or hind limb element) or the Rham- with characters that support principal nodes with- phorhynchinae (e.g., tips of mandibles fused into in the Pterosauria indicates a position at node (a) long prow-like symphysis that, together with in the shown in Figure 4. This suggests large fang-like teeth. forms a grab: rear edge a possible relationship with the Anurognathidae of wing-phalanges deeply grooved). and comparison of the two taxa reveals that they Dendrorhynchoides and Rhamphorhynchiis share at least four clear cut apomorphies: share two putative apomorphies (although other (1) Presence of a very short, broad, lightly con- members of the Rhamphorhynchidae exhibit the structed skull with large cranial openings. All primitive condition for these characters): other pterosaurs, even dimorphodontids, have (1) Wing-phalange one longer than ulna. Most rclatively longer, taller and narrower skulls. non-pterodactyloids exhibit the primitive Anurognathids and Dendrorhynchoides have Mitt. Mus. Nat.kd. Bed., Geowiss. Reihe 3 (2000) 193

an extremely unusual skull in which the prin- longer than the ulna (Fig. 5), which, as out- cipal bones form a slender frame of thin, group comparison with other diapsids shows, fine bars and rods of bone surrounding large is the primitive condition. Anurognathids are cranial vacuities. Many skull bones have a unusual in that the DSV is remarkably short, highly distinctive shape quite unlike that principally because of the relatively low seen in other pterosaurs. Thus, the fused pre- number of dorsal vertebrae (12, compared to maxillae have the form of an inverted “T” 14-16 in Rhamphorhynchus) and their rela- with the left and right maxillary processes tive shortness. diverging at an angle somewhat greater than (3) Sub-symmetric, angular profile of the proxi- 90, rather than less than 45, as in other pter- mal end of the humerus in dorsal view. In osaurs. In addition, the maxilla is “L” shaped anurognathids and Dendrorhynchoides the with a very short premaxillary process, and deltopectoral and medial crests on the proxi- the palatine is reduced to a very slender “Y” mal end of the humerus are of similar size shaped bone, the stem of which contacted and shape, with a broad base and straight, the maxilla on its medial surface. These fea- angular outline. These, and the practically tures are present in Anurognath~~(Welln- fiat proximal surface of the caput humeri hofer 1975a: fig. 38) and Batrachognathus give the humerus a highly distinctive profile (Riabinin 1948: fig. 1) and are also clearly that is unlike that found in other pterosaurs, evident in Dendrorhynchoides (Fig. 2). In- where, for example, the deltopectoral crest is deed, the anurognathid skull presents a large always considerably larger than the medial complex of derived characters that are diag- crest (Wellnhofer 1978: fig. 10). The humerus nostic of this clade, though they have yet to of Dendrorhynchoides (Fig. 3) is similar to be documented in detail. that of Anurognathus (Wellnhofer 1975a: The combined length of the dorsal + sacral fig. 37) and Batrachognathus (Riabinin 1948: vertebrae (= DSV) is of almost the same fig. l), but is not quite so symmetrical, and length as the ulna (Tab. 2). In pterosaurs the deltopectoral crest is directed a little other than anurognathids and Dendror- more proximally than in other anurog- hynchoides the DSV is always substantially nathids.

Table 3 Metric data for the principal elements of the fore and hind limbs of pterodactyloid pterosaurs. * = estimate, + = minimum length. Abbreviations as for Table 2.

Taxon Specimen hum ul mc4 wphl wph2 wph3 wph4 fem tib mt3

Ornithocheiroids Arthurdactylus conandoylei SMK 1132 PAL 230 312 227 445 402 312 275 190 234 60 Anhanguera santanae NSM PV 19892 257 384 257 462” 387” 270” 225” 236 283’ 59 Bennett 1991 250 344 548 615 499 346 194 250 328 NyctosaLiruT gracilis UNSM 2/85 94 142 255* 308 245 122 85 120 38

Ctenoehasmatoids Cvcnorhamphus suevicus MT 65.5 87 108 141 115.2 85 77.5 77 122 30 Cycnorhamphus canjuersensb MNHN S5+ 155 134 94 67+ 101 138 35 Pterodactylus antiquus BSP AS I 739 31.5 47 35 48.5 44.2 37 28.5 34.7 48.3 19 Pterodactylus kochi BSP 1937 I 18 29 39 27.5 38.5 36.5 32.5 27 28.5 38.5 14.5 Pterodactylus micronyx BSP 1911 I31 25 31 40 46 28.5 19.5 17.7 29 40.5 8.7 Pterodactylus longicollunz SMNS 78 104 130 160 109 77.5 65 99 149 30 ‘Pterodactylus’ elegans BSP 1867 I1 1 15 18.2 16.5 19.5 17.8 14.3 12.8 12.3 17.6 5.6 gracile BSP 1935 I24 38.5 52.5 35 66 57 3s 55 21 Pteroriaustro guinnzui PVL 3860 80 102 78 116 112 86 69 60 90 50 Eosipterus yangi GMV 21 17 34+ 95 73 96 95 80 73 60 96 38.5

Dsungaripteroids Gerrnanoductyliis rharnphastiniis BSP 1977 XIX 1 51 76 58 75 69.5 58.5 40.4 53 80.4 25.8 GerrnunoducqJlus cristatus BSP 1892 IV 1 56 75 66 84 77.5 65.5 55 56.5 87.5 24 Noripterus cornplicidens IVPPP 64043-3 76 104 142 174 124 44.5+ 87 160 32

Azhdarchoids Tupuxuara longicristatus IMCF 1052 234 291 359 505 301 208 298 398 88.2 Zheijiangopterus linhuiensis ZMNH M1323 137 234 336 322 220 222 265 sp. TMM 42422 250 358 620 602 305 ,156 39 604 194 Unwin. D. M.. Lu J. & N. N. Bakhurina. Chinese Dterosaurs

(4) The combined length of the caudal vertebrae In addition to the apomorphies discussed is shorter than either the cervical or dorsal above, the skeletal morphology of Dendror- series. By contrast to all other non-pterodac- hynchoides is remarkably similar to that of Anur- tyloids, which have a caudal series that is ognathus and Batrachognathus in many other re- longer than either the cervical or dorsal ser- spects, including its relative dimensions (Tab. 2). ies (and often longer than the two com- The following characters are putative synapo- bined), in anurognathids, the tail is reduced morphies uniting Dendrorhynchoides and anur- to a remarkably short, pygostyle-like struc- ognathids: ture (e.g. Riabinin 1948, Doderlein 1923: (1) Short, conical, sharp-pointed, widely spaced fig. 4). This is similar to the condition in teeth with slightly recurved tips showing lit- pterodactyloids, but is presumably the result tle size variation along the jaw. Teeth with a of homoplasy since the Rhamphorhynchidae, similar morphology occur in some species of members of which have long tails, appear to Pterodactylus, but are usually more closely be more closely related to the Pterodactyloi- spaced and do not have recurved tips. dea than the Anurognathidae (Unwin 1995; (2) Wing-phalange one longer than the com- Fig. 4). Dendrorhynchoides appears to have bined length of the ulna + wing-metacarpal. a tail that is of the same shape and propor- In all other non-pterodactyloids, with the ex- tions as those of anurognathids. ception of Rhamphorhynchus and Campylog-

Table 4 Classification, to the generic level. of all pterosaur taxa referred to in the text. Tables 1-3 and in Figures 1 and 4-7. The interrogative mark indicates that there is some ambiguity regarding the position of a partieular taxon within the next higher rank taxon

Preon da ctylus - Ctenochasmatoidea unnamed plesion 1 Pterodactylidae Dimorphodontidae Pterodactylus Cycnorhamphus Peteinosaurus Ctenochasmatidae unnamed plesion 2 Ctenochasmatinae Anu rognath idae Ctenochasma Anurognathus ? Eosipterus Ba trachogna thus Pte roda us fro Dendrorhynchoides Gnathosaurinae unnamed plesion 3 Ceara da ctylus Campylognathoididae Campylogna thoides Hua n h epterus f udimorphodon Pla taleorhynchus unnamed plesion 4 ?Lonchodectidae Rhamphorhynchidae Lonchodectes Rhamphorhynchinae unnamed plesion 6 Dsungaripteroidea Rhamphorhynchus Germanodactylidae Scaphognathinae Scaphogna thus Dsungaripteridae Sordes Domeykoda ctylus Pterodactyloidea Ds unga rip terus ?' Ornithostoma' Noripterus ?'Puntanipterus' Azhdarchoidea Ornithocheiroidea Tapejaridae ? Tapeja ra ' Ornithodesmus' Tupuxuara Ornithocheiridae Azhdarchidae Anhanguera Zh ejia ngopterus A rth urda ctyius Quefzalcoa tlus Bra siie oda cfy /us Colobo rhynch us Ornithocheirus ' Tropeognathus' Pteranodon unnamed plesion 5 Ctenochasmatoidea - Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 3 (2000) 195

nathoides, wing-phalange one is shorter than cluding almost complete right fore and hind the ulna + wing-metacarpal (Tab. 2). Basal limbs, partial left fore and hind limbs, gastralia, members of the Rhamphorhynchidae and and parts of the pelvis (Fig. 3). The skeleton is, Campylognathoididae exhibit the primitive for the most part, naturally articulated, although condition for this character suggesting that the left wing appears to have undergone some the derived condition was independently at- disruption. The current disposition of the fore tained in anurognathids and the other two and hind limbs is similar to that observed, for families. example, in pterodactyloids from the Solnhofen (3) Wing-phalange two longer than the ulna. Limestone (Wellnhofer 1970). A brief descrip- This character is only known for Dendror- tion together with measurements of the principal hynchoides among anurognathids, because a limb bones and illustrations was given by Ji & Ji complete wing-phalange two is not preserved (1997), and a sketch of the skeleton is repro- in specimens of Batrachognathus and Anur- duced here (Fig. 3). GMV 2117 represents a rela- ognathus - thus while it is a putative anur- tively small pterodactyloid with a wingspan of ognathid apomorphy, more complete materi- about 1.25 m, although, as Ji Shu’an et al. (1999) al of other taxa is needed to confirm this. In note, the absence of fusion of some elements all other pterosaurs, with the exception of suggests that it had not reached adulthood. Rhamphorhynchus, the ulna is longer than The single specimen of Eosipterus is far less wing-phalange two (Tab. 2). More basal complete than that of Dendrorhynchoides and members of the Rhamphorhynchidae exhibit lacks a skull, cervical vertebrae, or well pre- the primitive condition, thus the derived served humerus, all of which are important for state is presumably homoplastic for Rham- determining the systematic relationships of pter- phorhynchus and anurognathids. osaurs (Wellnhofer 1970, 1975a, 1978, Howse In summary, Dendrorhynchoides and anurog- 1986, Padian 1984, Unwin & Lu 1997). The main nathids are united by four unique characters source of evidence regarding the relationships of (one of which is a character complex consisting Eosipterus consists of comparative lengths of the of at least three characters) and this relationship major limb bones. This of data is often of is supported by a further three putative apomor- taxonomic significance, and has frequently been phies. In addition, these taxa also appear to have used in systematic studies of pterosaurs (e.g. a very similar general morphology and propor- Wellnhofer 1970, 1975a, Wild 1978, 1984, 1993, tions (Tab. 2). By contrast, Dendrorhynchoides Padian 1980), but is not without problems. Onto- does not share any derived characters in com- genetic variation, for example, can be quite mon with other non-pterodactyloid taxa, the sole marked in some pterosaurs (Bennett 1995, 1996). exception being Rhamphorhynchus, and in this but this variable has yet to be fully incorporated case character optimisation is ambiguous. In the into morphometric analyses. face of such overwhelming evidence we have no Previous studies (Ji & Ji 1997, Ji Shu’an et al. doubt in assigning Dendrorhynchoides to the An- 1999) have consistently identified Eosipterus as a urognathidae (Tab. 4). pterodactyloid and this is supported by the pre- sence of two unambiguous apomorphies of the Pterodactyloidea: Pterodactyloidea Plieninger, 1901 (1) The wing-metacarpal is more than 1.5 x the Ctenochasmatoidea Kuhn, 1967 length of the third metatarsal (in Eosipterus ?Ctenochasmatidae Nopcsa, 1928 this ratio = 1.9). In non-pterodactyloids the Eosipterus Ji & Ji, 1997 wing-metacarpal reaches, at most, 1.3 x the Eosipterus yangi Ji & Ji, 1997 length of the third metatarsal, but is usually Fig. 3 shorter than the latter element. (2) The fifth pedal digit is highly reduced and The holotype and only known specimen of Eosi- represented by just a single, short nubbin of pterus (GMV 2117) was collected from lacustrine bone. In non-pterodactyloids the fifth toc is sediments in the upper part of the Hengdaozi composed of two elongate phalanges (see Member of the Yixian Formation at the locality above). of Tuanshangou (Wang et al. 1999), in the Sihe- Pterodactyloids can be subdivided into four I tun region, south of Beipiao City, in western major : the Ornithocheiroidea, Ctenochas- Liaoning Province, China (Ji & Ji 1997). The matoidea, Dsungaripteroidea and Azhdarchoidea specimen consists of an incomplete skeleton in- (Unwin 1995, Unwin & Lu 1997; Fig. 4). In so 196 Unnin. D. M.. Lu J. & N. N. Bakhurina, Chinese pterosaurs

I

Fig. 3. Eosipterits yaizgi Ji & Ji. 1997. holotype (GMV 21 17) in dorsal view. Abbreviations as for Figure 2. except: g. gastralia. Scale bar = 50 nini far as comparisons can be made. Eosipteriis exhi- 1997). It does, however, have a relatively long bits the primitive condition for postcranial apo- third metatarsal, a character that is typical of morphies of ornithocheiroids. dsungaripteroids some ctenochasmatoids (Fig. 6a). Moreover, in and azhdarchoids (see Unwin 1995. Unwin & Lu its general proportions Eosipterus also compares well with ctenochasmatoids, as shown by the re- sults of the morphometric analysis carried out by Preondaclylus Ji Shu’an et al. (1999), which grouped Eosipterus Dirnorphodontidae with Pterodacryhis - a basal member of the Cte- Anurognathidae ]non-pterodacty\oids nochasmatoidea. ///, The Ctenochasmatoidea comprise a series of Carnpylognathoididae Late Jurassic and Early Cretaceous taxa. Basal Rhamphorhynchidae members of the clade include Pterodactylus and Cycrzorhanzphiis, traditionally placed in the Pter- Ornithocheiroidea 1 odactylidae (e.g. Wellnhofer 1978), although this Ctenochasmatoidea taxon is probably paraphyletic as some species Pterodactyloidea Dsungaripteroidea appear to be more closely related to the Cteno- chasmatidae than to other members of the Ptero- Azhdarchoidea dactylidae. The Ctenochasmatidae, characterised by remarkably elongate jaws and large numbers Fig. 4. Hypothesis of phylogenetic relationships of the main cladcs of ptcrosaur. After Unwin (1995) and I!n\vin & Lii of teeth that form a filtering apparatus (e.g.

( 1997) WpllnhqferI. u111111, 1970). consists of two clades: the Mitt. Mus. Nat.kd. Berl.. Geowiss. Reihe 3 (2000) 197

gnathosaurines comprising Gnathosaurus, Plata- 3g0T leorhynchus, and Cearadactylus; and the ctenochasmatines, comprising Ctenochas- ma and Pterodaustro. In the absence of skull material it is difficult to resolve the relationships of Eosipterus to other ctenochasmatoids. We note, however, that Eosipterus is of very similar size and proportions to Pterodaustro (Tab. 3). Moreover, in some re- spects, such as the relative length of the femur to the ulna (Fig. 6b) or to the wing-phalange two, and the length of the third metatarsal compared

I to the wing-phalange one, or the tibia (Fig. 6a), 0 20 40 60 80 Eosipterus exhibits proportions that, among cte- femur (mm) nochasmatoids, are most similar to those of Pter- odaustro and Ctenochasma. Fig. 5. Comparison of the relative lengths of the DSV (dorsal In summary, there is some evidence to suggest + sacral vertebrae) to the ulna, plotted against femur length, for non-pterodactyloid pterosaurs. Comparing the DSV with that Eosipterus is a ctenochasmatoid and prob- other limb bones results in the same basic distribution of ably belongs within the Ctenochasmatidae rather points. Bennett (1995) and Unwin (1995) have argued that than sharing a close relationship with Pterodacty- the five species of Rhamphorhynchus indicated here repre- sent a single species. Four species do indeed cluster separa- lus. However, confirmation of this assignment tely and show little variation, other than in size, but, by con- must await the discovery of more complete ma- trast, Rhamphorhynchus longiceps has a relatively long ulna terial of Eosipterus. and plots within the main cluster of points. Abbreviations: Aa, Aniirognathus ammoni; Db, Dorygnathus banthensis; Dc, Dendrorhynchoides curvidentatus; Dm, Dimorphodon macro- nyx; Er, Eudimorphodon ranzii; Pb, Preondactylus buffarinii; Discussion Pz, Peteinosaurus zambellii; Rg, ’Rhamphorhynchus gemmin- gi’; Ri ’Rhumphorhynchus interrnedius’; RI, ’Rhamphorhyn- chus longicaudus’; Rm, Rhamphorhynchus muensteri; Rp, Palaeoecology of Lower Cretaceous pterosaurs ’Rhanzphorhynchus longiceps’; Sc, crassiro- Sp, Sordes pilosiis stris; The vast majority of pterosaur remains have been recovered from marginal marine or marine

a b

A 0 A 4A

A

ulna femur

0 210t OPg 0 I I 4 I 0 100 200 300 0 200 400 600 800 femur (mm) wing-phalange 1 (mm)

Fig. 6. Comparison of the third metatarsal to the tibia, plotted against femur length (a) and of the femur to the ulna, plotted against length of wing-phalange one (b) for 18 species of pterodactyloid pterosaurs. Symbols: triangle = ornithocheiroids. circle = ctenochasmatoids, square = dsungaripteroids, diamond = azhdarchoids. In (b) ctenochasmatines are indicated by filled circles. Note that the location of ‘Pterodactylus elegans’ is consistent with Bennett’s contention (1996) that individuals assigned to this species are juveniles of Ctenochasma elegans. Abbreviations: Ey, Eosipterus yangi, Cg, Ctenochusma gracile. Pe, ‘Ptero- dactylus elegans’, Pg, Pterodaustro guinazui. The dispersion of points in plot (b) provides some further support for the syste- matic arrangement utilised in this paper: for example, azhdarchoids have a very low ulnaifemur ratio, while ornithocheiroids and ctenochasmatines have relatively high ratios 198 LJiiaiii. D. M.. Lii J. & N. N. Bakhurina. Chinese pterosaurs deposits and, as is well known. records from con- are consistent with what is known of the co-oc- tinental sediments are rare and generally frag- curring and vertebrate fauna. This mentary (Wellnhofer 1991a. Bell & Padian 1995. supports the idea that these pterosaurs inhabited Bakhurina & Unwin 1996). The fossil assem- the Sihetun region and formed an integral blage from the Yixian Formation thus provides a part of this Lower Cretaceous terrestrial ecosys- useful source for new insights into the palaeoe- tem. rather than representing accidental occur- cology of pterosaurs in terrestrial ecosystems. rences of individuals displaced from elsewhere With regard to the specific palaeoecologies of by unusual conditions, for example, storms, vol- the Yixian pterosaurs. it seems likely that Den- canic activity or disrupted migration patterns. drorhynchoides. which has a skull morphology A new compilation of the Lower Cretaceous remarkably similar to that of other anurog- pterosaur fossil record, including the taxa from nathids, was a small, aerial insectivore. This in- Liaoning (Tab. 1) indicates that, for this interval, ference is based on the occurrence. in anurog- pterosaurs are now known from more than 20 nathids, of relatively small, widely spaced peg- localities that represent fully terrestrial settings. like teeth, which seem well suited for gripping Moreover, these localities are widely distributed and puncturing the chitinous exoskeletons of in- across three continents, and represent a range sects (Doderlein 1923. Wellnhofer 1975b). In ad- of palaeoenvironments from shallow, fresh- dition, the jaws of anurognathids had a relatively brackish water, inland lakes with low diversity large gape, that was further extended by a fringe faunas and floras (e.g. Tsagaantsav Svita, Tatal, of "hair-like" structures (Bakhurina Br Unwin western Mongolia: Bakhurina 1983, 1984, 1986), 1995), in a fashion analogous to that of some to well vegetated lowland areas with large and caprimulgiforms such as Podargii~and Crrpriniiil- diverse invertebrate and vertebrate communities giis, which also hawk for insects (Tyne & Berger (e.g. Yixian beds, Liaoning: Barrett 2000 and 1976). That potential prey items were abun- refs therein). This reinforces the idea that ptero- dantly available is demonstrated by the large saurs were a typical, if not particularly diverse, and diverse fauna that has also been re- component of Lower Cretaceous terrestrial eco- covered from the Yixian deposits (Lin 1993). It systems. Consequently, their absence from many may also be significant that the other three de- Lower Cretaceous vertebrate localities can prob- posits that have yielded anurognathids (Bakhar ably be attributed to taphonomic bias rather Formation, Bakhar. central Mongolia: Karabas- than true absence. This interpretation is sup- tau Formation. Karatau. Kazakhstan and the ported by the discovery of pterosaur tracks at Solnhofen Limestones. southern Germany) have localities such as Las Hoyas (Fregenal-Martinez also yielded large insect faunas. & Moratalla 1995, Lockley et al. 199.5) that The palaeoecology of Eosipreriis is far less cer- have yielded a diverse terrestrial fauna, but, as tain. Assuming that it was a ctenochasmatine it yet. no skeletal remains of pterosaurs (Melkn- seems likely that this pterosaur had elongate jaws dez 1995). with numerous. long fine teeth that functioned as Comparison of the Yixian pterosaurs with as- a sieve or filtering device. It has recently been semblages from other Lower Cretaceous iocal- shown that pterodactyloid pterosaurs had a rea- ities (Tab. 1) provides an important new insight sonably effective quadrupedal stance and gait into pterosaur palaeoecology during this interval. (Bennett 1997. Unwin 1997. 1999. Henderson & Similar sedimentologic/biotic associations to Unwin 1999. Unwin & Henderson 1999). thus Eo- those of the Yixian Formation are rare, but a sipterzis may be envisioned proceeding on stilt- new fossil biota. from the Kuwajima Formation like fore and hind limbs through streams and of Shiramine. Ishikawa Prefecture, Japan (Man- along the shallow margins of lakes and rivers siev- abe et al. 2000, Hiroshige et al. 2000), forms one ing for small, soft-bodied crustaceans. the aquatic possible example. The Kuwajima Formation con- larvae of insects, other and perhaps sists of fluviatile and swamp sediments of Early even fish fry. Again. suitable prey items are Cretaceous (Berriasian) age that have yielded a known from sediments of the Jehol Group: Chen diverse freshwater assemblage (Evans et al. 1998, (I 988) has described a diverse freshwater inverte- Manabe et al. 2000, Hiroshige et al. 2000). Ptero- brate fauna, and fish that lived and presumably saurs are only known from teeth, most of which reproduced in the ancient Liaoning lakes have represent a medium to large sized ctenochasma- also been reported (Jin et al. 199.5. Jin 1996). toid. though dsungaripterid and ornithocheirid We emphasise. therefore. that the putative pa- teeth have also been identified. The Lagarcito laeoecologies proposed for the Yixian pterosaurs Formation of Argentina, currently dated as Low- Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 3 (2000) 199

Fig. 7. Stratigraphic range of main pterosaur clades. Tree topology follows that shown in Figure 4. Stage length based on Geological Society of America Timescale 1999. Fossil record data compiled from literature published up to March 2000. Each column represents the known extent of a particular lineage with blank sections indicating gaps in the fossil record. The age of most records can only be resolved, at best, to the stage level, thus the entire stage is shown as represented if any records fall within the stage. Abbreviations: Alb, Albian; Ani, Anisian; Apt, Aptian; Baj, Bajocian; Bar, Barremian; Bat, Bathonian; Ber, Berriasian; Cal, Callo- vian; Car, Carnian; Cen, ; Cmp, ; Hau, Hauterivian, Het, Hettangian; Kim, ; Lad, Ladinian; Maa, ; Nor, ; Oxf, , Sin, Sinemurian; Toa, Toarcian; Tth, ; Tur, Turonian; Val, Valanginian 200 Unwin. D. M.. Lu J. 13 N. N. Bakhurina. Chinese uterosaurs er Cretaceous (Albian) is also broadly analogous blages (Fig. 1). The Yixian taxa provide the first with the Yixian Formation in that it also repre- substantial evidence of pterosaurs from the late sents a large. shallow. freshwater lake that has Neocomian. yielded fish, conchostracans. and a cteno- The occurrence of a ctenochasmatine in the chasmatine. PferorfLrtrsfro (Chiappe et al. 1995. mid Lower Cretaceous is not unexpected. The 1998a. b). clade is known to have survived until the late In sharp contrast. other continental localities Early Cretaceous (Fig. 7) because of the pre- have produced strikingly different pterosaur as- sence of Pterodazistro in the Albian Lagarcito semblages. For example, shallow. fresh-brackish Formation (Chiappe et al. 1998a). Teeth of water lake sediments in western Mongolia and ctenochasmatines have been reported from the China. that were deposited in semi-arid. or arid basal Cretaceous (Berriasian) of Japan (Hir- conditions. have produced low diversity biotas oshige et al. ZOOO), but, if correctly identified, dominated by, or in some cases consisting solely Eosiptrrus currently represents the youngest re- of a single species of dsungaripterid pterosaur. cord for this clade in Eurasia. often represented by large numbers of remains The presence of an anurognathid seems, at (Young 1964, 1973. Bakhurina 1982. 1986. 1993. first glance, to be rather surprising, particularly Dong 1992). as it indicates that non-pterodactyloids, long Pterosaurs from marginal marine or fully ma- thought to have become extinct at the end of the rine settings are different again from those found Jurassic (e.g. Padian 1998) actually persisted into in continental environments. Pterosaur assem- the Early Cretaceous. Until now, anurognathids blages from deposits such as the Santana Forma- had only been reported from the Jurassic, tion of Brazil (Wellnhofer 1991b). the Toolebuc although some cladistic analyses (e.g. Unwin Formation of Australia (Molnar & Thulborn 1995) imply that the clade first appeared in 1980, Molnar 1987) and the Paw Paw Formation the Late Triassic (Fig. 7). Anurognathids have a of Texas (Lee 1994) are dominated by. or consist very patchy fossil record. however, which is al- entirely of ornithocheirids, medium to very large most certainly directly related to their occupa- pterosaurs specialised in aerial piscivory (Welln- tion of continental habitats. A range extension of hofer 1991a). 20 million is not, therefore, so surprising, In summary, this brief analysis of the pa- especially as this is the first Neocomian locality laeoenvironmental distribution of Lower Cretac- to produce well preserved, reasonably complete, eous pterosaurs suggests that two broad palaeoe- associated remains of small pterosaurs. Com- cological distinctions can be made: first. between bining this new evidence for the fossil record of pterosaurs inhabiting marine and continental cn- anurognathids with the predicted range of the vironments, and second. within continental envi- clade based on cladistic analysis, suggests that ronments, between lowland communities such as this clade may have persisted for more than the Liaoning assemblage, and more specialised 80 million years. Moreover, the remarkable simi- inland ecosystems such as the dsungaripterid larity of Batrrrchognathus, Anurognathus and dominated assemblages of western China and Dendrorf~ynchoidessuggests that there was little Mongolia. The apparent palaeoecological and modification of anurognathid skeletal anatomy systematic diversity of Lower Cretaceous ptero- over an interval of at least 25 million years. saur is in distinct contrast to the situation in the There are a number of possible explanations latest Cretaceous (Campanian-Maastrichtian) for the apparent absence of non-pterodactyloids wherein continental and marine assemblages are from the Lower Cretaceous: dominated by just a single family of pterosaurs. Partial . The apparent extinction the Azhdarchidae. may be in some part real, in that perhaps only anurognathids persisted into the Lower Cretaceous while the other two main Late Evolutionary significance of the Yixian pterosaurs Jurassic non-pterodactyloid lineages, the rhamphorhynchines and the scaphog- The Yixian pterosaurs also contribute signifi- nathines, had become extinct by the end of cantly to our understanding of the evolutionary the Jurassic. history of pterosaurs. As mentioned above. the Taphonomic bias. Deposits such as the Soln- Lower Cretaceous fossil record is patchy. with a hofen Limestone, that yield well prcscrvcd distinct gap between the better represented basal associated remains of small and Cretaceous and late Early Cretaceous assem- that have also been intensively exploited are Mitt. Mus. Nat.kd. Berl., Geowiss. Reihe 3 (2000) 20 1

rare in the Neocomian. The Yixian Forma- bile, with few if any restrictions on their disper- tion and associated units form the first exam- sal, thus it seems unlikely that they would have ple from this interval that can be considered, been confined to refugia. in taphonomic respects and sheer numbers of fossils recovered, as comparable to the Solnhofen Limestone. That they have Pterosaurs as stratigraphic indicators yielded a non-pterodactyloid only serves to emphasise the patchiness of the pterosaur Ji & Ji (1998) and Ji Shu’an et al. (1999) argued fossil record. that the similarity of the Yixian pterosaurs to the (3) Taxonomic and systematic biases. Anurog- Late Jurassic forms Pterodactylus and Rhnm- nathids must have been present in the Ber- phorhynchus supported the idea that the Yixian riasian-Hauterivian interval, and other clades Formation is Late Jurassic in age. of non-pterodactyloids may also have existed There are two difficulties with this argument. at this time, but the majority of remains re- The first concerns the identity of the Yixian pter- covered from the Neocomian, so far, are osaurs. We have shown, above, that Dendror- highly fragmentary (Tab. 1). Few isolated hynchoides is an anurognathid pterosaur. This bones can be reliably identified as non-pter- clade is certainly known from the Late Jurassic odactyloid (exceptions are the maxilla, cau- and probably originated in the Late Triassic. dal vertebrae, wing-metacarpal and [in rham- Thus the presence of this taxon in the Yixian phorhynchines] the wing-phalanges) while Formation could be taken as evidence in favour other remains, even though they might be of a Jurassic rather than Cretaceous age for this non-pterodactyloid, may have been assigned unit. By contrast, Eosipterus probably belongs to to the Pterodactyloidea on the grounds that the Ctenochasmatoidea (a clade that is known to non-pterodactyloids are “known” to have have existed in the Late Jurassic and Early Cre- become extinct at the end of the Jurassic. taceous), and appears to be most closely related Thus fossil remains of non-pterodactyloids to Pterodaustro, a form that is currently known may already be present in museum collec- only from the Albian (Chiappe 1998a). Thus, if tions, but remain unidentified and perhaps pterosaurs were to be employed for dating pur- unidentifiable. poses, the Yixian taxa could be used to support Luo (1999) has suggested, on the basis of the either a Late Jurassic or an Early Cretaceous occurrence in the Yixian Formation of “rham- age. phorhynchoid” pterosaurs, compsognathid thero- The second difficulty is more general and pod dinosaurs and primitive mammals, that, in more serious. Fossils selected as candidates for the Early Cretaceous, eastern Asia was a refu- dating purposes must satisfy three basic criteria. gium for typically Late Jurassic taxa. This idea They should be: (1) distinct and easily recognisa- has been challenged by Manabe etal. (2000) ble; (2) common, and with a wide horizontal dis- who argue that new fossil finds suggest that the tribution; and (3) have a clear, well defined and historical biogeography of this region may have short vertical range (Clarkson 1986). Pterosaurs been much more complex than previously do not satisfy any of these criteria. Unless criti- thought. cal parts of the skeletal anatomy are preserved Three aspects of the Yixian pterosaur assem- (specifically the skull, or at least a substantial blage also contradict the idea of a refugium. portion of the jaws and dentition) it is difficult to First, as noted above, pterosaurs have a patchy assign specimens to particular genera or species. and highly biased fossil record. In their case, ar- The Yixian pterosaurs provide a good example guments for a refugium depend upon the ab- of this - the skull of Dendrorhynchoides clearly sence of fossils, rather than a dense, continuous, demonstrates that it is an anurognathid, while and widely distributed sequence of fossils that the headless remains of Eosipterus are difficult show, for example, that in the Early Cretaceous to assign, even at the superfamily level. The relict forms similar to those from the Late Juras- pterosaur fossil record is dominated by fragmen- sic are restricted to eastern Asia. Second, while tary remains few of which can be assigned to a Dendrorhynchoides could be argued to be a re- particular taxon. Moreover, fossil remains of lict, Eosipterus could not, because other cteno- pterosaurs are rare and, with a few exceptions, chasmatoids are known from the Lower Cretac- most genera and species are known only from a eous and have also been reported from outside single locality (cf. Wellnhofer 1978, 1991a). At Asia (Tab. 1). Third, pterosaurs were highly mo- present, it is doubtful whether a reliable strati- 202 Unwin, D. M.. Lu J. & N. N. Bakhurina, Chinese pterosaurs graphic range can be specified for any clade. ir- Bennett. S. C. 1989. A pteranodontid pterosaur from the respective of its rank. The only conclusion that Early Cretaceous of Peru, with comments on the relation- ships of Cretaceous pterosaurs. - Journal of Paleontology can be drawn from these observations is that 63: 669-677. pterosaurs are highly unreliable as indicators of - 19Y4. and systematics of the stratigraphic age and it seems unlikely that this pterosaur Pferanodon (Pterosauria, Pterodactyloidea). - Occasional Papers of the Natural History Museum, The situation will improve in the near future. University of Kansas Lawrence, Kansas 169: 1-70. - 1995. A statistical study of Rhanzphorhynchus from the

Solnhofen Limestone of Germany ~ year-classes of a sin- gle large species. - Journal of Paleontology 69: 569-580. Acknowledgements - 1YY6. Year-classes of pterosaurs from the Solnhofen Limcstonc of Germany: taxonomic and systematic impli- We are very grateful to Jeff Hecht for providing aii original cations. - Journal of Vertebrate Paleontology 16: 432-444. copy of the USA Today article. Oliver Rauhut for help with - 1997. Terrestrial locomotion of pterosaurs: a reconstruc- translations and to Elke Siebert for assistance with the fig- tion based on Preraichniis trackways. - Journal of Verte- ures. Dave Martill kindly provided information on the age of brate Paleontology 17: 104-113. material from the Isle of Wight. Work by LJ was supported Benton. M. J. 1993. Reptiles. In Benton, M. J. (ed.). The Fos- by the Chinese Academy of Sciences and the National sil Record 2: 681-715. Chapman and Hall, London. Science Foundation. DMU thanks the Royal Society (UK) Benton. M. J.. Bouaziz. S., Buffetaut, E., Martill, D., Ouaja, for supporting early stages of the work reported here and M.. Soussi. M. & Trueman, C. 2000. Dinosaurs and other the staff of the Institut fur Paliontologie. Museum fur Natur- fossil vertebrates from fluvial deposits in the Lower Cre- kunde. Berlin for all their help and assistance. We are very taceous of southern Tunisia. - Palaeogeography, Palaeo- grateful to Paul Barrett. Chris Bennett. Mike Benton. Don climatology. Palaeoecology 157: 227-246. Henderson. Dave Martill and Peter Wellnhofer for their con- Benton. M. J., Cook. E., Grigorescu, D., Popa, E. & Tallodi, structive criticism of earlier versions of this paper and to E. 1997. Dinosaurs and other tetrapods in an Early Cre- Gloria Arratia for her editorial work on the MS. taceous bauxite-filled fissure, northwestern Romania. - Palaeogeography. Palaeoclimatology, Palaeoecology 130: 275-292. Bonaparte, J. F. 1970. Pterodairstro guinazui gen. et sp. nov. 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