First report of pterosaur remains from the Lower Cretaceous Quiricó Formation, São Francisco Basin (Minas Gerais), Brazil

FIALHO, FLÁVIA SIBELE F.; SANTUCCI, RODRIGO M.; GOMES, MARIANA S.; CARMO, DERMEVAL A. DO

doi:10.1590/0001-3765202520241140

Abstract

The record of pterosaur fossils in Brazil is significant, however, it remains limited to a few sedimentary basins from the Cretaceous period. This study describes an isolated pterosaur tooth assigned to Ornithocheiriformes, collected from the Quiricó Formation, São Francisco Basin, Early Cretaceous (Aptian), of Minas Gerais State. The tooth was found in association with fossil plants and articulate gonorynchiform fishes. The tooth has a narrow elliptical cross-section, with marked mesial and distal carinae bearing no denticles. In lateral view, the tooth is triangular, being slightly curved lingually. The enamel is thin with no striations. These features, coupled with the Lower Cretaceous age of the deposits it was found, suggest the tooth belongs to an ornithocheiriform pterosaur. This finding represents the first report of pterosaur material from the São Francisco Basin and fills a geographical gap when previous records of ornithocheiriforms in Brazil are considered (Araripe, Bauru, Grajaú, and Recôncavo basins), suggesting a broader distribution of these flying archosaurs in Brazil. Also, this new ornithocheiriform record in the São Francisco Basin strengthen the similarity of its vertebrate fauna to the vertebrates found in Kem Kem Beds in Morocco.

Key words
pterosaur; Quiricó Formation; Lower Cretaceous; Aptian; Ornithocheiriformes; tooth

INTRODUCTION

Several studies have reported pterosaur findings in Brazil, which vary in terms of number of taxa, but remain concentrated to few sedimentary basins: the Araripe Basin (Ceará), the Caiuá Group (Bauru Basin from Paraná), the Recôncavo Basin (Bahia), and the São Luís-Grajaú Basin (Maranhão). All these occurrences are, so far, restricted to the Cretaceous period. Among the earliest known findings, the Araripe Basin stands out as the most significant due to the high number of taxa, number and quality of its record, as demonstrated by numerous studies conducted over the past decades (e.g. Price 1971, Campos & Kellner 1985, Leonardi & Borgomanero 1985, Dalla Vecchia & Ligabue 1988, Kellner & Campos 1988, Kellner 1989, 2013, Kellner & Tomida 2000, Veldmeijer 2003, Sayão & Kellner 2006, Veldmeijer et al. 2009, Martill 2011, Elgin & Campos 2012, Pinheiro & Schultz 2012, Kellner et al. 2013, Aires et al. 2014, Pinheiro & Rodrigues 2017, Bantim et al. 2014, Leal et al. 2018, Cerqueira et al. 2021, Duque et al. 2023). This basin is recognized as one of the regions with the highest diversity of pterosaurs in both Brazil and the world, highlighting its importance for the study of this group. Notably, the Romualdo Formation is renowned as one of the most significant fossil deposits of the upper Aptian, Early Cretaceous (Guzmán et al. 2023), and is globally recognized as a Konservat-Lagerstätte (Maisey 1991, Martill 2007, Kellner et al. 2002). The unit hosts a significant variety of fossils, including vertebrates, invertebrates, and plants (Castro et al. 2006). Additionally, it has produced numerous species of pterosaurs, assigned to various genera and clades, like Anhanguera piscator, Cearadactylus atrox, Maaradactylus kellneri, Tapejara wellnhoferi, Caupedactylus ybaka, Thalassodromeus sethi, Ludodactylus sibbicki, among other species (Kellner & Tomida 2000, Leonardi & Borgomanero 1985, Bantim et al. 2014, Elgin & Campos 2012, Kellner et al. 2013, 2002, Pinheiro & Schultz 2012, Frey et al. 2003a, b, Vila Nova et al. 2011, Pinheiro & Rodrigues 2017).

The São Luís-Grajaú Basin, located on the Brazilian equatorial margin, comprises Cretaceous deposits formed during the Aptian-Albian and Albian-Cenomanian intervals (Rossetti 2001, Rossetti et al. 2004). The Alcântara Formation is considered to be formed during the late Albian to the Cenomanian time interval (Rossetti & Truckenbrodt 1997, Rossetti 2001). During this time, the opening of the Atlantic Ocean led to the separation of South America and Africa and the formation of sedimentary basins along their respective coasts, within a context of transgressive coastal deposits (Aranha et al. 1990, Góes & Rossetti 2001, Haq 2014). In the Alcântara Formation, isolated pterosaur teeth attributed to Ornithocheiroidea have been identified (Elias et al. 2007, Lindoso et al. 2011). The first study described nine isolated teeth from the Lage do Coringa locality, in the State of Maranhão. The second one highlighted the paleoecological relevance of a few hundreds of teeth collected from conglomerates and siltstones of the Alcântara Formation. In such work, the authors reviewed, described, and classified the teeth into four distinct morphotypes within the Ornithocheiroidea (Lindoso et al. 2011).

The Recôncavo Basin, located in the State of Bahia, is notable for the record of two isolated teeth found in association with coelacanth remains of the species Mawsonia gigas (Mawson & Woodward 1907, Rodrigues & Kellner 2010). The Recôncavo Basin is predominantly composed of lacustrine and fluvial deposits considered as Lower Cretaceous in age (Da Silva et al. 2007). Rodrigues & Kellner (2010) attributed this material to Anhangueria based on several morphological features like an elongated, narrow, and uniformly curved crown, oval cross-section, absence of visible carina, and the presence of longitudinal striations. According to the authors, these characteristics are consistent with the teeth of other Anhangueridae.

More recently, pterosaur records have been found in the Caiuá Group, Bauru Basin, represented by many specimens of different ontogenetic stages of at least two distinct species (Manzig et al. 2014, Kellner et al. 2019). Manzig et al. (2014) identified the species Caiuajara dobruskii in Cruzeiro do Oeste, State of Paraná, Brazil, from deposits of the Goio-Erê Formation. Posteriorly, from the same locality, Kellner et al. (2019) described a second species, Keresdrakon vilsoni. A third species, Torukjara bandeirae, was described by Pêgas (2024). These three species were discovered in a rare bone bed, located in a deposit considered as a paleodesert. The age of the Goio-Erê Formation (and the Caiuá Group as a whole) is disputed, with proposals regarding an Aptian-Albian (Batezelli 2015) or a Turonian-Campanian (Basilici et al. 2012) interval to these deposits.

This work aims to describe an isolated pterosaur tooth found in papyraceous shales, associated with Gonorynchiformes fish fossils and plants from the lacustrine sequence of the Quiricó Formation. Therefore, this study adds a new occurrence to the Brazilian pterosaur localities, suggesting that these archosaurs were more common than previously thought, with a more homogeneous geographical distribution. This record represents the first pterosaur occurrence in the São Francisco Basin, adding a new clade to the list of paleontological discoveries in the region. In addition to the description of the tooth, a comprehensive review of pterosaur records in Brazilian basins during the Cretaceous was conducted, aiming to understand the distribution of these animals throughout the period and to highlight the group diversity in Brazilian sedimentary basins.

GEOLOGICAL SETTING

The São Francisco Basin is an intracratonic unit, comprising geological strata ranging from the Upper Proterozoic to the Cenozoic. In lithostratigraphic terms, the basin is divided into nine groups: Espinhaço, Araí, Macaúbas, Paranoá, Bambuí, Santa Fé, Areado, Urucuia, and Mata da Corda (Campos & Dardenne 1997, Zalán & Romeiro-Silva 2007). Among these units, the Areado Group (Cretaceous) stands out for its high fossil content. This group crops out throughout the basin and shows some lateral lithological variations, reflecting different depositional systems. It is composed of the Abaeté, Quiricó, and Três Barras formations (Campos & Dardenne 1997, Zalán & Romeiro-Silva 2007).

The Abaeté Formation is the basal unit of the Areado Group, attributed to the Lower Cretaceous, comprising high-energy and gravity-driven deposits. In the southern São Francisco Basin, the Abaeté Formation presents polymictic conglomerates dominated by metasiltstone clasts from the local basement, while in other areas of the basin, they are oligomictic and clast-supported. The Três Barras Formation (the uppermost unit of the São Francisco Basin) is characterized by the deposition of sediments from a fluvio-deltaic system. By the end of the Early Cretaceous, the region underwent arid conditions, culminating in the formation of eolian dunes and interdunes in the upper portion, which are represented by the Três Barras Formation (Fragoso et al. 2011, Leite et al. 2018). It overlies the Quiricó Formation and is distinguished by its lithological diversity and significant volume of rocks. These rocks were deposited by fluvial, fluvio-deltaic, and eolian systems, resulting in a heterogeneous composition of sandstones (Campos & Dardenne 1997, Campos & Do Carmo 2005, Leite et al. 2018).

The Quiricó Formation represents a lacustrine system, presenting a stratigraphic sequence composed of siltstones at the base, sandstones in the middle portion, and micritic limestones and shales at the top. Among the formations of the Areado Group, it is the only one that contains ostracod fossils (Leite et al. 2018) (Fig. 1). In the region of Presidente Olegário, where fossils of Dastilbe moraesi are found, the shales exhibit a grayish/black coloration interbedded with laminae of siltstones with quartz grains and carbonate cement.

Figure 1
Geological map of the study area and location of the sampling site, lithostratigraphic column of the Quiricó Formation, Areado Group, on the riverbanks of the São José River, municipality of Presidente Olegário, Minas Gerais, Brazil.

Figure 2
Sampling scheme adopted for the collection of water, sediment, and fish in ten streams in the Amazon region. a. The letters (a-k) represent the transects marked every 15 meters; the longitudinal sections refer to the segments between transects, indicating the locations where water, sediment, and fish samples were collected; b. illustrates the water collection process; c. represents sediment collection; d. shows fish collection.

The top of the sequence is rich in fossils of plants, ostracods, fish, and other vertebrates (Leite et al. 2018, 2024, Leite & Do Carmo 2021, Campos & Do Carmo 2005). It has a wide geographic distribution and elongated morphology from south to north, in addition to isolated depocenters with varying thicknesses, indicating multiple water bodies connected during wet periods and isolated during arid periods, rather than forming a single lake (Sgarbi et al. 2001). The thickness of this formation varies from up to 100 m in the center of the basin to just a few meters at the edges (Fig. 1).

Figure 3
Isolated tooth (MAF 755), drawings. (a) lingual view, (b and d) mesial and distal views, (c) labial view, and (f) cross-section near the crown apex, scale bar = 1 cm. (e) detail of the carina, scale bar = 2 mm. Abbreviation: ca, carina

Figure 4
Geographic distribution map of pterosaur clades in Brazil with their respective geological units and taxonomic assignment. Taxonomic affinities based on: Anhanguera (Andres & Myers 2013), Araripedactylus (Wellnhofer 1977, Kellner & Tomida 2000), Arthurdactylus (Frey & Martill 1994), Aymberedactylus (Pêgas et al. 2016), Barbosania (Elgin & Frey 2011, Pêgas et al. 2019), Brasileodactylus (Veldmeijer et al. 2009), Caiuajara (Manzig et al. 2014), Caupedactylus (Kellner 2013), Cearadactylus (Holgado & Pêgas 2020), Coloborhynchus (Holgado & Pêgas 2020), Kariridraco (Cerqueira et al. 2021), Keresdrakon (Kellner et al. 2019), Lacusovagus (Wu et al. 2017), Ludodactylus (Holgado & Pêgas 2020), Maaradactylus (Holgado & Pêgas 2020), Tapejara (Campos & Kellner 1997), Thalassodromeus (Longrich et al. 2018), Torukjara (Pêgas 2024), Tropeognathus (Holgado & Pêgas 2020), Tupandactylus (Frey et al. 2003b, Andres et al. 2014), Tupuxuara (Kellner & Campos 1988), Unwindia (Martill 2011)

Figure 5
Paleogeographic reconstruction of two Cretaceous time intervals showing the occurrences of pterosaur taxa with their respective taxonomic affinities: a) Albian-Cenomanian; b) Barremian-Aptian. (paleogeographic reconstructions from Scotese 2014)

The lower contact commonly occurs with the Abaeté Formation or directly with the metasediments of the Bambuí Group, while the upper contact laterally interdigitates with the Três Barras Formation. Near the margins of the lacustrine system, evaporitic lithotypes and carbonate crusts are observed. Eolian influence is evident at several levels of laterally continuous sandstones, composed of eolian sand grains, coexisting with pelites and reworked dunes (Campos & Dardenne 1997, Leite et al. 2018, Leite & Do Carmo 2021).

Regarding the stratigraphic level where the material described here was found, it corresponds to the same level where fossil fish and leaves were previously reported and are considered to be Aptian in age (Arai et al. 1995, Leite & Do Carmo 2021, Leite et al. 2024). In the outcrop along the São José Stream, the deposits mainly represent a lacustrine sequence formed under more arid conditions, which is evidenced by the presence of evaporitic levels. The presence of shales indicates periods of lake flooding, leading to the deposition of fine sediments rich in organic matter. The presence of pelitic layers, shales, and well-preserved fish and leaf fossils suggests they underwent little transport (Leite et al. 2018). The pterosaur tooth was found at the stratigraphic level Am 3.6 meters, showing no signs of abrasion or transport (Fig. 1).

MATERIALS AND METHODS

The tooth described in this work (MAF 755) is housed in the macrofossil collection of the Geosciences Museum (MGEO) under the care of the Micropaleontology Laboratory (LabMicro) of the University of Brasília, Brasília, Brazil. The collection horizon is located in an outcrop of papyraceous shales from the Quiricó Formation, São José Farm, Municipality of Presidente Olegário, State of Minas Gerais (Fig. 1).

For imaging and morphological measurements, a DVM 6 digital microscope (LEICA®) with a high-magnification objective, maximum FOV of 12.55 mm, distance of 33 mm, and magnification range of 46x to 675x was used. These analyses were carried out at the National Institute of Criminalistics (INC) of the Federal Police in Brasília.

SYSTEMATIC PALEONTOLOGY

Pterosauria Kaup 1834

Pterodactyloidea Plieninger 1901

Pteranodontoidea Andres et al. 2014

Ornithocheiromorpha Andres et al. 2014

Material: An isolated tooth (MAF 755), represented by the complete crown lacking the root.

Locality: Papyraceous shales, banks of the Meloso River, São José Farm, Municipality of Presidente Olegário, Minas Gerais, Brazil. Other fossils are present in this locality, such as Gonorynchiformes fish and leaf impressions. Lower Cretaceous, Aptian (Arai et al. 1995, Leite & Do Carmo 2021, Leite et al. 2024).

Description: the tooth has a relatively well-preserved crown, with portions where the enamel is lacking (eroded). The dentin is black, and the enamel is grayish/beige in color and translucent. At the base of the crown, the dentin appears smoother and lacks striations, the aspect of the dentin surface, with some rugose areas, suggests it underwent some kind of dissolution. The total height of the crown is 22.6 mm. The dimensions at the base of the crown are 8.95 mm in mesio-distal length and 2.2 mm in width. The exact position of the tooth was not defined. However, based on its shape and comparisons with other Ornithocheiriformes, it likely belongs to the posterior portion of the skull or mandible, as these portions have a labio-lingually narrower cross-section. The tooth cross-section is more elongated mesiodistally and labio-lingually narrower, with an elliptical cross-section. A carina is present on both mesial and distal margins and slightly offset lingually at the top of the crown. The carinae are well-preserved and extend from the apex to the middle portion of the crown, displaying a slight undulation that does not form serrations (true denticles). In both labial and lingual views, the mesial and distal margins are symmetrical and straight, giving the tooth a symmetrical profile, assuming an acute triangle profile of approximately 21° at the crow apex. The labial and lingual sides differ, with the labial one being slightly convex while the lingual side is nearly flat. The crown shows a slight distal curvature. The enamel is smooth and lacks ornamentation or striations, with cracks due to diagenetic alteration. Longitudinal lines are present due to the cracking pattern of the enamel, with these filled by sediment. The enamel has an average thickness of 60 µm (Figs. 2 and 3).

DISCUSSION

Comparisons

The taxonomic assignment of isolated teeth can be challenging, especially when the material in question does not exhibit distinctive features unique to a specific group. To date, various vertebrate groups have been reported from the Quiricó Formation, including fish (Silva-Santos 1955, 1985, Carvalho 2002, Machado 2004, Cupello et al. 2016, Ribeiro et al. 2018, Fragoso et al. 2021, Carvalho & Santucci 2021, Ribeiro et al. 2022), sauropod dinosaurs (Titanosauriformes and Rebbachisauridae) and theropods (Carcharodontosauridae and Abelisauridae), as well as fragments of lizards such as Paramacellodidae and Borioteiioidea (Zaher et al. 2011, 2020, Carvalho & Santucci 2018, 2024, Bittencourt et al. 2020).

Regarding the fossil fish described for the Quiricó Formation, including groups such as Mawsoniidae (Carvalho 2002, Machado 2004), Hybodontidae (Cupello et al. 2016), Lepisosteidae (Carvalho & Santucci 2018, 2021, Fragoso et al. 2021), and Gonorynchiformes (Silva-Santos 1955, 1985, Ribeiro et al. 2018, 2022), it is important to highlight the significant morphological variation observed within these groups, both in size and shape. Specifically, Lepisosteidae teeth are small, styliform, and robust, with either flattened or rounded cusps. Hybodontidae teeth are also small, generally millimetric, with elevated crowns that are flattened labiolingually. In contrast, Gonorynchiformes lack teeth entirely. Mawsoniids, however, exhibit distinctly different dental plates. Due to these morphological differences, the tooth MAF 755 cannot be assigned to any fish group previously reported for the Quiricó Formation.

Among archosaurs, MAF 755 differs from the teeth of the sauropod groups found in the Quiricó Formation (Titanosauriformes and Rebbachisauridae) by being more laterally flattened, having a crown with a triangular apex, and relatively well-developed keels, which are not seen in the pencil- or spoon-like teeth typically found in these sauropods (Wilson & Sereno 1998, Wilson 2002, Upchurch et al. 2004, Zaher et al. 2011).

Regarding the teeth of Carcharodontosauridae, as described by Carvalho & Santucci (2018), they have an elliptical cross-section with denticles on the carinae. The mesial side is slightly curved, while enamel wrinkles are notable in the crown. These features differ significantly from tooth MAF 755. In comparison with Abelisauridae teeth, the main difference regards the shape of the crown, characterized by a mesial margin curved backward, while the distal portion is almost straight (Smith et al. 2005, Hendrickx & Mateus 2014, Carvalho & Santucci 2018), being different from MAF 755.

The tooth MAF 755, resembles the general morphology of some plesiosaur teeth—due to its triangular, symmetrical shape, apex of the crown exhibiting an acute angle, as well as its lingual curvature— for example, like Gronausaurus wegneri (Hampe 2013), Leptocleididae cf. Leptocleidus (Bunker et al. 2022), Scanisaurus (Kear et al. 2017) and Cryptoclididae (Weryński & Błażejowski 2023). However, plesiosaur teeth differ from MAF 755 by presenting a more circular cross-section, significantly thicker enamel (greater than 100 µm), and well-developed apicobasal ridges on the crown (e.g., Kear et al. 2017, Bunker et al. 2022, Weryński & Błażejowski 2023). Furthermore, plesiosaur occurrences in continental deposits are extremely rare and controversial (e.g., Bunker et al. 2022).

The clade Ornithocheiriformes encompasses various taxa that share similarities with specimen MAF 755, although unequivocal taxonomic identification based on dental characteristics is hindered by inter- and intraspecific heterodonty in this clade, as pointed by Ciaffi & Bellardini (2024). The absence of clear diagnostic features makes the taxonomy of isolated teeth, such as MAF 755, challenging. Nevertheless, the significance of this specimen in the context of the São Francisco Basin surpasses taxonomic limitations, representing the first record in the region. The morphological variability observed in the shape, cross-section, curvature, and size of the crowns in Ornithocheiriformes teeth from other Brazilian basins also tangles a proper identification and comparison.

However, features similar to pterosaur teeth of morphotype 2, described by Wellnhofer & Buffetaut (1999) in the Kem Kem Beds, including long, slender teeth with an elliptical and labiolingually narrow cross-section, and the presence of a developed mesial and distal carinae, show that MAF 755 is similar to Ornithocheiriformes pterosaur teeth from Morocco. Moreover, it is also quite similar in general morphology to species like Ludodactylus sibbicki (Frey et al. 2003a, Rodrigues & Kellner 2010) from the Araripe Basin.

Additionally, the enamel thickness and surface ornamentation of MAF 755 exhibit similarities with other Brazilian pterosaurs. Despite the morphological variation, certain features, such as the elliptical cross-section, elongated and slender teeth with slight distal curvature, and the presence of longitudinal ridges, not present in the tooth of the Quiricó Formation, are common among toothed pterosaurs from the Romualdo Formation (Kellner 2003, Rodrigues & Kellner 2010), allowing for the classification of MAF 755 within the clade Ornithocheiriformes.

Records of isolated pterosaur tooth from Lower Cretaceous deposits of Brazil are relatively rare. They are found in the Recôncavo Basin (Barrett et al. 2008, Rodrigues & Kellner 2010), the São Luís-Grajaú Basin (Elias et al. 2007), and the Araripe Basin (Wellnhofer 1985).

A notable example from the Crato Formation in the Araripe Basin is a complete and articulated skull of Ludodactylus sibbicki, an ornithocheiriform (Frey et al. 2003a, Rodrigues & Kellner 2010). The morphology of the teeth of Ludodactylus sibbicki and other ornithocheiriforms exhibit variation in both size and overall morphology. Posterior teeth are labiolingually narrow, with an elliptical cross-section, slender and elongated crowns, and a slight distal curvature. Additionally, like other Anhangueria, Ludodactylus has mesial and distal carina in posterior teeth like MAF 755 (Frey et al. 2003a, Sayão & Kellner 2006, Veldmeijer et al. 2009, Sweetman & Martill 2010, Lindoso et al. 2011, Martill et al. 2018). Some toothed anhanguerids species like Cearadactylus atrox and Guidraco venator also have similar posterior teeth (Leonardi & Borgomanero 1985, Kellner & Tomida 2000, Wang et al. 2012) however, they differ from MAF 755 in the cross-section, which is elliptical and has carinae on the mesial and distal margins. The presence of carinae appears to be a highly relevant feature within Ornithocheiriformes. Comparisons show a wide variation in the presence of this feature, being registered in species such as Anhanguera santanae (Wellnhofer 1985) and in Morphotype II mentioned by Wellnhofer & Buffetaut (1999) from Morocco. However, this feature is absent in Siroccopteryx moroccensis (Mader & Kellner 1999, Jacobs et al. 2020), and in Morphotype I of Wellnhofer & Buffetaut (1999). An isolated tooth from Australia assigned to Anhangueria lacks carinae, with both the mesial and distal surfaces being flat (Brougham et al. 2017).

The clade Ctenochasmatidae, a non-Ornithocheiroidea Pterodactyloidea, is characterized by needle-shaped, curved, and elongated teeth, like in Huanhepterus, Gegepterus, and Moganopterus (Dong 1982, Wang et al. 2007, Lü et al. 2012), for example. The teeth morphology of these genera is quite distinct from that observed in MAF 755.

During the Cretaceous, various pterosaur groups underwent significant diversification, with some lineages partially or completely losing their dentition. Among the clades exhibiting this feature are Azhdarchidae, Nyctosauridae, Pteranodontidae, Chaoyangopteridae, and Tapejaridae, which are excluded as possible candidates to which MAF 755 could be assigned. However, the clade Dsungaripteridae, an exception within Azhdarchoidea, retains teeth with short apicobasal ridges and rounded, pointed apices, restricted to the posterior portions of the upper and lower jaws (Young 1964, Unwin 2003, Brougham et al. 2017). The species Caiuajara dobruskii (Manzig et al. 2014) and Keresdrakon vilsoni (Kellner et al. 2019), from the Bauru Basin, belong to the clade Tapejaromorpha and are also edentulous pterosaurs.

The record of this pterosaur from the São Francisco Basin adds important information for understanding the geographic distribution of these flying reptiles in Brazilian sedimentary basins. Until now, most records were restricted to the Northeast region of Brazil (e.g. Price 1971, Campos & Kellner 1985, Dalla Vecchia & Ligabue 1988, Kellner & Tomida 2000, Veldmeijer 2003, Veldmeijer et al. 2009, Martill 2011, Pinheiro & Schultz 2012, Kellner 2013, Kellner et al. 2013, Aires et al. 2014, Bantim et al. 2014, Leal et al. 2018, Cerqueira et al. 2021, Duque et al. 2023, among others), with some occurrences in Paraná (Manzig et al. 2014, Kellner et al. 2019) and records of putative basal groups in southern Brazil (Bonaparte et al. 2010, Kellner et al. 2022). Thus, this discovery in the São Francisco Basin fills a significant gap between these regions, indicating that the distribution of Brazilian pterosaurs during the Cretaceous was broader and less local than previously thought. This discovery also opens new perspectives for future research on pterosaurs in the São Francisco Basin, since new findings could provide better information on its morphology and its phylogenetic relationships. Additionally, features such as the presence of mesial and distal carinae in MAF 755 are similar to those found in pterosaurs from the Kem Kem Beds in Morocco (Wellnhofer & Buffetaut 1999, Ibrahim et al. 2020) and some anhanguerids from the Araripe Basin (Frey et al. 2003a, Rodrigues & Kellner 2010). The similarity between the vertebrate faunas of the São Francisco Basin and African basins has already been highlighted in previous studies on sauropod dinosaurs (Carvalho & Santucci 2018) and Borioteiioidea lizards (Carvalho & Santucci 2024), reinforcing the idea that vertebrate faunas were remarkably similar between these continents.

Distribution and paleogeography

To highlight the distribution and diversity of Brazilian pterosaurs during the Early Cretaceous, a synthesis of the available records was conducted, as illustrated in Figs. 4 and 5. Based on a detailed review of the literature, this analysis aims to provide a comprehensive overview of these records, contributing to a deeper understanding of the presence and variety of these archosaurs in Brazilian territory during this period. The Araripe Basin stands out as the region with the greatest diversity and number of pterosaur species in Brazil. Ornithocheiriformes have been recorded in the Araripe, São Luís-Grajaú, Recôncavo, and São Francisco basins (this work).

The chronostratigraphic distribution of Ornithocheiriformes ranges from the Aptian to the Cenomanian. The tooth MAF 755, found in the Quiricó Formation, can be considered as one of the oldest records of this clade in Brazil, as this unit is considered to be Aptian in age (Arai et al. 1995, Campos & Dardenne 1997, Leite & Do Carmo 2021). This material is coeval to species like Arthurdactylus conandoylei (Sayão & Kellner 2006) and Ludodactylus sibbicki (Frey et al. 2003a) from the Crato Formation, as well as representatives of ornithocheiriforms in the Recôncavo Basin (Rodrigues & Kellner 2010).

Other records of Ornithocheiriformes, such as for the genus Anhanguera from the Romualdo Formation (Kellner & Tomida 2000, Fastnacht 2001), are Albian in age, being coeval with pterosaur fossils from the Kem Kem Beds in North Africa. The taxa from this region comprise genera such as Anhanguera, Ornithocheirus, and Coloborhynchus (Ibrahim et al. 2020). However, the age of the Santana Group has been a subject of recent debate, since new data on ostracods and foraminifera suggested that sedimentation occurred during the development of a transitional to marine environment throughout the Aptian (Guzmán et al. 2023).

Ibrahim et al. (2020) reported at least nine pterosaur taxa from the Kem Kem Beds, including five Ornithocheiriformes and four azhdarchids, suggesting the coexistence of toothed and edentulous pterosaurs in an environment with a wide diversity of food resources and suitable areas for breeding and nesting (Ibrahim et al. 2020, Jacobs et al. 2020). These records are crucial for understanding the distribution of Ornithocheiriformes in Brazil, Africa, and Europe as well as they provide insights into the dispersal of these pterosaurs over large distances (Barrett et al. 2008, Ibrahim et al. 2020, Martill & Smith 2024).

Additionally, it is important to highlight that the diversity of pterosaurs in Brazilian sedimentary basins is significantly greater than previously described in earlier reviews, especially in the context of the opening of the South Atlantic (Pentland & Poropat 2023, Martill & Smith 2024).

CONCLUSIONS

The tooth MAF 755 exhibits a set of characteristics that allow it to be assigned as an ornithocheiriform pterosaur, including an elliptical crown cross-section with mesial and distal carinae, a triangular crown in lateral view with slight labial curvature, a smooth surface without striations, and relatively thin enamel.

This specimen represents the first pterosaur record from the Quiricó Formation, São Francisco Basin, Aptian, Early Cretaceous. The occurrence of this fossil in low-energy lacustrine depositional systems, associated with gonorynchiform fish, suggests new avenues for research in the Quiricó Formation, addressing taxonomic, systematic, paleoecological, and behavioral aspects of pterosaurs. Moreover, this discovery adds to the existing data on pterosaurs in Brazilian sedimentary basins a new occurrence, the first for the Minas Gerais State, filling a geographical gap in the Lower Cretaceous pterosaur distribution in Brazil.

Acknowledgements

We thank the owners of Fazenda São José, in the municipality of Presidente Olegário-MG, for allowing us to conduct fossil collection on their property. We are in debt to Luísa Torres Madeira for her contributions to the scientific illustrations depicted in this work and to Manuel Pereira de Oliveira Júnior for his assistance in preparing the map of Figure 1. We also thank the AABC editors, Felipe Lima Pinheiro, and an anonymous reviewer whose comments greatly improved an earlier draft of this work. Felipe Lima Pinheiro is also thanked for sharing with us some images of Ludodactylus sibbicki. We thank the financial support provided by the Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES), funding code 001, and the Decanato de Pesquisa e Inovação of the University of Brasília for supporting this research.

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Publication Dates

Publication in this collection
19 Sept 2025
Date of issue
2025

History

Received
11 Oct 2024
Accepted
4 May 2025

This is an open-access article distributed under the terms of the Creative Commons Attribution License

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[3] ANDRES B & MYERS TS. 2013. Lone Star Pterosaurs. Earth Environ Sci Trans R Soc Edinb 103(3-4): 383-398.

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[7] BARRETT PM, BUTLER RJ, EDWARDS NP & MILNER AC. 2008. Pterosaur distribution in time and space: Zitteliana 28: 61-107.

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