Anhanguera taxonomy revisited: is our understanding of Santana Group pterosaur diversity biased by poor biological and stratigraphic control?

Background Anhanguerids comprise an important clade of pterosaurs, mostly known from dozens of three-dimensionally preserved specimens recovered from the Lower Cretaceous Romualdo Formation (northeastern Brazil). They are remarkably diverse in this sedimentary unit, with eight named species, six of...

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Main Authors: Felipe L. Pinheiro, Taissa Rodrigues
Format: Article
Language:English
Published: PeerJ Inc. 2017-05-01
Series:PeerJ
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Online Access:https://peerj.com/articles/3285.pdf
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author Felipe L. Pinheiro
Taissa Rodrigues
author_facet Felipe L. Pinheiro
Taissa Rodrigues
author_sort Felipe L. Pinheiro
collection DOAJ
description Background Anhanguerids comprise an important clade of pterosaurs, mostly known from dozens of three-dimensionally preserved specimens recovered from the Lower Cretaceous Romualdo Formation (northeastern Brazil). They are remarkably diverse in this sedimentary unit, with eight named species, six of them belonging to the genus Anhanguera. However, such diversity is likely overestimated, as these species have been historically diagnosed based on subtle differences, mainly based on the shape and position of the cranial crest. In spite of that, recently discovered pterosaur taxa represented by large numbers of individuals, including juveniles and adults, as well as presumed males and females, have crests of sizes and shapes that are either ontogenetically variable or sexually dimorphic. Methods We describe in detail the skull of one of the most complete specimens referred to Anhanguera, AMNH 22555, and use it as a case study to review the diversity of anhanguerids from the Romualdo Formation. In order to accomplish that, a geometric morphometric analysis was performed to assess size-dependent characters with respect to the premaxillary crest in the 12 most complete skulls bearing crests that are referred in, or related to, this clade, almost all of them analyzed first hand. Results Geometric morphometric regression of shape on centroid size was highly statistically significant (p = 0.0091) and showed that allometry accounts for 25.7% of total shape variation between skulls of different centroid sizes. Premaxillary crests are both taller and anteroposteriorly longer in larger skulls, a feature consistent with ontogenetic growth. A new diagnosis is proposed for Anhanguera, including traits that are nowadays known to be widespread within the genus, as well as ontogenetic changes. AMNH 22555 cannot be referred to “Anhanguera santanae” and, in fact, “Anhanguera santanae”, “Anhanguera araripensis”, and “Anhanguera robustus” are here considered nomina dubia. Discussion Historically, minor differences in crest morphology have been used in the definition of new anhanguerid species. Nowadays, this practice resulted in a considerable difficulty in referring well-preserved skulls into known taxa. When several specimens are analyzed, morphologies previously believed to be disparate are, in fact, separated by a continuum, and are thus better explained as individual or temporal variations. Stratigraphically controlled excavations on the Romualdo Formation have showed evidence for faunal turnover regarding fish communities. It is thus possible that some of the pterosaurs from this unit were not coeval, and might even represent anagenetic morphotypes. Unfortunately, amateur collecting of Romualdo Formation fossils, aimed especially at commerce, resulted in the lack of stratigraphic data of virtually all its pterosaurs and precludes testing of these further hypotheses.
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spelling doaj.art-c916552d7cc44ef1965892ec4b25288a2023-12-03T10:55:37ZengPeerJ Inc.PeerJ2167-83592017-05-015e328510.7717/peerj.3285Anhanguera taxonomy revisited: is our understanding of Santana Group pterosaur diversity biased by poor biological and stratigraphic control?Felipe L. Pinheiro0Taissa Rodrigues1Laboratório de Paleobiologia, Universidade Federal do Pampa, São Gabriel, RS, BrazilDepartamento de Ciências Biológicas, Universidade Federal do Espírito Santo, Vitória, ES, BrazilBackground Anhanguerids comprise an important clade of pterosaurs, mostly known from dozens of three-dimensionally preserved specimens recovered from the Lower Cretaceous Romualdo Formation (northeastern Brazil). They are remarkably diverse in this sedimentary unit, with eight named species, six of them belonging to the genus Anhanguera. However, such diversity is likely overestimated, as these species have been historically diagnosed based on subtle differences, mainly based on the shape and position of the cranial crest. In spite of that, recently discovered pterosaur taxa represented by large numbers of individuals, including juveniles and adults, as well as presumed males and females, have crests of sizes and shapes that are either ontogenetically variable or sexually dimorphic. Methods We describe in detail the skull of one of the most complete specimens referred to Anhanguera, AMNH 22555, and use it as a case study to review the diversity of anhanguerids from the Romualdo Formation. In order to accomplish that, a geometric morphometric analysis was performed to assess size-dependent characters with respect to the premaxillary crest in the 12 most complete skulls bearing crests that are referred in, or related to, this clade, almost all of them analyzed first hand. Results Geometric morphometric regression of shape on centroid size was highly statistically significant (p = 0.0091) and showed that allometry accounts for 25.7% of total shape variation between skulls of different centroid sizes. Premaxillary crests are both taller and anteroposteriorly longer in larger skulls, a feature consistent with ontogenetic growth. A new diagnosis is proposed for Anhanguera, including traits that are nowadays known to be widespread within the genus, as well as ontogenetic changes. AMNH 22555 cannot be referred to “Anhanguera santanae” and, in fact, “Anhanguera santanae”, “Anhanguera araripensis”, and “Anhanguera robustus” are here considered nomina dubia. Discussion Historically, minor differences in crest morphology have been used in the definition of new anhanguerid species. Nowadays, this practice resulted in a considerable difficulty in referring well-preserved skulls into known taxa. When several specimens are analyzed, morphologies previously believed to be disparate are, in fact, separated by a continuum, and are thus better explained as individual or temporal variations. Stratigraphically controlled excavations on the Romualdo Formation have showed evidence for faunal turnover regarding fish communities. It is thus possible that some of the pterosaurs from this unit were not coeval, and might even represent anagenetic morphotypes. Unfortunately, amateur collecting of Romualdo Formation fossils, aimed especially at commerce, resulted in the lack of stratigraphic data of virtually all its pterosaurs and precludes testing of these further hypotheses.https://peerj.com/articles/3285.pdfPterosauriaAnhangueridaeRomualdo formationCretaceousAllometryGeometric morphometrics
spellingShingle Felipe L. Pinheiro
Taissa Rodrigues
Anhanguera taxonomy revisited: is our understanding of Santana Group pterosaur diversity biased by poor biological and stratigraphic control?
PeerJ
Pterosauria
Anhangueridae
Romualdo formation
Cretaceous
Allometry
Geometric morphometrics
title Anhanguera taxonomy revisited: is our understanding of Santana Group pterosaur diversity biased by poor biological and stratigraphic control?
title_full Anhanguera taxonomy revisited: is our understanding of Santana Group pterosaur diversity biased by poor biological and stratigraphic control?
title_fullStr Anhanguera taxonomy revisited: is our understanding of Santana Group pterosaur diversity biased by poor biological and stratigraphic control?
title_full_unstemmed Anhanguera taxonomy revisited: is our understanding of Santana Group pterosaur diversity biased by poor biological and stratigraphic control?
title_short Anhanguera taxonomy revisited: is our understanding of Santana Group pterosaur diversity biased by poor biological and stratigraphic control?
title_sort anhanguera taxonomy revisited is our understanding of santana group pterosaur diversity biased by poor biological and stratigraphic control
topic Pterosauria
Anhangueridae
Romualdo formation
Cretaceous
Allometry
Geometric morphometrics
url https://peerj.com/articles/3285.pdf
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