BANTIM ET AL.212 · Zootaxa 3869 (3) © 2014 Magnolia Press

Kellner (2003), these two clades include very fragmentary material, which became a "mixed bag" containing

several species mainly from Cambridge Greensand, some of which may belong to different groups of pterosaurs

(Kellner 1989a; Rodrigues and Kellner 2008). The results of our phylogenetic analysis were similar to those of

Kellner (2003) and Wang et al. (2009), as Ornithocheirus compressirostris is closely related to the Anhangueridae,

being its sister taxon.

Comparing our results with existing phylogenies for pterosaurs, we observed that few changes occurred for the

topology of the Anhangueridae clade. The phylogeny published by Andres and Ji (2008) positioned Tropeognathus

mesembrinus as the sister taxon to Anhanguera, which is the same result returned by Wang et al. (2009). This is

due to the presence of sagittal crests in the premaxilla and dentary (characters 15 and 46 respectively); this

character has been regarded as one of the main factors in grouping the individuals belonging to Anhangueridae

(Kellner 2003; Wang et al. 2009). In the Andres and Ji (2008) phylogeny "Lonchodectes" compressirostris Seeley,

1870 is reovered as sister taxon of Anhangueridae, and Liaoningopterus gui Wang and Zhou, 2003 is included in

the Anhanguera group, because of the possession of a sagittal crest on the premaxilla, but not on the dentary.

In the phylogeny of Lü et al. (2010) Ornithocheirus was positioned as the sister taxon to Coloborhynchus +

Anhanguera. Earlier versions of this matrix considered distinct groups of individuals as belonging to the same

clade (Unwin 2001; Unwin 2003; Lü et al. 2008.). Tropeognathus mesembrinus and Ornithocheirus

compressirostris are named only as Ornithocheirus. Species of the genus Anhanguera are all treated as

Coloborhynchus, producing difficulties in defining the phylogeny that unites several distinct taxonomic units as an

OTU (Fastnacht 2001; Unwin 2001; Unwin 2003; Lü et al. 2008; Lü et al. 2009) and therefore differs from the

present analysis, which has Ornithocheirus compressirostris as sister group of Anhangueridae.

FIGURE 7. Reconstruction of Maaradactylus kellneri.

Conclusion

Comparisons with other species and phylogenetic analysis support Maaradactylus kellneri as a new genus and a

new species (Fig. 7). Comparisons of M. kellneri with pterosaur skulls from the Araripe Basin shows that this new

taxon is different from all previously described forms from the region. The main diagnostic features of this new

taxon are related to the dentition, including the possession of groups of three teeth starting from the 18th

pair of

alveoli. This feature was never observed before in pterosaurs. A phylogenetic analysis placed Tropeognathus

mesembrinus and Maaradactylus kellneri in a polytomy, as sister taxa of the genus Anhanguera within

Pteranodontoidea (sensu Kellner 2003). This new discovery and its implications for understanding pterosaur