The oldest South American occurrence of Spinosauridae (Dinosauria, Theropoda)

Accepted Manuscript The oldest South American occurrence of spinosauridae (Dinosauria, Theropoda) Marcos A.F. Sales, Alexandre Liparini, Marco B. De Andrade, Paulo R.L. Aragão, Cesar L. Schultz PII:

S0895-9811(16)30131-6

DOI:

10.1016/j.jsames.2016.10.005

Reference:

SAMES 1622

To appear in:

Journal of South American Earth Sciences

Received Date: 23 July 2016 Accepted Date: 19 October 2016

Please cite this article as: Sales, M.A.F., Liparini, A., De Andrade, M.B., Aragão, P.R.L., Schultz, C.L., The oldest South American occurrence of spinosauridae (Dinosauria, Theropoda), Journal of South American Earth Sciences (2016), doi: 10.1016/j.jsames.2016.10.005. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.

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THE OLDEST SOUTH AMERICAN OCCURRENCE OF SPINOSAURIDAE

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(DINOSAURIA, THEROPODA)

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Marcos A. F. Sales,*,1 Alexandre Liparini,*,2 Marco B. De Andrade,3 Paulo R. L. Aragão,2 and

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Cesar L. Schultz1

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Federal do Rio Grande do Sul, Av. Bento Gonçalves, 9500, Agronomia, 91501-970, Porto

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Alegre, Rio Grande do Sul, Brazil, [email protected]; 2Departamento de Biologia,

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Departamento de Paleontologia e Estratigrafia, Instituto de Geociências, Universidade

Centro de Ciências Biológicas e da Saúde, Universidade Federal de Sergipe, Av. Marechal

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Rondon, s/n, Jardim Rosa Elze, 49100-000, São Cristóvão, Sergipe, Brazil,

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[email protected]; 3Faculdade de Biociências, Pontifícia Universidade

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Católica do Rio Grande do Sul, Av. Ipiranga, 6681, Partenon, 90619-900, Porto Alegre, Rio

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Grande do Sul, Brazil, [email protected]; [email protected];

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[email protected]

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Corresponding authors

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ACCEPTED MANUSCRIPT ABSTRACT

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A new fossil site, called ‘Canafístula 01’, has yielded the first archosaur remains from

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the Berriasian–Valanginian Feliz Deserto Formation, Sergipe-Alagoas Basin, northeastern

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Brazil. Most of them comprise crocodylomorph teeth and osteoderms. However, the most

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remarkable specimen is a partial tooth assigned to Spinosauridae, based on the unique

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combination of the following features: (1) unfluted root almost as wide as the crown base,

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with a large pulp cavity; (2) straight and more regularly spaced flutes of the crown, formed by

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both the enamel and the dentine; and (3) unserrated carina on a mesiodistal plane coinciding

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with the main plane of curvature of the crown. This is the oldest occurrence of a spinosaurid

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from South America. In addition, given the unserrated distal carina, this tooth is referred to

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the subfamily Spinosaurinae, representing also the oldest spinosaurine record worldwide. It

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indicates that the cladogenetic event between Baryonychinae and Spinosaurinae must have

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occurred even before the Barremian, and that the latter was already present in South America

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in pre-Aptian ages. Thus, the occurrence of a spinosaurid in the Feliz Deserto Formation

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points to a latent potential for new relevant findings in northeastern Brazil and the necessity

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for greater collection efforts in this region.

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KEYWORDS: Spinosauridae, Sergipe-Alagoas Basin, Feliz Deserto Formation, South

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America, Early Cretaceous

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ACCEPTED MANUSCRIPT INTRODUCTION

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Northeastern Brazil has plenty of sedimentary basins. Most of them underwent depositional histories related to the tectonic processes that led to the separation between

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Africa and South America. The break-up process began as the result of the reactivation of

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faults in the Precambrian crystalline basement from Middle Jurassic onwards, but mainly

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during the Early Cretaceous (Valença et al., 2003). This fact implies that most fossil sites of

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this region are Jurassic–Cretaceous in age. Among these are the Araripe and São Luís-Grajaú

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basins, from which comes much of the vertebrate paleontological record of northeastern

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Brazil. Particularly, the Aptian–Albian Crato and Romualdo formations of the former basin

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can be placed among the ten best Lagerstätten worldwide and have yielded many

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scientifically relevant specimens, e.g., the most complete Cretaceous snake and bird skeletons

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from South America and the first fossilized heart of a vertebrate (Martill, 1990; Maisey, 1991;

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Carvalho et al., 2015; Martill et al., 2015; Maldanis et al., 2016). On the other hand, the

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Cenomanian Alcântara Formation of the latter basin is one of the few extensive bone beds in

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northern South America (Medeiros et al., 2014). In addition, other smaller basins have

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contributed with many important Early Cretaceous dinosaur ichnofossils (Carvalho, 2004).

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However, the vertebrate fossil record of northeastern Brazilian intracratonic basins,

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notably those encompassing pre-Aptian stages, is still poorly known (Valença et al., 2003;

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Arai, 2006). Such a fossil record is better sampled in marginal basins like the Sergipe-Alagoas

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Basin, which is mainly known due to its post-Aptian marine strata and its important and

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abundant record of ammonites and other invertebrates, along with vertebrate remains,

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including teeth, scales, and vertebrae of fishes and mosasaurs (Koutsoukos and Bengtson,

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1993; Bengtson and Lindgren, 2005). Its depositional sequence also includes Early

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Cretaceous continental and transitional settings for some units, whose paleontological

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ACCEPTED MANUSCRIPT potential has been barely explored, possibly due to local environmental conditions and

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diagenetic control (Garcia et al., 2005). The Feliz Deserto Formation is one of such units and

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is of particular importance because its age is inferred to be Berriasian–Valanginian. So far, its

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fossil record comprises gastropod and bivalve mollusks and, among vertebrates, only remains

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of the lepisosteiform fish Lepidotes have been found (Brito, 1984).

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However, recent collections in a new locality where the Feliz Deserto Formation crops out—the outcrop called ‘Canafístula 01’—have revealed also the presence of archosaur

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remains in this unit, mostly crocodylomorph teeth and osteoderms (see below; Fig. 1). These

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new records represent an important addition to the paleontological knowledge of northeastern

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Brazil, especially regarding the pre-Aptian Cretaceous time interval. Thus, the present

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contribution reports for the first time tetrapod remains from the Feliz Deserto Formation and

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describes the most remarkable tooth among those collected so far. This specimen enabled the

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recognition of the first spinosaurid theropod from the Sergipe-Alagoas Basin, with

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paleobiogeographic implications. The new outcrop along with the remaining specimens—

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basically the crocodylomorph remains—is to be described elsewhere.

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MATERIALS AND METHODS

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All collected specimens are housed at the collections of Laboratório de Paleontologia

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of the Universidade Federal de Sergipe (LPUFS), São Cristóvão, Sergipe State, Brazil,

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including the one described below. Terminology for dental features follows Hendrickx et al.

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(2015). Measurements were taken with a digital caliper. The abbreviated description of the

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provenance follows Bengtson (1983).

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FIGURE 1. Outcrop Canafístula 01, Jaboatã municipality, Sergipe State, Brazil. A, Location

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map of the outcrop, which is indicated by the black star; B, Photograph of the northeastern

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side of the road cut with the spots where the partial tooth LPUFS 5737 and one

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crocodylomorph tooth crown were found indicated by the upper and lower hammers,

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respectively.

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RESULTS

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SYSTEMATIC PALEONTOLOGY

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DINOSAURIA Owen, 1842

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SAURISCHIA Seeley, 1887

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THEROPODA Marsh, 1881

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TETANURAE Gauthier, 1986

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SPINOSAURIDAE Stromer, 1915

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Gen. et sp. indet.

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(Fig. 2)

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Referred Specimen—LPUFS 5737, a partial tooth.

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Locality and Age— Outcrop known as ‘Canafístula 01’ (UTM DATUM WGS 84,

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Zone 24 L, 8 857 550 N/ 739 980 E, GPS error +/- 4 m) after the homonymous farm property

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nearby. The outcrop is located 2 km NW from Japoatã municipality, Sergipe State, Brazil, on

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the SE-202 Road. It comprises exposures on the northeastern and southwestern faces of a road

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cut, 200 m NW from a bridge; altitude ca. 30 m. Serviço Geológico do Brazil GEOBANK

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lithology code K1bi: yellow, fine-grained sandstones interbedded with grayish-to-rosaceous

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siltstones.

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The exposure is ca. 200 m wide with a maximum height of ca. 6 m. Its lithology

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basically encompasses layers of fine sandstones 30 to 50 cm high intercalated with fine

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siltstones no more than 10 cm thick. Three siltstone levels are each overlain by bone beds

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with many ganoid fish scales and other fragmentary bone remains. Overall, the teeth here

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reported were found in the outcrop, next to the upper-most bone bed, but detached from the

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rock. LPUFS 5737 was recovered from the upper half of the northeastern exposure (Fig. 1).

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One of the crocodylomorph teeth was collected in situ from the base of the same exposure,

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still embedded in the matrix, whereas the others came from the southwest-facing side, one

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meter above the outcrop base, among sediments eroded from upper layers.

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Schaller (1969) determined an Early Cretaceous age for the deltaic–lacustrine

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sandstones, siltstones and shales of the Barra de Itiúba Formation—later subdivided into Feliz

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Deserto and Barra de Itiúba formations—based on non-marine ostracodes and palynomorphs,

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correlating it to beds of the same age of the Recôncavo and Tucano basins. Borba et al. (2011)

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estimated a duration of approximately 5.8 Ma for the deposition of the Feliz Deserto

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Formation (sequences K10/10 to K10/40), being equivalent to the lower and middle Rio da

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Serra local stage. This is considered to be within the Berriasian–Valanginian interval. Description—LPUFS 5737 is a fragmentary tooth preserving the basal-most part of

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the crown and likely most of its root. The break that led to the loss of the apical part of the

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crown seems to have occurred along a plane oblique with respect to the tooth main axis,

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sloping downwards distally if the tooth is oriented with its apical portion upwards. Despite

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that, it is a well-preserved element; actually, the best preserved one among those collected at

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Canafístula 01. Apicobasally, the whole specimen measures 21.23 mm, but the preserved

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apical length of the crown is only 0.8 mm. The crown presents a subcircular cross-section,

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measuring 6.98 mm in length and 5.98 mm in width at the level of the cervix. In general, the

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tooth shows a conical morphology and in lateral view it has a slightly mesiodistal curvature,

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which is more obvious in its mesial outline than in the distal one. In addition, there is a

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labiolingual curvature, which is even fainter than the mesiodistal one. In fact, it is the

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labiolingual curvature that allows the identification of the labial and lingual sides, the latter

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being that towards which the crown very gently inclines. Regarding the mesial and distal

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margins of LPUFS 5737, only the latter bears a conspicuous carina, which is not serrated and

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seems to be placed on a plane that symmetrically divides the crown labiolingually, although

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the mesial carina may have been present in the lost apical portion or worn by tooth-to-food

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contact. The enamel is well-preserved and clearly reached the cervix on both sides. Also,

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LPUFS 5737 clearly presents ten and eleven flutes respectively on the labial and lingual sides.

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Interestingly, the distance between the distal-most flute and the distal carina differs between

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sides, being greater on the lingual face. Besides the flutes, no other major ornamentation, such

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as marginal undulations, can be seen. Finally, concerning the root, it presents a smooth

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surface, without any discernible ornamentation or other attribute, and its cross-section is also

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subcircular.

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FIGURE 2. Specimen LPUFS 5737. A, labial view; B, lingual view; C, Apicodistal view of

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the apical-most preserved portion of the tooth crown, with the flutes in the enamel indicated

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by white arrows; D, apical view of the tooth root. Scale bars equal 10 (for A, B, and D) and 5

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mm (for C).

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Remarks—The preserved root makes LPUFS 5737 attributable to groups with a

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typical thecodont dentition. Despite bearing at least one carina and being slightly

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mesiodistally curved, the generally conical shape of the crown, along with its fluted condition,

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places it within the conidont category of Hendrickx et al. (2015), rather than in the ziphodont

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category (Langston, 1975; Andrade et al., 2010; Hendrickx et al., 2015). In fact, the

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combination of thecodont tooth attachment and conidonty are found in the dentition of some

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typically aquatic to semiaquatic taxa and/or with a piscivorous diet (e.g., Massare, 1987;

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Charig and Milner, 1997).

ACCEPTED MANUSCRIPT However, LPUFS 5737 can be set apart from sauropterygians and other marine

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reptiles based on the unique combination of the following features: (1) unfluted root almost as

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wide as the crown base, with a large pulp cavity; (2) straight and more regularly spaced flutes

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of the crown, formed by both the enamel and the dentine; and (3) unserrated carina on a

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mesiodistal plane coinciding with the main plane of curvature of the crown (Owen, 1851;

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Brown, 1981; Buffetaut and Ingavat, 1986; Massare, 1987; Buffetaut et al., 2008; Andrade et

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al., 2010; Hone et al., 2010). Regarding toothed pterosaurs, especially the piscivores, their

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flutes, when present, are usually found only on the dentine. Those on the enamel are

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ocasionally found in crowns much more labiolingually compressed and slender than LPUFS

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5737. Another feature commonly seen in pterosaur teeth but absent in the tooth described

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herein is the enamel often covering the crown only partially, exposing the dentine in basal

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portions, especially on the lingual side (Owen, 1851; Wellnhofer and Buffetaut, 1999;

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Sánchez-Hernández et al., 2007). All of this clearly precludes an attribution of LPUFS 5737

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to Pterosauria.

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The only conidont taxon showing all major features described in the previous section is Spinosauridae, including the typical theropod feature of the mesiodistal plane as the main

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plane of curvature of the crown (Buffetaut and Ingavat, 1986; Currie et al., 1990; Sánchez-

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Hernández et al., 2007; Buffetaut et al., 2008). Although crocodylomorph teeth may be

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reminiscent of spinosaurid teeth, this resemblance is indeed shallow. For instance, in most

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crocodylomorphs, including conical-toothed neosuchians, the main plane of curvature of the

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crown is labiolingual (Buffetaut and Ingavat, 1986; Sánchez-Hernández et al., 2007; Buffetaut

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et al., 2008; Hone et al., 2010). The more refined and dendritic pattern of flutes in

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neosuchians also contrasts with LPUFS 5737 (MAFS and MAB, personal observations).

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Some coelurosaurian theropods show a sort of conidonty, but their crowns are not fluted

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(Pérez-Moreno et al., 1994; Hendrickx and Mateus, 2014; Hendrickx et al., 2015).

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With respect to other features of LPUFS 5737, a faint or an indiscernible mesial carina seems to be more common among Spinosauridae than previously thought, especially

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regarding those with unserrated teeth (Sánchez-Hernández et al., 2007), as shown by some

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illustrations of cross-sections of spinosaur crowns (Stromer, 1915:pl. 1, fig. 7c; Medeiros,

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2006:fig. 2f,g). Also, despite being found in some spinosaurid taxa or isolated teeth, marginal

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undulations are not reported in other specimens, as in LPUFS 5737 (Stromer, 1915; Sues et

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al., 2002; Medeiros, 2006; Sánchez-Hernández et al., 2007; Hendrickx and Mateus, 2014).

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Other traits, like the degree of labiolingual compression and the distribution of flutes, can

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vary within the tooth row (Dal Sasso et al., 2005; Hendrickx and Mateus, 2014). Finally,

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although exhibiting flutes only on the lingual side, the sixth right premaxillary crown of the

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spinosaurid Angaturama limai is similar to LPUFS 5737 both in size and the general aspect of

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the flutes (Kellner, 1996; Kellner and Campos, 1996).

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DISCUSSION

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Given the Berriasian–Valanginian age of the Feliz Deserto Formation, LPUFS 5737

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represents a spinosaurid theropod in the pre-Aptian Early Cretaceous of northeastern Brazil,

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being the oldest occurrence of a South American spinosaur. Known and undisputed

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spinosaurid records from South America include three described taxa, Irritator challengeri,

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Angaturama limai, and Oxalaia quilombensis, along with post-cranial remains and isolated

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teeth, from the Albian and the Albian–Cenomanian of Araripe and São Luís-Grajaú basins,

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respectively (Fig. 3; Kellner and Campos, 1996; Sues et al., 2002; Medeiros, 2006; Machado

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and Kellner, 2007, 2008; Kellner et al., 2011; Medeiros et al., 2014; Sales et al., 2016).

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FIGURE 3. A, orthographic projection of the world showing punctual spinosaurid

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occurrences. The South American records are indicated by triangles, the black and white ones

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representing São Luís-Grajaú and Araripe basins, respectively. The record of LPUFS 5737

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from the Sergipe-Alagoas Basin is indicated by the black star. Undisputed Barremian

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spinosaurid records are shown by black circles. B, Cretaceous fossil record of Spinosauridae

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after the database of Sales et al. (2016). Only formally published records with images of the

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specimens were included here. Spinosaurine and baryonychine fossil records are indicated by

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high-spined and low-spined spinosaur body icons, respectively. The black bars represent more

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complete specimens assigned at least to generic taxa, whereas the white ones comprise more

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fragmentary or putative records. The dashed lines represent the extended history of each clade

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due to the Berriasian–Valanginian age of LPUFS 5737 and its attribution to a spinosaurine

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spinosaur. The numerical ages (in millions of years ago) of the Cretaceous subdivisions are

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indicated by numbers at the left and follow Cohen et al. (2013). The map in A is from Alroy

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(2013) and the body icons in B were modified from Sales et al. (2016).

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Traditionally, spinosaurid taxa have been classified in two subfamilies, Baryonychinae

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and Spinosaurinae (Sereno et al., 1998; Carrano et al., 2012). Besides other craniodental

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features, the former is characterized by serrated teeth, whereas the latter shows the unserrated

ACCEPTED MANUSCRIPT condition. Following this taxonomic proposal, LPUFS 5737 would be referable to

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Spinosaurinae, based on its unserrated distal carina. It is consistent with the similarity

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between this tooth and Angaturama limai, another spinosaurine spinosaurid from northeastern

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Brazil. Thus, it might also correspond to the oldest spinosaurine occurrence, whereas the

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previous oldest spinosaurine records are referred to the Barremian of Spain (Fig. 3, Sánchez-

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Hernández et al., 2007). If later further supported, the occurrence of a spinosaurine in the

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Berriasian–Valanginian of Gondwana would indicate that the cladogenetic event between

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Spinosaurinae and Baryonychinae occurred even before the previously suggested minimum

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age based on Barremian remains (e.g., Charig and Milner, 1997; Sánchez-Hernández et al.,

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2007; Le Loeuff et al., 2010; Carrano et al., 2012). Furthermore, a litteral interpretation of the

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fossil record would suggest western Gondwana as the center of origin and dispersal for

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Spinosaurinae, and that dispersal events of this clade throughout Gondwana may have

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occurred well before the ~Aptian split between Africa and South America. However, a recent analysis has shown spinosaurid taxa forming a politomy, and the

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boundary between typical features of each subfamily may not be so clear-cut (Evers et al.,

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2015). Actually, given the proposed scenario for the evolution of spinosaurid teeth, passing

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from the plesiomorphic coarsely-serrated teeth to the finely-serrated teeth and then to the

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unserrated teeth (Buffetaut, 2011; Serrano-Martínez et al., 2016), it is possible that in the

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future baryonychines are shown to be, in fact, successive outgroups of Spinosaurinae.

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Another possible scenario would be that unserrated teeth evolved more than once during the

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spinosaurid evolutionary history. Anyway, LPUFS 5737 indicates that unserrated-toothed

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spinosaurs had already evolved and were present in Gondwana—more specifically South

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America—during the Berriasian–Valanginian.

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Spinosauridae is the first identified tetrapod taxa from the Feliz Deserto Formation, based on described remains. The crocodylomorph teeth and osteoderms here reported are yet

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enlarge the diversity of the Feliz Deserto fossil assemblage. Considering that most of the

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knowledge on Early Cretaceous faunas of northeastern Brazil, especially regarding body

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fossils, come from mid-Cretaceous (Aptian–Cenomanian) strata (e.g., Martill, 1990; Maisey,

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1991; Medeiros et al., 2014), the Feliz Deserto Formation represents an important source of

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data on the faunal diversity of northern South America during pre-Aptian ages. It also points

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to a latent potential for new relevant findings in northeastern Brazil outside the Lagerstätten

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Crato and Romualdo formations of the Araripe Basin and the necessity for greater collection

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efforts in this region.

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ACKNOWLEDGMENTS

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LPUFS 5737 photographs were skillfully taken by Luís Flávio Lopes (Universidade Federal do Rio Grande do Sul). We warmly thank Antonio J. V. Garcia (Universidade Federal

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de Sergipe), whose knowledge helped us distinguish Feliz Deserto Formation from Barra de

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Itiúba Formation, and Rainer Schoch (Staatliche Museum für Naturkunde Stuttgart) for

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providing access to the specimens under his care. Soki Hattori (University of Tokyo),

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Kenneth Monsch (Naturalis Biodiversity Center), Felipe L. Pinheiro (Universidade Federal do

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Pampa), Heitor Francischini (UFRGS), and Maria E. P. Batista (Universidade Federal do

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Ceará) kindly read and commented on early drafts of the manuscript. Support was provided

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by a Ph.D. scholarship grant (to MAFS) from the Conselho Nacional de Desenvolvimento

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Científico e Tecnológico (CNPq), a short-term grant (to MAFS) from the Deutscher

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Akademischer Austauschdienst (DAAD), and scientific initiation scholarship grants (to

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PRLA) from the CNPq and Coordenação de Pesquisa da Universidade Federal de Sergipe.

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