A forged 'chimera' including the second specimen of the protostegid sea turtle Santanachelysgaffneyi and shell parts of the pleurodire Araripemys from the Lower Cretaceous Santana Group of Brazil.

Fossils of Cretaceous sea turtles adapted to an open marine lifestyle remain rare finds to date. Furthermore, the relationships between extant sea turtles, chelonioids, and other Mesozoic marine turtles are still contested, with one key species being Santanachelys gaffneyi Hirayama, 1998, long considered the earliest true sea turtle. The species is an Early Cretaceous member of Protostegidae, a controversial clade either placed within or closely related to Chelonioidea or, alternatively, along the stem lineage of hidden-neck turtles (Cryptodira) and representing an independent open marine radiation. Santanachelys gaffneyi is one of the most completely preserved early protostegids and is therefore critical for establishing the global phylogenetic position of the group. However, the single known specimen of this taxon is yet to be described in detail. Here we describe a second specimen of Santanachelys gaffneyi from its type horizon, the Romualdo Formation (late Aptian) of the Santana Group of the Araripe basin, NE Brazil. The skeletal elements preserved include the posterior part of the skull, neck vertebrae, shoulder girdle, anterior-most and left/central part of the carapace with few peripherals, and plastron lacking most of the hyoplastra. The remaining part of the carapace was apparently completed by fossil dealers using an anterior part of the pleurodiran Araripemydidae, tentatively identified as a shell portion of cf. Araripemys barretoi, a more common Santana fossil turtle, among other indeterminate turtle shell fragments. The purpose of this paper is to report the repatriation of the specimen to Brazil and to provide a preliminary description. Supplementary Information: The online version contains supplementary material available at 10.1186/s13358-023-00271-9.

Georeferencing fossiliferous localities from Solimões and Acre Basins (Brazil) - what we know so far about Solimões Formation and future perspectives.

The Solimões and Acre basins are complex geological units related to the Andean uplift, covering the Northwestern region of Brazil, being one of the most important units due to their fossil diversity. In order to produce a document that integrates part of the fossil records of this region, we compiled/georeferenced localities from literature on which tetrapods are described, focusing on Solimões Formation but not restricted to this unit. We were able to recognize 208 localities, documented in over two centuries of reports of fossils from several taxonomic groups from the proto-Amazonia, 199 new entries in Paleobiology Database. We summarize, for each locality, its geographical position, geological information, age, and data of the paleodiversity. Most outcrops in the Amazonia region are located on river banks (~96%), while road cuts and other non-riverside outcrops represent the remainder (~4%). Most tetrapod are Mammalia, followed by Testudinata, and Crocodyliformes. This work reinforces the need for a more controlled and refined prospecting at the Solimões/Acre Basins, especially in the Solimões Formation, which represents the majority of fossiliferous records, to help answer old questions, such as dating, and new ones, here discussed, such as the paleodiversity patterns and temporal distribution among the mapped localities.

Intra-specific variation and allometry of the skull of Late Cretaceous side-necked turtle Bauruemys elegans (Pleurodira, Podocnemididae) and how to deal with morphometric data in fossil vertebrates.

BACKGROUND: Previous quantitative studies on Bauruemys elegans (Suárez, 1969) shell variation, as well as the taphonomic interpretation of its type locality, have suggested that all specimens collected in this locality may have belonged to the same population. We rely on this hypothesis in a morphometric study of the skull. Also, we tentatively assessed the eating preference habits differentiation that might be explained as due to ontogenetic changes. METHODS: We carried out an ANOVA testing 29 linear measurements from 21 skulls of B. elegans taken by using a caliper and through images, using the ImageJ software. First, a Principal Components Analysis (PCA) was performed with 27 measurements (excluding total length and width characters; =raw data) in order to visualize the scatter plots based on the form variance only. Then, a second PCA was carried out using ratios of length and width of each original measurement to assess shape variation among individuals. Finally, original measurements were log-transformed to describe allometries over ontogeny. RESULTS: No statistical differences were found between caliper and ImageJ measurements. The first three PCs of the PCA with raw data comprised 70.2% of the variance. PC1 was related to size variation and all others related to shape variation. Two specimens plotted outside the 95% ellipse in PC1∼PC2 axes. The first three PCs of the PCA with ratios comprised 64% of the variance. When considering PC1∼PC2, all specimens plotted inside the 95% ellipse. In allometric analysis, five measurements were positively allometric, 19 were negatively allometric and three represented enantiometric allometry. Many bones of the posterior and the lateral emarginations lengthen due to increasing size, while jugal and the quadratojugal decrease in width. DISCUSSION: ImageJ is useful in replacing caliper since there was no statistical differences. Yet iterative imputation is more appropriate to deal with missing data in PCA. Some specimens show small differences in form and shape. Form differences were interpreted as occuring due to ontogeny, whereas shape differences are related to feeding changes during growth. Moreover, all outlier specimens are crushed and/or distorted, thus the form/shape differences may be partially due to taphonomy. The allometric lengthening of the parietal, quadrate, squamosal, maxilla, associated with the narrowing of jugal and quadratojugal may be related to changes in feeding habit between different stages of development. This change in shape might represent a progressive skull stretching and enlargement of posterior and lateral emargination during ontogeny, and consequently, the increment of the feeding-apparatus musculature. Smaller individuals may have fed on softer diet, whereas larger ones probably have had a harder diet, as seen in some living species of Podocnemis. We conclude that the skull variation might be related to differences in feeding habits over ontogeny in B. elegans.

The tale of the headless turtle.

Pelomedusoides is the most diverse clade of side-necked turtles and there is an extensive fossil record (de Broin, 1988; Lapparent de Broin, 2000; Gaffney et al., 2006, 2011) that dates back at least to the Barremian (Lower Cretaceous) (Romano et al., 2014). Its large fossil record evidences a greater diversity in the past, particularly at the end of the Mesozoic, and exhibits a good sampling of species that are represented by skull material (Gaffney et al., 2006, 2011). As a consequence, the most complete and recent phylogenetic hypotheses for this clade (e.g. Romano et al., 2014; Cadena, 2015) are based on matrices comprising a great amount of cranial characters derived largely from Gaffney et al. (2006, 2011). In addition, it is well established that shell characters show a lot of phenotypic plasticity, even in the fossil species (Romano, 2008; Gaffney et al., 2006, 2011). In most cases it consequently is not justified to rely on "diagnostic features" of poorly informative shell-only material for describing a new species. Because of that, most authors remark new morphotypes in the literature when such aberrant specimens are recovered, but do not make any nomenclatural act by proposing a new yet poorly supported species (e.g. Romano et al., 2013; Ferreira & Langer, 2013; Menegazzo et al., 2015). Unfortunately, such a supposedly new bothremydid turtle (Pleurodira: Bothremydidae) from the Early Paleocene of Brazil was recently described based on poorly diagnostic remains (Carvalho et al., 2016; hereafter CGB, for the authors initials) and a correction of this unfounded nomenclatural act is required. In addition I present some comments on shell only material from Brazil in order to guide splitter-taxonomists to stop describing poorly preserved fossil specimens as new species.

Atolchelys lepida, a new side-necked turtle from the Early Cretaceous of Brazil and the age of crown Pleurodira.

We report a new pleurodiran turtle from the Barremian Morro do Chaves Formation, Sergipe-Alagoas Basin, Brazil. We tested the phylogenetic position of Atolchelys lepida gen. et sp. nov. by including it in a comprehensive cladistic analysis of pleurodires. The new species is a basal member of Bothremydidae and simultaneously the oldest unambiguous crown Pleurodira. The biogeographic and chronostratigraphic significance of the finding has implications for the calibration of molecular clocks studies by pushing back the minimum age of crown Pleurodira by more than 12 Ma (ca 125 Ma). The reanalysis of Pelomedusoides relationships provides evidence that the early evolution and relationships among the main lineages of side-necked turtles can be explained, at least partially, by a sequence of vicariance events.