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Description
Anhanguera is one of the best-known pterosaurs from the Santana Group, and like a lot of its brethren in Brazil, the anhanguerids, all fossils assigned to the genus stem from the early Albian Romualdo Formation (113-110 mya). Multiple specimens, including many well-preserved skulls, have been attributed to Anhanguera, and several species have been erected within or subsequently sunk into the genus , six in total: A. blittersdorffi, A. santanae, A. araripensis (1985), A. robustus (1987), A. piscator (2000), and A. spielbergi (2003). However, the validity of most of these species has been questioned in recent years, for a variety of reasons.

Anhanguera is the type genus of the anhanguerids, the group that would traditionally have been called ornithocheirids, and in a weird way, it's quite fitting, since a lot of the anatomical features that were used to erect the type species, Anhanguera blittersdorffi , are (according to more recent studies) plesiomorphic traits widely distributed across the Anhangueridae family, making it hard to pinpoint which other species meet the criteria of belonging in this genus, outside of A. blittersdorffi itself, who is known from a complete holotype skull (nearly 50 cm long) and a second skull with a jawbone. Not helping matters is that other species of Anhanguera were originally named based on incomplete material but later, more complete specimens were (rather arbitrarily in hindsight) assigned to some of these species, muddying things.

Those are Anhanguera santanae, originally named as a species of Araripesaurus, and Anhanguera araripensis, originally named as a species of Santanadactylus, before both got reclassified by Alexander Kellner in 1990. Araripesaurus (1971) and Santanadactylus (1980) both predate Anhanguera but were both named from fragmentary postcranial material, making their validity dubious today, with the type species of Santanadactylus (S. brasilensis) even being suggested to be a chimera.
A. santanae was originally described based on the back half of a skull and bits of wing bone , but a much more complete skull (45 cm) and postcranial skeleton (AMNH 22555) were later assigned to it, and this specimen became the go-to anatomical reference for Anhanguera for many years. A. araripensis was named based on an incomplete skull (just the midsection) but later, two very complete skulls (MN 4735-V and SAO 16494) were assigned to the species, with MN 4735-V having a taller crest than most other specimens. There is also Tropeognathus robustus, described from a jawbone (the same year as T. mesembrinus) and later the tip of a snout, and was later renamed as Anhanguera robustus in 2013. However, since all three of these species were described based on very incomplete and thus not particularly diagnostic material, a 2017 study found them all to be nomen dubia, while other, more complete specimens like AMNH 22555 and MN 4735-V are relegated to Anhanguera sp., since many of the subtle differences between various skull specimens once used to distinguish them on a specific level could simply represent ontogenetic stages and/or sexual dimorphism, or even just individual variation (mirroring the taxonomic history of Pteranodon).

Two other, more recently described species with a more robust foundation are Anhanguera piscator (2000) and Anhanguera spielbergi (2003), who happen to be the largest alleged species of Anhanguera, both having a skull length of circa 65 cm respectively, and both have holotypes that are a largely complete skeleton, giving us a clear idea of their anatomy, which is why they are still considered distinct from A. blittersdorffi. A. spielbergi was originally named as Coloborhynchus spielbergi by André Veldmeijer, who also insisted that A. piscator and A. araripensis should be called Coloborhynchus piscator and Coloborhynchus araripensis. As I've talked about before, Coloborhynchus (just like Ornithocheirus) is an ill-defined wastebin taxon, and its type species (C. clavirostris) is a very small species described from fragmentary remains from Valanginian strata in England, much older than the various large, Mid Cretaceous fossils from across the world (often just snout tips) later attributed to the genus by some workers, but are nowadays classed as Uktenadactylus (Texa, dubiously England ), Nicorhynchus (England, Morroco), and Siroccopteryx (Morocco), as well as the aforementioned (much more comple) fossls from Brazil, though in 2019, A. spielbergi was reclassified as Maaradactylus spielbergi, the type species being the slightly larger M. kellneri (2014), so...yeah, the controversy is far from settled.

Despite the many species that have been named over the years, A. blittersdorffi and A. piscator might turn out to be the only valid species of Anhanguera, with others being ill-defined nomen dubia and various minor differences in the better-preserved specimens possibly being the result of individual variation, sexual dimorphism and/or growth stages, though of course, future studies will likely shake things up further and continue this controversy, and a larger sample size (which is likely going to turn up at Romualdo sooner or latter) would also help validate/dispute the former suggestions, as would more thorough studies on additional Anhanguera fossils that are still only known from preliminary descriptions or none. That said, it still seems plausible that many of the fossils found so far do indeed belong to the genus (just not as many species as once suggested) and Anhanguera seems to have outnumbered the giant Tropeognathus by a considerable margin on the ancient coasts of South America 110 million years ago, which makes sense for a smaller taxon.

Oh, and you'll occasionally see references to Anhanguera having survived into the Cenomanian, but there is no real evidence for it, as this was based on fragmentary pterosaur fossils from the upper Albian-Cenomanian of England, specifically Cambridge Greensand and the Chalk Group, being lumped into Anhanguera, including "Ornithocheirus fittoni" and Cimoliopterus cuvieri being identified as "Anhanguera fittoni" and "Anhanguera cuvieri" by some workers, but Cimoliopterus is now recognized as a distinct genus, belonging to the newly erected cimoliopterid targaryendraconians, distant cousins to the anhanguerid anhanguerians, and any other fossils are far too fragmentary to be confidently assigned to Anhanguera.

Description
Anhanguera is one of the best-known pterosaurs from the Santana Group, and like a lot of its brethren in Brazil, the anhanguerids, all fossils assigned to the genus stem from the early Albian Romualdo Formation (113-110 mya). Multiple specimens, including many well-preserved skulls, have been attributed to Anhanguera, and several species have been erected within or subsequently sunk into the genus , six in total: A. blittersdorffi, A. santanae, A. araripensis (1985), A. robustus (1987), A. piscator (2000), and A. spielbergi (2003). However, the validity of most of these species has been questioned in recent years, for a variety of reasons.

Anhanguera is the type genus of the anhanguerids, the group that would traditionally have been called ornithocheirids, and in a weird way, it's quite fitting, since a lot of the anatomical features that were used to erect the type species, Anhanguera blittersdorffi , are (according to more recent studies) plesiomorphic traits widely distributed across the Anhangueridae family, making it hard to pinpoint which other species meet the criteria of belonging in this genus, outside of A. blittersdorffi itself, who is known from a complete holotype skull (nearly 50 cm long) and a second skull with a jawbone. Not helping matters is that other species of Anhanguera were originally named based on incomplete material but later, more complete specimens were (rather arbitrarily in hindsight) assigned to some of these species, muddying things.

Those are Anhanguera santanae, originally named as a species of Araripesaurus, and Anhanguera araripensis, originally named as a species of Santanadactylus, before both got reclassified by Alexander Kellner in 1990. Araripesaurus (1971) and Santanadactylus (1980) both predate Anhanguera but were both named from fragmentary postcranial material, making their validity dubious today, with the type species of Santanadactylus (S. brasilensis) even being suggested to be a chimera.
A. santanae was originally described based on the back half of a skull and bits of wing bone , but a much more complete skull (45 cm) and postcranial skeleton (AMNH 22555) were later assigned to it, and this specimen became the go-to anatomical reference for Anhanguera for many years. A. araripensis was named based on an incomplete skull (just the midsection) but later, two very complete skulls (MN 4735-V and SAO 16494) were assigned to the species, with MN 4735-V having a taller crest than most other specimens. There is also Tropeognathus robustus, described from a jawbone (the same year as T. mesembrinus) and later the tip of a snout, and was later renamed as Anhanguera robustus in 2013. However, since all three of these species were described based on very incomplete and thus not particularly diagnostic material, a 2017 study found them all to be nomen dubia, while other, more complete specimens like AMNH 22555 and MN 4735-V are relegated to Anhanguera sp., since many of the subtle differences between various skull specimens once used to distinguish them on a specific level could simply represent ontogenetic stages and/or sexual dimorphism, or even just individual variation (mirroring the taxonomic history of Pteranodon).

Two other, more recently described species with a more robust foundation are Anhanguera piscator (2000) and Anhanguera spielbergi (2003), who happen to be the largest alleged species of Anhanguera, both having a skull length of circa 65 cm respectively, and both have holotypes that are a largely complete skeleton, giving us a clear idea of their anatomy, which is why they are still considered distinct from A. blittersdorffi. A. spielbergi was originally named as Coloborhynchus spielbergi by André Veldmeijer, who also insisted that A. piscator and A. araripensis should be called Coloborhynchus piscator and Coloborhynchus araripensis. As I've talked about before, Coloborhynchus (just like Ornithocheirus) is an ill-defined wastebin taxon, and its type species (C. clavirostris) is a very small species described from fragmentary remains from Valanginian strata in England, much older than the various large, Mid Cretaceous fossils from across the world (often just snout tips) later attributed to the genus by some workers, but are nowadays classed as Uktenadactylus (Texa, dubiously England ), Nicorhynchus (England, Morroco), and Siroccopteryx (Morocco), as well as the aforementioned (much more comple) fossls from Brazil, though in 2019, A. spielbergi was reclassified as Maaradactylus spielbergi, the type species being the slightly larger M. kellneri (2014), so...yeah, the controversy is far from settled.

Despite the many species that have been named over the years, A. blittersdorffi and A. piscator might turn out to be the only valid species of Anhanguera, with others being ill-defined nomen dubia and various minor differences in the better-preserved specimens possibly being the result of individual variation, sexual dimorphism and/or growth stages, though of course, future studies will likely shake things up further and continue this controversy, and a larger sample size (which is likely going to turn up at Romualdo sooner or latter) would also help validate/dispute the former suggestions, as would more thorough studies on additional Anhanguera fossils that are still only known from preliminary descriptions or none. That said, it still seems plausible that many of the fossils found so far do indeed belong to the genus (just not as many species as once suggested) and Anhanguera seems to have outnumbered the giant Tropeognathus by a considerable margin on the ancient coasts of South America 110 million years ago, which makes sense for a smaller taxon.

Oh, and you'll occasionally see references to Anhanguera having survived into the Cenomanian, but there is no real evidence for it, as this was based on fragmentary pterosaur fossils from the upper Albian-Cenomanian of England, specifically Cambridge Greensand and the Chalk Group, being lumped into Anhanguera, including "Ornithocheirus fittoni" and Cimoliopterus cuvieri being identified as "Anhanguera fittoni" and "Anhanguera cuvieri" by some workers, but Cimoliopterus is now recognized as a distinct genus, belonging to the newly erected cimoliopterid targaryendraconians, distant cousins to the anhanguerid anhanguerians, and any other fossils are far too fragmentary to be confidently assigned to Anhanguera.