The first ornithocheiromorph humerus from Wuerho (Urho), China, with a new isotopic age of the Tugulu Group

SONG, JUNYI; ZHONG, YUTING; JIANG, SHUNXING; WANG, XIAOLIN

doi:10.1590/0001-3765202520240557

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PALEONTOLOGY • An. Acad. Bras. Ciênc. 97 (suppl 1) • 2025 • https://doi.org/10.1590/0001-3765202520240557 linkcopy

The first ornithocheiromorph humerus from Wuerho (Urho), China, with a new isotopic age of the Tugulu Group

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Pterosaur remains are rare from the lowermost Cretaceous, hampering our understanding of the taxonomic and morphological diversities of pterosaurs during this period. The Lower Cretaceous Tugulu Group in Wuerho, China is renowned for hosting the Wuerho Pterosaurian Fauna (WPF), which has so far yielded numerous fossil remains of two dsungaripterid pterosaurs, Dsungaripterus weii and Noripterus complicidens. Here we report a partial ornithocheiromorph humerus from the WPF, representing a deeply divergent clade from Dsungaripteridae. The scarcity of ornithocheiromorphs from the WPF might be interpreted by niche partitioning with dsungaripterids. Meanwhile, we also report a U-Pb zircon age of 134.27 ± 0.36 Ma dated by LA-ICP-MS for the tuffaceous layer at the uppermost part of the Shengjinkou Formation, confirming the Valanginian age of the WPF presented by a previous study. The Wuerho region is one of the few localities producing abundant pterosaur fossils and the only one with an earliest Cretaceous age. The new finding here also suggests that Ornithocheiromorpha had rapidly diversified and achieved a global distribution during the earliest Cretaceous, presumably through a series of modifications on the locomotor apparatus including the warped deltopectoral crest of the humerus, which might substantially improve their flight efficiency.

Key words
ornithocheiromorpha; Tugulu Group; Valanginian; Wuerho (Urho); China

	IVPP V 33251
Length	158*
Proximal articular facet, a-p length	31.08
Proximal articular facet, d-v depth	13.16
Deltopectoral crest, a-p length	33.02
Deltopectoral crest, p-d length	34.07
Distal end, a-p length	39*

Occurrence	Age	Taxonomy	Reference	Remarks
Matute Formation, Spain	Tithonian–Berriasian	Pteranodontoidea indet. Pterodactyloidea indet.	Puértolas-Pascual et al. (2023)	–
Purbeck Group, England	Berriasian	Ctenochasmatidae Gnathosaurus macrurus Plataleorhynchus streptorophodon Ctenochasmatidae indet. Ornithocheiromorpha Ornithocheiriformes indet. Azhdarchoidea ?Azhdarchidae indet. Pterosauria indet. (‘Doratorhynchus validus’)	Howse & Milner (1995), Martill et al. (2013)	“Ornithocheirus species A” (NHMUK PV R 11958) was originally referred to Ornithocheiridae Seeley, 1870 by Howse & Milner (1995), the extent of which is subequal to Ornithocheiriformes Andres 2021.
Münder Formation, Germany	Berriasian	Ctenochasmatidae Ctenochasma roemeri	Bennett (2007), Abel et al. (2021)	–
Cornet bauxite, Romania	Berriasian	?Dsungaripteridae indet. (or Azhdarchoidea indet.) ?Azhdarchidae indet. Pterodactyloidea indet.	Dyke et al. (2011), Averianov (2014)	The pterosaur remains from Cornet bauxite have highly controversial affinities. Dyke et al. (2011) assigned one rostrum to Dsungaripteridae based on its edentulous tip. Averianov (2014) interpreted the purported alveoli behind the edentulous tip as neurovascular foramina, and accordingly referred it to Azhdarchoidea. Besides, the midline groove in this specimen is not commonly seen on the occlusal surface of the rostrum but the mandibular symphysis. Thus the phylogenetic position of this specimen needs to be revised.
Hastings Group, England	Berriasian–Valanginian	Ornithocheiromorpha ?Lonchodectidae Serradraco sagittirostris cf. Lonchodraco (or S. sagittirostris) ?Istiodactylidae indet. Anhangueridae Coloborhynchus clavirostris ?Azhdarchoidea indet. (‘Palaeornis cliftii’)	Witton et al. (2009), Rigal et al. (2018), Sweetman (2023)	Rigal et al. (2018) placed Serradraco sagittirostris in Lonchodectidae, but this was rebutted by Averianov (2020), who argued that Serradraco did not preserve diagnostic features of Lonchodectidae. The latter hypothesis was proved by the cladistic analysis of Andres (2021). ‘Palaeornis cliftii’ (NHMUK PV R 2253) was referred to Lonchodectidae by Witton et al. (2009) and considered to be distinct from Ornithocheiroidea sensu Unwin 2003 (which is roughly equal to Pteranodontoidea sensu Kellner 2003). Lonchodectidae has a controversial phylogenetic position but recently it was generally resolved as a subclade within Ornithocheiromorpha, which is in turn a deeply diverging lineage of Pteranodontoidea sensu Kellner 2003. Averianov (2012) instead considered NHMUK PV R 2253 to be an azhdarchoid. These authors reached the consensus that this specimen did not belong with Pteranodontoidea.
Mengyin Formation, China	Berriasian–Valanginian	Ornithocheiromorpha idet.	Young (1964), Xu & Li (2015)	Young (1964) indicated that a first wing phalanx from Mengyin Formation pertained to Dsungaripterus. However, the conspicuous proximal groove of the extensor tendon process, and the posteroventral pneumatic foramen, suggest its affinity with Ornithocheiromorpha instead of Dsungaripteridae.
Stadthagen Formation, Germany	Early Valanginian	Ornithocheiromorpha Anhangueria indet.	Abel et al. (2021)	–
Tugulu Group, Wuerho, China	Valanginian	Dsungaripteridae Dsungaripterus weii Noripterus complicidens Ornithocheiromorpha indet.	Young (1964, 1973), this article	–
Rosablanca Formation, Columbia	Late Valanginian	Ornithocheiromorpha Ornithocheiriformes indet. Pterodactyloidea indet.	Cadena et al. (2020)	–
Sao Khua Formation, Thailand	Late Valanginian– early Hauterivian	Ornithocheiromorpha Ornithocheiriformes indet.	Buffetaut (2003), Tucker et al. (2022)	–

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[tfnI] *Estimated value