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On the Systematic and Stratigraphic Significance of Pterosaurs from the Lower Cretaceous Yixian Formation (Jehol Group) of Liaoning, China

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On the Systematic and Stratigraphic Significance of Pterosaurs from the Lower Cretaceous Yixian Formation (Jehol Group) of Liaoning, China Mitt. Mus. Nat.kd. Berl.. Geowiss. Reihe 3 (2000) 181-206 10. 11.2000 On the systematic and stratigraphic significance of pterosaurs from the Lower Cretaceous Yixian Formation (Jehol Group) of Liaoning, China 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 (“Rhamphorhynchoidea”) and that Eosipterus possibly belongs within Ctenochasmatidae (Pterodactyloidea). These pterosaurs formed an integral part of a di- verse community that inhabited lowland terrestrial environments in the region of northeast China in the Early Cretaceous. 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 pterosaur fossil 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, Dsungaripteroidea) 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 year interval, ran- Pterosaurs, Mesozoic flying reptiles, have a rela- ging from the Late Triassic to the end of the tively modest fossil record compared to most Cretaceous (e.g. Unwin 2000). The rest of the other vertebrate groups (Benton 1993). The re- pterosaur fossil record consists, largely, of iso- cord is also highly uneven: only about 30 species, 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 (Aptian + Albian) 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 Crato Formation 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, ostracods, conchostracans, insects, (Martill & Unwin 1989, Martill et al. 1990. Kell- shrimps, fishes, frogs, turtles, lizards, non-avian ner 1997. Martill & Frey 1998). By contrast. the dinosaurs, birds, mammals and pterosaurs (see first half of the Lower Cretaceous. the Neoco- Wang et al. [1998, 19991, Barrett [2000] and mian (= Berriasian to Barremian) 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 Jurassic date while Mongolia (Bakhurina 1982. 1983. 1984. 1986. others argue for an Early Cretaceous age (see 1993), the Tugulu group of Xinjiang. 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 + South America Australasia :ambridge Greensand, England 4nhanguera Coloborhynchus, Ziiunbayan Svita, Khuren-Dukh, Paw Paw Fm., Texas, USA 'oolebuc Fm., Australia !onchodectes, Ornrthocheirus Mongolia Coloborhynchus Inhanguera Ornithostoma' Coloborhynchus Lagarcito Fm., San Luis, Argentina Pterodaustro Santana Fm., Araripe, Brazil Anhanguera, flrasfleodacfylus, Tropeognathus', Cearadactylus, Tapelara, Tupuxuara Crato Fm., Araripe, Brazil Tapejara, Arthurdactylus. Jectis Fm., Isle-of-Wight, England 7Arhdarchidae Ornithodesrnus' Yixian Formation Dendrorhynchoides Eosrpterus Dterodactyloidea genus a lastings Beds, Sussex, England ?oloborhynchus. 7 Lonchcdectes 'urbeck Limestone 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. Valanginian 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 Rhamphorhynchus, a from the Yixian Formation (Ji & Ji 1997, 1998, Ji well known Late Jurassic 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 Rhamphorhynchidae 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
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