Anhanguera santanae, copyright Tim Evanson.

Belongs within: Pterodactyloidea.

The Anhangueria are a clade of pterosaurs including all species more closely related to Anhanguera blittersdorffi than Istiodactylus latidens or Cimoliopterus cuvieri. Its members are united by an anterior expansion of the premaxillary tip with the jaw end tall, and larger teeth at the tip of the rostrum (Rodrigues & Kellner 2013).

    |  i. s.: Camposipterus Rodrigues & Kellner 2013RK13
    |           |--*C. nasutus (Seeley 1870) [=Ornithocheirus nasutus, Ptenodactylus nasutus Seeley 1869 (n. n.)]RK13
    |           |--C. colorhinus (Seeley 1870) [=Ornithocheirus colorhinus, Ptenodactylus colorhinus Seeley 1869 (n. n.)]RK13
    |           `--C. sedgwickii (Owen 1859) (see below for synonymy)RK13
    |--+--‘Cearadactylus’ ligabuei Dalla Vecchia 1993RK13
    |  `--Ludodactylus Frey, Martill & Buchy 2003RK13, FMB03
    |       `--*L. sibbicki Frey, Martill & Buchy 2003FMB03
    `--+--Cearadactylus Leonardi & Borgomanero 1985RK13, U03 [Cearadactylidae]
       |    `--C. atrox Leonardi & Borgomanero 1985B93
            |  i. s.: ‘Ornithocheirus’ capito Seeley 1870 (see below for synonymy)RK13
            |--Caulkicephalus trimicrodonRK13
            |--Uktenadactylus wadleighiRK13
            |--Liaoningopterus Wang & Zhou 2003RK13, A08
            |    `--L. guiRK13
            |--Tropeognathus Wellnhofer 1987RK13, W91
            |    `--T. mesembrinus Wellnhofer 1987 [=Ornithocheirus mesembrinus, Criorhynchus mesembrinus]RK13
            |--Siroccopteryx Mader & Kellner 1999RK13, FMB03
            |    `--*S. moroccensis Mader & Kellner 1999 [=Coloborhynchus moroccensis]FMB03
            |--Brasileodactylus Kellner 1984FMB03
            |    `--B. araripensis Kellner 1984RK13
            |--Coloborhynchus Owen 1874RK13
            |    |--*C. clavirostris Owen 1874 [=Criorhynchus clavirostris]RK13
            |    |--C. gui Wang & Zhou 2003A08
            |    `--C. wadleighi Lee 1994A08
            `--Anhanguera Campos & Kellner 1985RK13
                 |  i. s.: *A. blittersdorffi Campos & Kellner 1985FMB03 [=Coloborhynchus blittersdorfiA08]
                 |         A. araripensis (Wellnhofer 1985)RK13, FMB03 (see below for synonymy)
                 |         A. robustus (Wellnhofer 1987)RK13, FMB03 [=Tropeognathus robustusFMB03, Coloborhynchus robustusFMB03]
                 |--A. spielbergi (Veldmeijer 2003)RK13, A08 [=Coloborhynchus spielbergiA08]
                 `--+--A. piscator Kellner & Tomida 2001RK13, FMB03 [=Coloborhynchus piscatorFMB03]
                    `--A. santanae (Wellnhofer 1985)RK13, FMB03 [=Araripesaurus santanaeFMB03, Coloborhynchus santanaeA08]

Anhanguera araripensis (Wellnhofer 1985)RK13, FMB03 [=Santanadactylus araripensisFMB03, Coloborhynchus araripensisA08]

Camposipterus sedgwickii (Owen 1859) [=Pterodactylus sedgwickii, Coloborhynchus sedgwickii, Ornithocheirus sedgwicki, Ornithochirus (l. c.) sedgwicki, Ptenodactylus sedgwicki]RK13

Ornithocheirus’ capito Seeley 1870 [=Coloborhynchus capito, Criorhynchus capito, Ptenodactylus capito Seeley 1869 (n. n.); incl. Ornithocheirus reedi Seeley 1870, Criorhynchus reedi, Cr. reedii, Ornithocheirus readi (l. c.), O. reedii]RK13

*Type species of generic name indicated


[A08] Averianov, A. O. 2008. Superorder Pterosauria. In: Ivakhnenko, M. F., & E. N. Kurochkin (eds) Fossil Vertebrates of Russia and Adjacent Territories. Fossil Reptiles and Birds. Part 1 pp. 319–342. GEOS: Moscow.

[B93] Benton, M. J. 1993. Reptilia. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 681–715. Chapman & Hall: London.

[FMB03] Frey, E., D. M. Martill & M.-C. Buchy. 2003. A new crested ornithocheirid from the Lower Cretaceous of northeastern Brazil and the unusual death of an unusual pterosaur. In: Buffetaut, E., & J.-M. Mazin (eds) Evolution and Palaeobiology of Pterosaurs. Geological Society Special Publications 217: 55–63. The Geological Society: London.

[RK13] Rodrigues, T., & A. W. A. Kellner. 2013. Taxonomic review of the Ornithocheirus complex (Pterosauria) from the Cretaceous of England. ZooKeys 308: 1–112.

[U03] Unwin, D. M. 2003. On the phylogeny and evolutionary history of pterosaurs. In: Buffetaut, E., & J.-M. Mazin (eds) Evolution and Palaeobiology of Pterosaurs. Geological Society Special Publications 217: 139–190. The Geological Society: London.

[W91] Wellnhofer, P. 1991. The Illustrated Encyclopedia of Pterosaurs. Salamander Books: London (reprinted in: Norman, D., & P. Wellnhofer. 2000 The Illustrated Encyclopedia of Dinosaurs. Salamander Books).

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