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UFRJ Spinosaurid Anuário do Instituto de Geociências - UFRJ www.anuario.igeo.ufrj.br Spinosaurid Dinosaurs from the Early Cretaceous of North Africa and Europe: Fossil Record, Biogeography and Extinction Dinossauros Espinossaurídeos do Cretáceo Inicial do Norte da África e Europa: Registro Fossilífero, Biogeograia e Extinção. Carlos Roberto A. Candeiro1; Stephen L. Brusatte2 & André Luis de Souza1 1Universidade Federal de Goiás, Campus Aparecida de Goiânia, Laboratório de Paleontologia e Evolução, Rua Mucuri, s/n, Área 03 – St., Conde dos Arcos, 74968-755, Aparecida de Goiânia, Goiás State, Brazil. 2University of Edinburgh, Grant Institute, School of GeoSciences, James Hutton Road, Edinburgh, EH9 3FE, United Kingdom. E-mails: [email protected], [email protected], [email protected] Recebido em: 16/10/2017 Aprovado em: 14/11/2017 DOI: http://dx.doi.org/10.11137/2017_3_294_302 Abstract We review the fossil records of spinosaurid dinosaurs in order to discuss this group’s evolution and distribution in Europe and North Africa during the Early Cretaceous. Along with their eastern Laurasian distribution during the Cretaceous, these theropods have been found in coastal deposits of Europe and North Africa dated from the Barremian to the Cenomanian. The main occurrences of spinosaurid remains are in the deposits of northern Gondwana and western Laurasia, which suggests that these regions were very important in spinosaurid evolution prior to the Cenomanian. Later, spinosaurids were seemingly replaced in northern Gondwana by other top predator groups, including the abelisauroids. Keywords: theropod spinosaurids; “middle” Cretaceous; biogeography Resumo Aqui nos revisamos os registros de dinossauros de spinosaurídeos e discutimos a evolução e distribuição deste grupo na Europa e no Norte da África durante o Cretáceo Inferior. Juntamente com sua distribuição Laurasiana oriental durante o Cretáceo, esses terópodes foram encontrados em depósitos costeiros da Europa e do Norte da África datados do Barremiano ao Cenomaniano. As principais ocorrências de restos de espinossáuridos estão nos depósitos do norte de Gondwana e Laurasia ocidental, o que sugere que estas regiões foram importante de sua evolução antes do Cenomaniano. Posteriormente, os espinossáuridos foram aparentemente substituídos no norte de Gondwana por outros grupos predadores de topo, incluindo abelissauroideos. Palavras-chave: espinossaurídeos terópodes; Cretáceo “médio”; biogeograia Anuário do Instituto de Geociências - UFRJ 294 ISSN 0101-9759 e-ISSN 1982-3908 - Vol. 40 - 3 / 2017 p. 294-302 Spinosaurid Dinosaurs from the Early Cretaceous of North Africa and Europe: Fossil Record, Biogeography and Extinction Carlos Roberto A. Candeiro; Stephen L. Brusatte & André Luis de Souza 1 Introduction 2 The Spinosaurids of Western Laurasia and Northern Gondwana Spinosaurids – the group of long-snouted and possibly semiaquatic theropods – are one of the Spinosaurids may have been a globally most distinctive dinosaur groups. Fossil remains of distributed group (Barret et al., 2011). However, these predators are known from many localities from their best fossils come from the Early-mid the Lower Cretaceous of Africa (Algeria, Egypt, Cretaceous of Northern Africa and Europe. Many Niger, Sudan, Tanzania, Tunisia), Europe (England, records of spinosaurid dinosaurs have been reported Portugal, Spain), South America (Brazil), Asia from diferent locations in Algeria, Egypt, Morocco, (China, Laos, Thailand), and Australia (e.g., Kellner Niger, Portugal, Spain, Tunisia and the United & Campos, 1996; Sereno et al., 1998; Benton et al., Kingdom (Figure 2). The geological units bearing 2000; Bufetaut & Ouaja, 2002; Sues et al., 2002; these specimens range in age from the Barremian to Medeiros, 2006; Bufetaut, 2008, 2012; Bufetaut the Cenomanian. For many decades, and particularly et al., 2008; Hone et al., 2010; Barrett et al., 2011; over the last 20 years, fossils of basal and derived Kellner et al., 2011; Allain et al., 2012; Medeiros spinosaurids have been found in these strata. et al., 2014; Hendrickx et al., 2016; Sales et al., 2017). Knowledge about spinosaurids has increased All known spinosaurid species have large signiicantly in the last few decades as new fossil body sizes, elongated and laterally compressed remains, including various new species, have been skulls (some specimens also have cranial crests), found in several new Gondwanan and Laurasian and large claws on the manus, and some species also localities (Bufetaut & Ingavat, 1986; Serenoet al., possess elongated neural spines on the vertebrae 1998; Benson et al., 2009; Bufetaut, 2012; Allain et that supported some type of sail or hump, which al., 2012; Hendrickx et al., 2016). The most diverse was probably used for some combination of display, and best preserved records of these dinosaurs are thermoregulation, and perhaps even swimming from the Early-mid Cretaceous strata of North behaviors (Bailey, 1997). Africa and Europe. Spinosaurids and two other groups of large theropods, the abelisauroids and Currently, there are 11 known species of pos- carcharodontosaurids, comprise the most important sible spinosaurid dinosaurs, whose records come members of the large-bodied terrestrial predator from northern Gondwana and Laurasia. Unlike other fauna of these regions. Among these large theropods,top predators that also inhabited Gondwana (Carcha- spinosaurids have the most diverse and abundant rodontosauridae and Abelisauroidea), the spinosau- fossil record in the Early Cretaceous deposits rids show a more restricted distribution in northern of northern Brazil, northern Africa, and Europe and western Gondwana, from the Barremian to the (England, Portugal, and Spain). Therefore, it is nowCenomanian (Table 1). Spinosauridae includes two possible and desirable to place these specimens in a subfamily-level subclades, Spinosaurinae and Baryo- biogeographic context in order to understand their nychinae (sensu Hendrickx et al., 2016) (Figure 1). turnover prior to the Cenomanian. As such, the focus Their evolutionary history dates from the Early Cre- of this review is on the best-known spinosaurids of taceous, and they were mainly distribution in north- North Africa and Europe. ern Gondwana and western Laurasia. The absence of spinosaurids after the Cenomanian is an interesting Several anatomical, taxonomic and systematic observation, with evolution and biogeographic impli- studies of Spinosauridae have been published over cations, which is discussed below. the last few decades (e.g., Bufetaut & Ingavat, 1986; Sereno et al., 1998; Benton et al., 2000; Bufetaut, 2008, 2012; Bensonet al., 2009; Allain 2.1 Europe et al., 2012; Hendrickx et al., 2016). Nevertheless, the biogeographic distribution of this group in The only spinosaurid species thus far Africa and Europe has received much less attention.described from England is Baryonyx walkeri from Accordingly, the aim of this paper is to document the Barremian Weald Clay Formation (Charig & and analyze the fossil record, biogeography and Milner, 1997). The associated material by Charig extinction of spinosaurids in western Europe and & Milner (1986) contains a premaxilla, vomers, an North Africa during Cretaceous Period. anterior portion of the left maxilla, some skull bones, Anuário do Instituto de Geociências - UFRJ ISSN 0101-9759 e-ISSN 1982-3908 - Vol. 40 - 3 / 2017 p. 294-302 295 Spinosaurid Dinosaurs from the Early Cretaceous of North Africa and Europe: Fossil Record, Biogeography and Extinction Carlos Roberto A. Candeiro; Stephen L. Brusatte & André Luis de Souza Figure 1 Phylogenetic relationships of the species of Spinosauridae family from middle Cretaceous (modiied from Hendrickx et al., 2016). Taxa both dentaries, teeth, an axis, most of the cervical Formation Age Country Selected vertebrae, some dorsal vertebrae, one caudal vertebra, Europe References a cervical rib, dorsal ribs, gastralia, chevrons, both Weald Clay Charig & scapulae, a coracoid, both humeri, manual phalanges Baryonyx walker Formation Barremian England Milner (1986) (including the ungual phalanges), an incomplete Charig & ilium, a pubis, an ischium, the proximal end of the Enciso Milner (1997), Baryonyx walker Group Barremian Spain Vieira & left femur and the distal end of the right femur, an Torres (1995) incomplete left ibula, the right calcaneum, the distal Baryonychinae Wessex Barremian England Martill & Hutt ends of the metatarsals, and the pedal phalanges of indet. Formation (1996) both feet (including the ungual ones). Papo Seco Formation Portugal At irst, Charig & Milner (1986) did not Africa consider B. walkeri as belonging to Spinosauridae, Spinosaurus Bahariya Stromer aegyptiacus Formation Cenomanian Egypt (1915) and it was Paul (1988) and Bufetaut (1989a,b) who Russel were the irst authors to associate this species with Spinosaurus Kem Kem early (1996), maroccanus Beds Cenomanian Morocco Sereno et al. Spinosaurus aegyptiacus Stromer, 1915. It is now (1998) widely accepted that Baryonyx is an exemplary Taquet & taxon of spinosaurid. Many authors (e.g., Serenoet Russel Spinosaurus Kem Kem al., 1998; Hendrickx et al., 2016) have considered maroccanus Beds Albian Algeria (1998), Sereno et al. Baryonyx as belonging to a spinosaurid subclade, (1998) called Baryonychinae, which also includes Taquet & Cristatusaurus Russel Suchomimus tenerensis. lapparenti Elrhaz Aptian Niger (1998), nomen nudum Formation Sereno et al. (1998) There is also additional material of Sereno et Baryonyx, in addition to the type specimen from
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  • DOI: 10.1126/Science.282.5392.1298 , 1298 (1998); 282 Science Et Al
    A Long-Snouted Predatory Dinosaur from Africa and the Evolution of Spinosaurids Paul C. Sereno et al. Science 282, 1298 (1998); DOI: 10.1126/science.282.5392.1298 This copy is for your personal, non-commercial use only. If you wish to distribute this article to others, you can order high-quality copies for your colleagues, clients, or customers by clicking here. Permission to republish or repurpose articles or portions of articles can be obtained by following the guidelines here. The following resources related to this article are available online at www.sciencemag.org (this information is current as of September 6, 2013 ): Updated information and services, including high-resolution figures, can be found in the online version of this article at: on September 6, 2013 http://www.sciencemag.org/content/282/5392/1298.full.html This article cites 29 articles, 3 of which can be accessed free: http://www.sciencemag.org/content/282/5392/1298.full.html#ref-list-1 This article has been cited by 90 article(s) on the ISI Web of Science This article has been cited by 15 articles hosted by HighWire Press; see: http://www.sciencemag.org/content/282/5392/1298.full.html#related-urls This article appears in the following subject collections: www.sciencemag.org Geochemistry, Geophysics http://www.sciencemag.org/cgi/collection/geochem_phys Downloaded from Science (print ISSN 0036-8075; online ISSN 1095-9203) is published weekly, except the last week in December, by the American Association for the Advancement of Science, 1200 New York Avenue NW, Washington, DC 20005. Copyright 1998 by the American Association for the Advancement of Science; all rights reserved.
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