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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">A large abelisaurid (Dinosauria, Theropoda) from Morocco and comments on the Cenomanian theropods from North Africa</title><author><name sortKey="Chiarenza, Alfio Alessandro" sort="Chiarenza, Alfio Alessandro" uniqKey="Chiarenza A" first="Alfio Alessandro" last="Chiarenza">Alfio Alessandro Chiarenza</name><affiliation><nlm:aff id="aff-1"><institution>Department of Earth Science and Engineering, Imperial College London</institution>,<addr-line>London</addr-line>,<country>United Kingdom</country></nlm:aff></affiliation></author><author><name sortKey="Cau, Andrea" sort="Cau, Andrea" uniqKey="Cau A" first="Andrea" last="Cau">Andrea Cau</name><affiliation><nlm:aff id="aff-2"><institution>Earth, Life and Environmental Sciences, University of Bologna</institution>,<addr-line>Bologna</addr-line>,<country>Italy</country></nlm:aff></affiliation><affiliation><nlm:aff id="aff-3"><institution>Geological and Palaeontological Museum “G. Capellini,”</institution><addr-line>Bologna</addr-line>,<country>Italy</country></nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">26966675</idno><idno type="pmc">4782726</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4782726</idno><idno type="RBID">PMC:4782726</idno><idno type="doi">10.7717/peerj.1754</idno><date when="2016">2016</date><idno type="wicri:Area/Pmc/Corpus">000039</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">000039</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">A large abelisaurid (Dinosauria, Theropoda) from Morocco and comments on the Cenomanian theropods from North Africa</title><author><name sortKey="Chiarenza, Alfio Alessandro" sort="Chiarenza, Alfio Alessandro" uniqKey="Chiarenza A" first="Alfio Alessandro" last="Chiarenza">Alfio Alessandro Chiarenza</name><affiliation><nlm:aff id="aff-1"><institution>Department of Earth Science and Engineering, Imperial College London</institution>,<addr-line>London</addr-line>,<country>United Kingdom</country></nlm:aff></affiliation></author><author><name sortKey="Cau, Andrea" sort="Cau, Andrea" uniqKey="Cau A" first="Andrea" last="Cau">Andrea Cau</name><affiliation><nlm:aff id="aff-2"><institution>Earth, Life and Environmental Sciences, University of Bologna</institution>,<addr-line>Bologna</addr-line>,<country>Italy</country></nlm:aff></affiliation><affiliation><nlm:aff id="aff-3"><institution>Geological and Palaeontological Museum “G. Capellini,”</institution><addr-line>Bologna</addr-line>,<country>Italy</country></nlm:aff></affiliation></author></analytic><series><title level="j">PeerJ</title><idno type="eISSN">2167-8359</idno><imprint><date when="2016">2016</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p>We describe the partially preserved femur of a large-bodied theropod dinosaur from the Cenomanian “Kem Kem Compound Assemblage” (KKCA) of Morocco. The fossil is housed in the Museo Geologico e Paleontologico “Gaetano Giorgio Gemmellaro” in Palermo (Italy). The specimen is compared with the theropod fossil record from the KKCA and coeval assemblages from North Africa. The combination of a distally reclined head, a not prominent trochanteric shelf, distally placed lesser trochanter of stout, alariform shape, a stocky shaft with the fourth trochanter placed proximally, and rugose muscular insertion areas in the specimen distinguishes it from <italic>Carcharodontosaurus</italic>, <italic>Deltadromeus</italic> and <italic>Spinosaurus</italic> and supports referral to an abelisaurid. The estimated body size for the individual from which this femur was derived is comparable to <italic>Carnotaurus</italic> and <italic>Ekrixinatosaurus</italic> (up to 9 meters in length and 2 tons in body mass). This find confirms that abelisaurids had reached their largest body size in the “middle Cretaceous,” and that large abelisaurids coexisted with other giant theropods in Africa. We review the taxonomic status of the theropods from the Cenomanian of North Africa, and provisionally restrict the Linnean binomina <italic>Carcharodontosaurus iguidensis</italic> and <italic>Spinosaurus aegyptiacus</italic> to the type specimens. Based on comparisons among the theropod records from the Aptian-Cenomanian of South America and Africa, a partial explanation for the so-called “Stromer’s riddle” (namely, the coexistence of many large predatory dinosaurs in the “middle Cretaceous” record from North Africa) is offered in term of taphonomic artifacts among lineage records that were ecologically and environmentally non-overlapping. Although morphofunctional and stratigraphic evidence supports an ecological segregation between spinosaurids and the other lineages, the co-occurrence of abelisaurids and carcharodontosaurids, two groups showing several craniodental convergences that suggest direct resource competition, remains to be explained.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Accarie, H" uniqKey="Accarie H">H Accarie</name></author><author><name sortKey="Beaudoin, B" uniqKey="Beaudoin B">B Beaudoin</name></author><author><name sortKey="Dejax, J" uniqKey="Dejax J">J Dejax</name></author><author><name sortKey="Fries, G" uniqKey="Fries G">G Fries</name></author><author><name sortKey="Michard, Jc" uniqKey="Michard J">JC Michard</name></author><author><name sortKey="Taquet, P" uniqKey="Taquet P">P Taquet</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Allain, R" uniqKey="Allain R">R Allain</name></author><author><name 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<front><journal-meta><journal-id journal-id-type="nlm-ta">PeerJ</journal-id><journal-id journal-id-type="iso-abbrev">PeerJ</journal-id><journal-id journal-id-type="pmc">PeerJ</journal-id><journal-id journal-id-type="publisher-id">PeerJ</journal-id><journal-title-group><journal-title>PeerJ</journal-title></journal-title-group><issn pub-type="epub">2167-8359</issn><publisher><publisher-name>PeerJ Inc.</publisher-name><publisher-loc>San Francisco, USA</publisher-loc></publisher></journal-meta><article-meta><article-id pub-id-type="pmid">26966675</article-id><article-id pub-id-type="pmc">4782726</article-id><article-id pub-id-type="publisher-id">1754</article-id><article-id pub-id-type="doi">10.7717/peerj.1754</article-id><article-categories><subj-group subj-group-type="heading"><subject>Paleontology</subject></subj-group><subj-group subj-group-type="heading"><subject>Zoology</subject></subj-group></article-categories><title-group><article-title>A large abelisaurid (Dinosauria, Theropoda) from Morocco and comments on the Cenomanian theropods from North Africa</article-title></title-group><contrib-group><contrib id="author-1" contrib-type="author"><contrib-id contrib-id-type="orcid" authenticated="false">http://orcid.org/0000-0001-5525-6730</contrib-id><name><surname>Chiarenza</surname><given-names>Alfio Alessandro</given-names></name><xref ref-type="aff" rid="aff-1">1</xref></contrib><contrib id="author-2" contrib-type="author" corresp="yes"><name><surname>Cau</surname><given-names>Andrea</given-names></name><xref ref-type="aff" rid="aff-2">2</xref><xref ref-type="aff" rid="aff-3">3</xref><email>cauand@gmail.com</email></contrib><aff id="aff-1"><label>1</label><institution>Department of Earth Science and Engineering, Imperial College London</institution>,<addr-line>London</addr-line>,<country>United Kingdom</country></aff><aff id="aff-2"><label>2</label><institution>Earth, Life and Environmental Sciences, University of Bologna</institution>,<addr-line>Bologna</addr-line>,<country>Italy</country></aff><aff id="aff-3"><label>3</label><institution>Geological and Palaeontological Museum “G. Capellini,”</institution><addr-line>Bologna</addr-line>,<country>Italy</country></aff></contrib-group><contrib-group><contrib contrib-type="editor"><name><surname>Sues</surname><given-names>Hans-Dieter</given-names></name></contrib></contrib-group><pub-date pub-type="epub" date-type="pub" iso-8601-date="2016-02-29"><day>29</day><month>2</month><year iso-8601-date="2016">2016</year></pub-date><pub-date pub-type="collection"><year>2016</year></pub-date><volume>4</volume><elocation-id>e1754</elocation-id><history><date date-type="received" iso-8601-date="2016-01-13"><day>13</day><month>1</month><year iso-8601-date="2016">2016</year></date><date date-type="accepted" iso-8601-date="2016-02-12"><day>12</day><month>2</month><year iso-8601-date="2016">2016</year></date></history><permissions><copyright-statement>© 2016 Chiarenza & Cau</copyright-statement><copyright-year>2016</copyright-year><copyright-holder>Chiarenza & Cau</copyright-holder><license xlink:href="http://creativecommons.org/licenses/by/4.0/"><license-p>This is an open access article distributed under the terms of the <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/4.0/">Creative Commons Attribution License</ext-link>, which permits unrestricted use, distribution, reproduction and adaptation in any medium and for any purpose provided that it is properly attributed. For attribution, the original author(s), title, publication source (PeerJ) and either DOI or URL of the article must be cited.</license-p></license></permissions><self-uri xlink:href="https://peerj.com/articles/1754"></self-uri><abstract><p>We describe the partially preserved femur of a large-bodied theropod dinosaur from the Cenomanian “Kem Kem Compound Assemblage” (KKCA) of Morocco. The fossil is housed in the Museo Geologico e Paleontologico “Gaetano Giorgio Gemmellaro” in Palermo (Italy). The specimen is compared with the theropod fossil record from the KKCA and coeval assemblages from North Africa. The combination of a distally reclined head, a not prominent trochanteric shelf, distally placed lesser trochanter of stout, alariform shape, a stocky shaft with the fourth trochanter placed proximally, and rugose muscular insertion areas in the specimen distinguishes it from <italic>Carcharodontosaurus</italic>, <italic>Deltadromeus</italic> and <italic>Spinosaurus</italic> and supports referral to an abelisaurid. The estimated body size for the individual from which this femur was derived is comparable to <italic>Carnotaurus</italic> and <italic>Ekrixinatosaurus</italic> (up to 9 meters in length and 2 tons in body mass). This find confirms that abelisaurids had reached their largest body size in the “middle Cretaceous,” and that large abelisaurids coexisted with other giant theropods in Africa. We review the taxonomic status of the theropods from the Cenomanian of North Africa, and provisionally restrict the Linnean binomina <italic>Carcharodontosaurus iguidensis</italic> and <italic>Spinosaurus aegyptiacus</italic> to the type specimens. Based on comparisons among the theropod records from the Aptian-Cenomanian of South America and Africa, a partial explanation for the so-called “Stromer’s riddle” (namely, the coexistence of many large predatory dinosaurs in the “middle Cretaceous” record from North Africa) is offered in term of taphonomic artifacts among lineage records that were ecologically and environmentally non-overlapping. Although morphofunctional and stratigraphic evidence supports an ecological segregation between spinosaurids and the other lineages, the co-occurrence of abelisaurids and carcharodontosaurids, two groups showing several craniodental convergences that suggest direct resource competition, remains to be explained.</p></abstract><kwd-group kwd-group-type="author"><kwd>Cenomanian</kwd><kwd>Morocco</kwd><kwd>Theropoda</kwd></kwd-group><funding-group><funding-statement>The authors received no funding for this work.</funding-statement></funding-group></article-meta></front><body><sec sec-type="intro"><title>Introduction</title><p>The dinosaurs from the Aptian-Cenomanian of North Africa are mainly known from a few articulated skeletons and several isolated bones, the majority of which are referred to medium- to large-sized theropod clades (i.e., Abelisauroidea, Carcharodontosauridae, Spinosauridae; <xref rid="ref-66" ref-type="bibr">Stromer, 1915</xref>; <xref rid="ref-67" ref-type="bibr">Stromer, 1931</xref>; <xref rid="ref-68" ref-type="bibr">Stromer, 1934</xref>; <xref rid="ref-57" ref-type="bibr">Russell, 1996</xref>; <xref rid="ref-63" ref-type="bibr">Sereno et al., 1996</xref>; <xref rid="ref-24" ref-type="bibr">Dal Sasso et al., 2005</xref>; <xref rid="ref-45" ref-type="bibr">Mahler, 2005</xref>; <xref rid="ref-6" ref-type="bibr">Brusatte & Sereno, 2007</xref>; <xref rid="ref-62" ref-type="bibr">Sereno & Brusatte, 2008</xref>; <xref rid="ref-65" ref-type="bibr">Smith et al., 2010</xref>; <xref rid="ref-18" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2012</xref>; <xref rid="ref-19" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2013</xref>; <xref rid="ref-36" ref-type="bibr">Ibrahim et al., 2014</xref>; <xref rid="ref-29" ref-type="bibr">Evers et al., 2015</xref>; <xref rid="ref-32" ref-type="bibr">Hendrickx, Mateus & Buffetaut, 2016</xref>). Whether the abundance of large theropods compared to other dinosaurs reflects a real ecological signal (i.e., an unusually unbalanced ecosystem; <xref rid="ref-40" ref-type="bibr">Läng et al., 2013</xref>) or a preservational, taphonomic or collecting biases (<xref rid="ref-49" ref-type="bibr">McGowan & Dyke, 2009</xref>) is still to be assessed. Here we describe an additional fossil specimen, adding further information on the known diversity of large-bodied African theropods. The fossil comes from the region of Taouz (Errachidia Province, Morocco, near the Moroccan-Algerian border) and was donated in 2005 to the Museo Geologico e Paleontologico “Gaetano Giorgio Gemmellaro” in Palermo (Italy) by a donor who had purchased it from a Moroccan fossil dealer. Many dinosaurian remains have been collected from the Tafilalt and Kem Kem regions (SE Morocco) by local inhabitants and fossil dealers and deposited in public institutions all over the world (<xref rid="ref-49" ref-type="bibr">McGowan & Dyke, 2009</xref>). As is usually the case (e.g. <xref rid="ref-28" ref-type="bibr">Evans et al., 2015</xref>; <xref rid="ref-18" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2012</xref>; <xref rid="ref-32" ref-type="bibr">Hendrickx, Mateus & Buffetaut, 2016</xref>), this specimen was found by local people, and its exact horizon and locality is unknown. On the other hand, some information may be gleaned from the most recent and exhaustive review on the sedimentary geology of the Late Cretaceous North Africa dinosaur-rich units, also known as “Kem Kem Compound Assemblage” (KKCA <italic>sensu</italic><xref rid="ref-20" ref-type="bibr">Cavin et al., 2010</xref>). These units are represented by the Ifezouane Formation and the overlying Aoufous Formation (<xref rid="ref-20" ref-type="bibr">Cavin et al., 2010</xref>), which are Cenomanian in age, and have been deposited along the south-western Tethyan margin before the late Cenomanian global marine transgression, represented in this region by the limestone unit of the Akrabou Formation (<xref rid="ref-20" ref-type="bibr">Cavin et al., 2010</xref>). The units included in the KKCA are the only dinosaur-bearing levels in the region of Taouz (<xref rid="ref-20" ref-type="bibr">Cavin et al., 2010</xref>). The matrix still encrusting the specimen (i.e., a consolidated red sandstone) closely recalls that present in other dinosaur fossils from the KKCA (e.g., <xref rid="ref-19" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2013</xref>; <xref rid="ref-32" ref-type="bibr">Hendrickx, Mateus & Buffetaut, 2016</xref>; personal observations on material housed in the Natural History Museum in Milan; see <xref rid="ref-36" ref-type="bibr">Ibrahim et al., 2014</xref>). Based on its documented provenance and the lithological features mentioned above, we thus refer the fossil to the KKCA. In this study, we describe this specimen, compare it to other North African theropods, assess its phyletic relationships, and infer its body size.</p></sec><sec><title>Abbreviations</title><p>KKCA, Kem Kem Compound Assemblage; OLPH, Olphin collection of the Museo Geologico e Paleontologico “Gaetano Giorgio Gemmellaro,” Università degli Studi di Palermo, Palermo, Sicily, Italy; NMC, Canadian Museum of Nature, formerly National Museum of Canada, Ottawa, Canada; ROM, Royal Ontario Museum, Toronto, Canada; SGM, Ministère de l’Énergie et des Mines, Rabat, Morocco.</p></sec><sec><title>Systematic Palaeontology</title><list list-type="simple" id="list-1"><list-item><p>Dinosauria <xref rid="ref-70" ref-type="bibr">Owen (1842)</xref>.</p></list-item><list-item><p>Theropoda <xref rid="ref-71" ref-type="bibr">Marsh (1881)</xref>.</p></list-item><list-item><p>Abelisauridae <xref rid="ref-72" ref-type="bibr">Bonaparte (1991)</xref>.</p></list-item></list><sec><title>Locality and age</title><p>Based on the registry of the OLPH, the specimen was collected nearby the Moroccan-Algerian boundary just south of Taouz (Errachidia Province, Meknès−Tafilalet Region), Morocco. Following <xref rid="ref-20" ref-type="bibr">Cavin et al. (2010)</xref>, the age of this fossil is considered as Late Cretaceous (Cenomanian).</p></sec><sec><title>Material</title><p>OLPH 025, partial proximal portion of a right femur (<xref ref-type="fig" rid="fig-1">Fig. 1</xref>).</p><fig id="fig-1" orientation="portrait" position="float"><object-id pub-id-type="doi">10.7717/peerj.1754/fig-1</object-id><label>Figure 1</label><caption><title>Abelisauridae indet. femur OLPH 025.</title><p>(A) proximal view, (B) anterior view, (C) medial view, (D) posterior view, (E) lateral view, (F) distal view (not at same scale as other views). Scale bars, 5 cm. Abbreviations: gt, greater trochanter; iMie, insertion for the <italic>M. iliofemoralis externus</italic>; fn, femoral neck; s, shallow sulcus.</p></caption><graphic xlink:href="peerj-04-1754-g001"></graphic></fig></sec><sec><title>Description and Comparison</title><p>Measurements for the specimen are included in <xref ref-type="table" rid="table-1">Table 1</xref>. OLPH 025 is the proximal end of a femur, including the head, neck and trochanteric region. The preserved shaft is sigmoid in both anterior and posterior views (<xref ref-type="fig" rid="fig-1">Fig. 1</xref>), as in <italic>Berberosaurus liassicus</italic> (<xref rid="ref-2" ref-type="bibr">Allain et al., 2007</xref>) and <italic>Majungasaurus crenatissimus</italic> (<xref rid="ref-12" ref-type="bibr">Carrano, 2007</xref>), and differs from the straighter shape in <italic>Carcharodontosaurus saharicus</italic> (<xref rid="ref-67" ref-type="bibr">Stromer, 1931</xref>). The femoral head (<xref ref-type="fig" rid="fig-1">Fig. 1A</xref>) is anteroposteriorly compressed, subcircular in medial view (<xref ref-type="fig" rid="fig-1">Fig. 1C</xref>), and has a narrow neck that curves anteriorly, placing the head anteromedially in proximal view, similar to the condition in <italic>Carnotaurus</italic>, <italic>Ekrixinatosaurus</italic>, <italic>Rahiolisaurus</italic>, <italic>Xenotarsosaurus</italic> and all other non-tetanuran theropods (<xref rid="ref-5" ref-type="bibr">Bonaparte, Novas & Coria, 1990</xref>; <xref rid="ref-52" ref-type="bibr">Novas et al., 2010</xref>). In anterior view (<xref ref-type="fig" rid="fig-1">Fig. 1B</xref>), the dorsal margin of the femoral head is angled slightly distally rather than mainly perpendicular to the shaft, recalling <italic>Masiakasaurus</italic> and abelisaurids (<xref rid="ref-14" ref-type="bibr">Carrano, Sampson & Forster, 2002</xref>; <xref rid="ref-15" ref-type="bibr">Carrano, Wilson & Barrett, 2010</xref>; <xref rid="ref-12" ref-type="bibr">Carrano, 2007</xref>; <xref rid="ref-28" ref-type="bibr">Evans et al., 2015</xref>), whereas in <italic>Carcharodontosaurus saharicus</italic> and <italic>Deltadromeus agilis</italic> the head projects considerably proximally (<xref rid="ref-67" ref-type="bibr">Stromer, 1931</xref>; <xref rid="ref-28" ref-type="bibr">Evans et al., 2015</xref>). The lesser trochanter is broad anteroposteriorly and anteriorly projected, as in <italic>Ceratosaurus, Masiakasaurus</italic>, abelisaurids and basal tetanurans, set apart from the femoral head by a shallow sulcus as in <italic>Ceratosaurus</italic> and <italic>Berberosaurus</italic> and unlike the wide and deep cleft present in <italic>Carcharodontosaurus saharicus</italic> (<xref rid="ref-67" ref-type="bibr">Stromer, 1931</xref>). The lesser trochanter is positioned distally relative to the articular end, approaching proximally the level of the base of the head, differing from the more proximally placed trochanter present in <italic>Deltadromeus</italic> and most tetanurans (<xref rid="ref-43" ref-type="bibr">Madsen, 1976</xref>; <xref rid="ref-28" ref-type="bibr">Evans et al., 2015</xref>). The distal placement of the lesser trochanter is a plesiomorphic condition shared by coelophysoid-grade theropods (e.g., <italic>Sarcosaurus</italic>, <xref rid="ref-4" ref-type="bibr">Andrews, 1921</xref>), ceratosaurids (<xref rid="ref-44" ref-type="bibr">Madsen & Welles, 2000</xref>), and abelisauroids (<xref rid="ref-5" ref-type="bibr">Bonaparte, Novas & Coria, 1990</xref>; <xref rid="ref-41" ref-type="bibr">Le Loeuff & Buffetaut, 1991</xref>; <xref rid="ref-1" ref-type="bibr">Accarie et al., 1995</xref>; <xref rid="ref-46" ref-type="bibr">Martínez & Novas, 1997</xref>; <xref rid="ref-14" ref-type="bibr">Carrano, Sampson & Forster, 2002</xref>; <xref rid="ref-12" ref-type="bibr">Carrano, 2007</xref>). There is no evidence of a trochanteric shelf, although the posterolateral surface of the shaft at the level of the lesser trochanter appears damaged, so that any trace of even a faint trochanteric shelf (as in <italic>Majungasaurus</italic>; <xref rid="ref-12" ref-type="bibr">Carrano, 2007</xref>) may have been obliterated by erosion. Similar to <italic>Berberosaurus</italic> and <italic>Majungasaurus</italic> (<xref rid="ref-2" ref-type="bibr">Allain et al., 2007</xref>; <xref rid="ref-12" ref-type="bibr">Carrano, 2007</xref>), OLPH 025 does not show any evidence of the accessory trochanter, a feature widely present among neotetanuran theropods (<xref rid="ref-35" ref-type="bibr">Hutchinson, 2001</xref>) and illustrated on a femur referred to <italic>Bahariasaurus</italic> by <xref rid="ref-68" ref-type="bibr">Stromer (1934)</xref> and to <italic>Deltadromeus</italic> by <xref rid="ref-63" ref-type="bibr">Sereno et al. (1996)</xref>. The anterior margin of the lesser trochanter bears a mound-like rugosity, interpreted as the insertion for the <italic>M. iliofemoralis externus</italic> (<xref rid="ref-35" ref-type="bibr">Hutchinson, 2001</xref>; <xref rid="ref-12" ref-type="bibr">Carrano, 2007</xref>). The distal (apical) and lateral surface of the lesser trochanter is extremely rugose, as in <italic>Majungasaurus</italic> (<xref rid="ref-12" ref-type="bibr">Carrano, 2007</xref>). In posterior view (<xref ref-type="fig" rid="fig-1">Fig. 1D</xref>), toward the distal surface of the femur, a thin crista, proximodistally oriented, is set closer to the medial margin of the femur, extending gradually from the bone surface and oriented subparallel to the proximodistal axis of the diaphysis. This crest is interpreted as the proximal end of the ridge-like fourth trochanter. As in <italic>Ceratosaurus</italic> and abelisauroids (e.g., <xref rid="ref-44" ref-type="bibr">Madsen & Welles, 2000</xref>; <xref rid="ref-12" ref-type="bibr">Carrano, 2007</xref>), the fourth trochanter is placed more proximally than in tetanurans (e.g., <italic>Allosaurus</italic>, <xref rid="ref-43" ref-type="bibr">Madsen, 1976, plate 50</xref>). The fourth trochanter is more medially than centrally set along the posterior surface, as in <italic>Ceratosaurus</italic> (<xref rid="ref-44" ref-type="bibr">Madsen & Welles, 2000</xref>). In proximal view, the femur head appears “kidney-shaped” with the lesser trochanter barely visible on the anteromedial corner, differing from the condition in tetanurans and noasaurids, where the lesser trochanter is more widely exposed in proximal view (e.g., <italic>Allosaurus fragilis</italic>, personal observations; <italic>Masiakasaurus</italic>, <xref rid="ref-14" ref-type="bibr">Carrano, Sampson & Forster, 2002</xref>). In distal view (<xref ref-type="fig" rid="fig-1">Fig. 1F</xref>), the femoral shaft is slightly more anteroposteriorly compressed, with an approximately triangular to rhomboidal outline in section at the level of the fourth trochanter, and with the apex pointing anteriorly, as in <italic>Ceratosaurus, Masiakasaurus,</italic> and abelisaurids (<xref rid="ref-44" ref-type="bibr">Madsen & Welles, 2000</xref>; <xref rid="ref-12" ref-type="bibr">Carrano, 2007</xref>; <xref rid="ref-16" ref-type="bibr">Carrano, Loewen & Sertich, 2011</xref>). This shape differs from the more rounded cross-section of tetanuran femora (e.g., <xref rid="ref-43" ref-type="bibr">Madsen, 1976, fig. 24B</xref>). As in the vast majority of theropods, but differing from a KKCA femur referred to <italic>Spinosaurus</italic> by <xref rid="ref-36" ref-type="bibr">Ibrahim et al. (2014)</xref>, the medullary cavity is large (using the better preserved anteromedial quarter of the section, the radius of the medullary cavity is about half the length of both principal section axes).</p><table-wrap id="table-1" orientation="portrait" position="float"><object-id pub-id-type="doi">10.7717/peerj.1754/table-1</object-id><label>Table 1</label><caption><title>Selected measurements (in mm) of OLPH 025.</title></caption><alternatives><graphic xlink:href="peerj-04-1754-g004"></graphic><table frame="hsides" rules="groups"><colgroup span="1"><col span="1"></col><col span="1"></col></colgroup><tbody><tr><td rowspan="1" colspan="1">Proximal surface, anteroposterior length from acetabular surface to greater trochanter</td><td rowspan="1" colspan="1">170</td></tr><tr><td rowspan="1" colspan="1">Proximal surface, minimum transverse width at mid-length</td><td rowspan="1" colspan="1">100</td></tr><tr><td rowspan="1" colspan="1">Anterior view, maximum proximodistal length of preserved bone</td><td rowspan="1" colspan="1">330</td></tr><tr><td rowspan="1" colspan="1">Anterior view, proximodistal depth of articular surface</td><td rowspan="1" colspan="1">95</td></tr><tr><td rowspan="1" colspan="1">Head, articular surface anteroposterior diameter</td><td rowspan="1" colspan="1">150</td></tr><tr><td rowspan="1" colspan="1">Greater trochanter, maximum anteroposterior diameter</td><td rowspan="1" colspan="1">90</td></tr><tr><td rowspan="1" colspan="1">Greater trochanter, proximodistal depth above lesser trochanter base</td><td rowspan="1" colspan="1">105</td></tr><tr><td rowspan="1" colspan="1">Shaft, preserved distal surface, anteroposterior diameter vs preserved width</td><td rowspan="1" colspan="1">120 × 90</td></tr></tbody></table></alternatives></table-wrap></sec></sec><sec><title>Results and Discussion</title><sec><title>Taxonomy and inclusiveness of the KKCA theropod taxa</title><p>Most African theropod taxa are based on isolated material, often single bones, or include referred material that in many cases lacks overlapping elements with the type specimens (e.g., <italic>Kryptops palaios</italic>, <xref rid="ref-62" ref-type="bibr">Sereno & Brusatte, 2008</xref>; see discussion in <xref rid="ref-73" ref-type="bibr">Carrano, Benson & Sampson (2012)</xref>; <italic>Eocarcharia dinops</italic>, <xref rid="ref-62" ref-type="bibr">Sereno & Brusatte, 2008</xref>). Since referral of isolated and non-overlapping specimens to the same taxon is a hypothesis itself, we briefly review here the taxonomic status of the known theropod taxa from the KKCA and–where relevant to the discussion–from penecontemporaneous assemblages from North Africa (<xref ref-type="fig" rid="fig-2">Fig. 2</xref>).</p><fig id="fig-2" orientation="portrait" position="float"><object-id pub-id-type="doi">10.7717/peerj.1754/fig-2</object-id><label>Figure 2</label><caption><title>Main theropod faunal assemblages from the Aptian-Cenomanian of North Africa.</title><p>Taxa enclosed in rectangles have been considered as synonyms by some authors and distinct by others (see <xref rid="ref-57" ref-type="bibr">Russell, 1996</xref>; <xref rid="ref-61" ref-type="bibr">Sereno, Wilson & Conrad, 2004</xref>; <xref rid="ref-19" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2013</xref>; <xref rid="ref-36" ref-type="bibr">Ibrahim et al., 2014</xref>; <xref rid="ref-29" ref-type="bibr">Evers et al., 2015</xref>).</p></caption><graphic xlink:href="peerj-04-1754-g002"></graphic></fig></sec><sec><title>Carcharodontosauridae</title><p>The original types of “<italic>Megalosaurus saharicus</italic>” were two isolated teeth from the Late Cretaceous “Continental Intercalaire” units of Algeria (<xref rid="ref-25" ref-type="bibr">Depéret & Savornin, 1925</xref>; see discussion by <xref rid="ref-6" ref-type="bibr">Brusatte & Sereno (2007)</xref>). The genoholotype of <italic>Carcharodontosaurus</italic> is based on a partial skeleton from the Baharjie assemblage of Egypt that includes teeth comparable to those of “<italic>M. saharicus</italic>,” and, among other elements, a well-preserved femur (<xref rid="ref-67" ref-type="bibr">Stromer, 1931</xref>; <xref ref-type="fig" rid="fig-3">Fig. 3C</xref>). That material (and all other theropod bones described by <xref rid="ref-66" ref-type="bibr">Stromer (1915)</xref>, <xref rid="ref-67" ref-type="bibr">Stromer (1931)</xref> and <xref rid="ref-68" ref-type="bibr">Stromer (1934)</xref>) was destroyed during World War II. <xref rid="ref-6" ref-type="bibr">Brusatte & Sereno (2007)</xref> designated a partial skull from the Cenomanian of the KKCA (see <xref rid="ref-63" ref-type="bibr">Sereno et al., 1996</xref>) as the neotype of <italic>Carcharodontosaurus saharicus</italic>. This material lacks a femur, preventing direct comparison with the Palermo specimen. Although in overall morphology the neotype of <italic>C. saharicus</italic> (<xref rid="ref-63" ref-type="bibr">Sereno et al., 1996</xref>; <xref rid="ref-6" ref-type="bibr">Brusatte & Sereno, 2007</xref>) closely matches the overlapping cranial material of the destroyed Egyptian specimen (<xref rid="ref-67" ref-type="bibr">Stromer, 1931</xref>), the two specimens differ in the shape of the maxillary interdental plates, that are quadrangular in medial view and apically flattened in the Moroccan specimen (<xref rid="ref-6" ref-type="bibr">Brusatte & Sereno, 2007, fig. 2</xref>; <xref rid="ref-31" ref-type="bibr">Hendrickx & Mateus, 2014, supplementary information</xref>), whereas are depicted as subtriangular in medial view and apically pointed in the Egyptian specimen (<xref rid="ref-68" ref-type="bibr">Stromer, 1934, plate 1, fig. 6a</xref>). This difference may be taxonomically significant because it also differentiates the holotype of <italic>Carcharodontosaurus iguidensis</italic> from the neotype of <italic>Carcharodontosaurus saharicus</italic> (<xref rid="ref-6" ref-type="bibr">Brusatte & Sereno, 2007, fig. 2</xref>), and is a phylogenetically informative feature among theropod species (see <xref rid="ref-31" ref-type="bibr">Hendrickx & Mateus, 2014, supplementary information</xref>). The type material of <italic>Carcharodontosaurus iguidensis</italic> includes an isolated maxilla from the Echkar Formation of Niger (<xref rid="ref-6" ref-type="bibr">Brusatte & Sereno, 2007</xref>). The referred material (partial skull and vertebrae) was discovered three kilometers away from the type maxilla and lacks overlapping elements with the latter (<xref rid="ref-6" ref-type="bibr">Brusatte & Sereno, 2007</xref>). <xref rid="ref-6" ref-type="bibr">Brusatte & Sereno (2007: 905)</xref> referred isolated bones from the Echkar Formation to <italic>C. iguidensis</italic> because they “closely match the morphology of <italic>C. saharicus</italic> and because it is unlikely that there would be more than three contemporaneous large-bodied carnivores in the same formation (<italic>Rugops primus</italic>, <italic>Spinosaurus</italic> sp., <italic>Carcharodontosaurus iguidensis</italic>).” We see no reason why the number of large-bodied carnivores in a geological formation should be limited to three, or to refer all carcharodontosaurid specimens from the same formation to a single species when no overlapping material is available (see <xref rid="ref-19" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2013</xref> and reference therein). This raises doubts about the referral of that material to <italic>C. iguidensis</italic>. In particular, the referred material of <italic>C. iguidensis</italic> includes vertebrae referable to the spinosaurid <italic>Sigilmassasaurus</italic> or a closely related taxon (<xref rid="ref-48" ref-type="bibr">McFeeters et al., 2013</xref>; <xref rid="ref-29" ref-type="bibr">Evers et al., 2015</xref>), indicating that it represents a multitaxic association. Among the material referred to <italic>C. iguidensis</italic>, a dentary and braincase were discovered in situ embedded in sandstone of the Echkar Formation and closely associated in a small area (<xref rid="ref-6" ref-type="bibr">Brusatte & Sereno, 2007</xref>), supporting their referral to a single individual. This material shares synapomorphies of Carcharodontosauridae (<xref rid="ref-6" ref-type="bibr">Brusatte & Sereno, 2007</xref>) but lacks synapomorphies of the subclade Carcharodontosaurinae present in both <italic>Carcharodontosaurus</italic> and <italic>Giganotosaurus</italic> (<xref rid="ref-22" ref-type="bibr">Coria & Currie, 2002</xref>): the thickened lacrimal facet of frontal, the invaginated anteromedial corner of the supratemporal fossa, and the exit of the trigeminal foramen posterior to the nuchal crest.</p><fig id="fig-3" orientation="portrait" position="float"><object-id pub-id-type="doi">10.7717/peerj.1754/fig-3</object-id><label>Figure 3</label><caption><title>Theropod femora from the Cenomanian of Egypt and the ‘Kem Kem Compound Assemblage’.</title><p>OLPH 025 in anterior (A) posterior (B) and distal (L) views; scale bar 5 cm. (C) <italic>Carcharodontosaurus saharicus</italic> femur in anterolateral view (re-drawn from <xref rid="ref-67" ref-type="bibr">Stromer, 1931, table I</xref>), scale bar, 5 cm. (D) and (E) cf. <italic>Baharisaurus ingens</italic> femur, referred to <italic>Deltadromeus</italic> by <xref rid="ref-63" ref-type="bibr">Sereno et al. (1996)</xref>, re-drawn from <xref rid="ref-68" ref-type="bibr">Stromer (1934, table III)</xref> in anterior (D) and lateral (E) views; scale bar, 5 cm. (G) left femur of a theropod (ROM 64666, reversed from right) referred to <italic>Deltadromeus agilis</italic> by <xref rid="ref-28" ref-type="bibr">Evans et al. (2015)</xref>, in anterior view; scale bar, 1 cm. (H) left type femur of <italic>Deltadromeus agilis</italic> (SGM Din-2, reversed from right) in posterior view; scale bar, 5 cm. NMC 41869, a right femur referred to <xref rid="ref-57" ref-type="bibr">Russell (1996)</xref> to Theropoda indet. (“bone taxon M”) in (I) anterior, (J) distal, (K) posterior views; scale bar, 5 cm.</p></caption><graphic xlink:href="peerj-04-1754-g003"></graphic></fig><p>As stated by <xref rid="ref-6" ref-type="bibr">Brusatte & Sereno (2007)</xref>, the braincase referred to <italic>C. iguidensis</italic> shows the facet for contact with the prefrontal-lacrimal on the frontal is shallower than in both <italic>C. saharicus</italic> and <italic>Giganotosaurus</italic> (note that, in derived carcharodontosaurids, the prefrontal is reduced and fused to the lacrimal; therefore, the lacrimal facet of frontal in carcharodontosaurids is homologous to the prefrontal facet of basal allosauroids, <xref rid="ref-62" ref-type="bibr">Sereno & Brusatte (2008)</xref>). This feature was listed by <xref rid="ref-6" ref-type="bibr">Brusatte & Sereno (2007)</xref> among the three features differentiating the frontal of <italic>C. iguidensis</italic> from that of <italic>C. saharicus</italic>, “the latter probably exhibiting the derived condition” (<xref rid="ref-6" ref-type="bibr">Brusatte & Sereno, 2007: 907</xref>). Thus, <xref rid="ref-6" ref-type="bibr">Brusatte & Sereno (2007)</xref> implicitly noted that <italic>C. iguidensis</italic> shows the plesiomorphic condition compared to <italic>C. saharicus</italic>. In particular, the lacrimal facet in the neotype frontal of <italic>C. saharicus</italic> is 65 mm deep, about 40% the length of the frontal (a bone stated by <xref rid="ref-6" ref-type="bibr">Brusatte & Sereno (2007)</xref>, to be identical in length to the 150 mm long frontal of <italic>C. iguidensis</italic>). In the frontal referred to <italic>C. iguidensis</italic>, the same facet is reported to be 35 mm deep (<xref rid="ref-6" ref-type="bibr">Brusatte & Sereno, 2007: 908</xref>), about 23% the length of the bone. <xref rid="ref-22" ref-type="bibr">Coria & Currie (2002)</xref> reported that on the 200 mm long frontal of <italic>Giganotosaurus carolinii</italic> holotype, the prefrontal [-lacrimal] facet is 67.5 mm deep, about 33% the length of the frontal. In the braincase of <italic>Acrocanthosaurus atokensis</italic> described by <xref rid="ref-27" ref-type="bibr">Eddy & Clarke (2011)</xref>, the depth of the prefrontal facet of the frontal is about 40–45 mm deep (<xref rid="ref-27" ref-type="bibr">Eddy & Clarke, 2011, fig. 12</xref>), about 23% the length of the frontal (<xref rid="ref-27" ref-type="bibr">Eddy & Clarke, 2011, table 1</xref>). Note that the latter is the same value as for the frontal referred to <italic>C. iguidensis</italic>. In <italic>Shaochilong maortuensis</italic>, the depth of the same facet is 25% the length of the frontal (based on measurements provided by <xref rid="ref-69" ref-type="bibr">Brusatte et al. (2010)</xref>). In more basal allosauroids, the depth of the prefrontal facet of frontal is about 20–25% the length of the bone (e.g., <italic>Sinraptor dongi</italic>, see <xref rid="ref-23" ref-type="bibr">Currie & Zhao, 1994, figs. 7B–7D</xref>). Therefore, <italic>C. saharicus</italic> and <italic>Giganotosaurus</italic> share a prefrontal-lacrimal facet that is more than 30% the length of the frontal, and this derived feature may represent a synapomorphy of Carcharodontosaurinae absent in the frontals of other allosauroids, including that referred to <italic>C. iguidensis.</italic> Although a deep lacrimal facet of frontal is present also in <italic>Sauroniops</italic> (<xref rid="ref-18" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2012</xref>; <xref rid="ref-19" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2013</xref>), this feature is probably not homologous to the condition in other carcharodontosaurids because in the latter the facet is thickest in its posterior margin, not along its anterior margin, as in <italic>Eocarcharia</italic> and <italic>Sauroniops</italic> (<xref rid="ref-62" ref-type="bibr">Sereno & Brusatte, 2008</xref>; <xref rid="ref-19" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2013</xref>).</p><p>Furthermore, <xref rid="ref-6" ref-type="bibr">Brusatte & Sereno (2007)</xref> reported that “the anteromedial corner of the supratemporal fossa is deeply invaginated in <italic>C. saharicus</italic>, but forms a near vertical, broadly arched wall in [the braincase referred to] <italic>C. iguidensis</italic>” (<xref rid="ref-6" ref-type="bibr">Brusatte & Sereno, 2007: 908</xref>). <italic>Carcharodontosaurus saharicus</italic> shows the derived condition, which is due to the extensive development of a medial shelf overlapping the anteromedial corner of the supratemporal fossa (<xref rid="ref-22" ref-type="bibr">Coria & Currie, 2002</xref>). The latter feature is only shared by <italic>Giganotosaurus carolinii</italic> among allosauroids (<xref rid="ref-22" ref-type="bibr">Coria & Currie, 2002</xref>), including other carcharodontosaurids (<xref rid="ref-62" ref-type="bibr">Sereno & Brusatte, 2008</xref>; <xref rid="ref-69" ref-type="bibr">Brusatte et al., 2010</xref>; <xref rid="ref-27" ref-type="bibr">Eddy & Clarke, 2011</xref>), and is thus interpreted as a synapomorphy of Carcharodontosaurinae. Although a medial shelf is incipiently developed in other carcharodontosaurids (e.g., <italic>Acrocanthosaurus</italic>, <xref rid="ref-22" ref-type="bibr">Coria & Currie, 2002</xref>), only <italic>C. saharicus</italic> and <italic>Giganotosaurus</italic> show a deeply invaginated anteromedial corner of the supratemporal fossa due to the extreme development of the shelves. The absence of the invaginated anteromedial corner of the supratemporal fossa on the braincase from Niger is an additional feature challenging its referral to a species of Carcharodontosaurinae.</p><p>In their phylogenetic analysis of Allosauroidea, <xref rid="ref-7" ref-type="bibr">Brusatte & Sereno (2008)</xref> used the position of the trigeminal foramen exit in the braincase relative to the nuchal crest as a phylogenetically informative character, and defined the states as: “braincase, trigeminal (nerve V) foramen, location relative to nuchal crest: anterior or ventral (0); posterior (1) (<xref rid="ref-7" ref-type="bibr">Brusatte & Sereno, 2008: 26</xref>).” According to this character statement, the braincase referred to <italic>C. iguidensis</italic> should be scored as “0,” as its trigeminal foramen is reported to be ventral to the nuchal crest (<xref rid="ref-6" ref-type="bibr">Brusatte & Sereno, 2007: 910</xref>), as in <italic>Sinraptor</italic>, and not “1,” as in <italic>C. saharicus</italic>, <italic>Giganotosaurus</italic> and <italic>Shaochilong</italic> (<xref rid="ref-69" ref-type="bibr">Brusatte et al., 2010</xref>), the latter three showing a more posteriorly placed foramen. Therefore, according to <xref rid="ref-7" ref-type="bibr">Brusatte & Sereno (2008)</xref>, the position of the trigeminal foramen in the braincase referred to <italic>C. iguidensis</italic> is plesiomorphic relative to the conditions in both <italic>C. saharicus</italic> and <italic>Giganotosaurus</italic>, further challenging the referral of that specimen (regardless to the placement of the taxa <italic>C. iguidensis</italic>, based on the type maxilla, and <italic>Shaochilong</italic>) to Carcharodontosaurinae.</p><p>Therefore, the braincase referred to <italic>C. iguidensis</italic> shows a combination of features intermediate between carcharodontosaurine (e.g., <xref rid="ref-22" ref-type="bibr">Coria & Currie, 2002</xref>) and non-carcharodontosaurine (e.g., <xref rid="ref-27" ref-type="bibr">Eddy & Clarke, 2011</xref>) carcharodontosaurids. Some of these features were considered by <xref rid="ref-6" ref-type="bibr">Brusatte & Sereno (2007)</xref> autapomorphies of <italic>C. iguidensis</italic>, and thus, accepting the referral of the braincase to the latter taxon, should be considered reversals to the non-carcharodontosaurine (plesiomorphic) condition. We cannot dismiss that some of these differences between the Nigerine braincase and the carcharodontosaurines are ontogenetic in nature (implying that the braincase described by <xref rid="ref-6" ref-type="bibr">Brusatte & Sereno (2007)</xref>, pertains to an individual ontogenetically less mature than the Moroccan neotype of <italic>C. saharicus</italic>). Nevertheless, assuming that the material belongs to a mature individual (<xref rid="ref-6" ref-type="bibr">Brusatte & Sereno, 2007</xref>), this plesiomorphic combination of features challenges the referral of the braincase from Niger to <italic>Carcharodontosaurus</italic>. <xref rid="ref-6" ref-type="bibr">Brusatte & Sereno (2007)</xref> listed the presence of large internal carotid foramina and deep paracondylar pneumatic foramina and the presence of a deep basisphenoid fossa as diagnostic features of <italic>Carcharodontosaurus</italic>, supporting the referral of the Nigerine braincase to the latter genus. Nevertheless, this combination of features is also shared by <italic>Giganotosaurus</italic> (<xref rid="ref-22" ref-type="bibr">Coria & Currie, 2002</xref>), indicating that they are synapomorphies of a clade more inclusive than <italic>Carcharodontosaurus</italic> and thus not diagnostic for the latter genus alone. In North African fossil assemblages, it is not uncommon to have two similarly-sized and closely related theropod taxa occurring in the same unit (e.g., <xref rid="ref-68" ref-type="bibr">Stromer, 1934</xref>; <xref rid="ref-18" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2012</xref>; <xref rid="ref-19" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2013</xref>; <xref rid="ref-30" ref-type="bibr">Fanti et al., 2014</xref>; <xref rid="ref-29" ref-type="bibr">Evers et al., 2015</xref>; <xref rid="ref-32" ref-type="bibr">Hendrickx, Mateus & Buffetaut, 2016</xref>). Therefore, in the absence of overlapping material with the type of <italic>C. iguidensis</italic> (i.e., maxillae), and lacking unambiguous braincase autapomorphies of <italic>Carcharodontosaurus</italic>, we cannot exclude that the associated dentary-braincase material pertains to a carcharodontosaurid species distinct from (and more basal than) <italic>C. iguidensis</italic>. Alternatively, if the referral of that material to <italic>C. iguidensis</italic> is confirmed, its combination of features may support a more basal placement for the latter taxon relative to Carcharodontosaurinae. In conclusion, in order to avoid the introduction of a possible chimera (in particular, in phylogenetic analyses), we provisionally exclude the referred material from <italic>C. iguidensis</italic>, restricting the latter name to the type maxilla.</p><p><italic>Sauroniops pachytholus</italic> is based on a large, isolated frontal from the KKCA (<xref rid="ref-18" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2012</xref>; <xref rid="ref-19" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2013</xref>). The specimen differs from all other known theropod frontals from the “mid-Cretaceous” of North Africa, in particular <italic>Carcharodontosaurus saharicus</italic> and the braincase referred to <italic>C. iguidensis</italic> (<xref rid="ref-63" ref-type="bibr">Sereno et al., 1996</xref>; <xref rid="ref-6" ref-type="bibr">Brusatte & Sereno, 2007</xref>; <xref rid="ref-62" ref-type="bibr">Sereno & Brusatte, 2008</xref>), and shares a set of unique features with the frontals from the Aptian of Niger referred to the basal carcharodontosaurid <italic>Eocarcharia dinops</italic> (<xref rid="ref-7" ref-type="bibr">Brusatte & Sereno, 2008</xref>).</p></sec><sec><title>Spinosauridae</title><p><italic>Sigilmassasaurus brevicollis</italic> is based on isolated presacral vertebrae from the KKCA (<xref rid="ref-57" ref-type="bibr">Russell, 1996</xref>) and was recently rediagnosed by <xref rid="ref-48" ref-type="bibr">McFeeters et al. (2013)</xref> and <xref rid="ref-29" ref-type="bibr">Evers et al. (2015)</xref>, including material that was referred to <italic>Spinosaurus</italic> (as <italic>Sp. maroccanus</italic>) by <xref rid="ref-57" ref-type="bibr">Russell (1996)</xref>. <xref rid="ref-36" ref-type="bibr">Ibrahim et al. (2014)</xref> suggested the referral of several specimens from the Cenomanian of Morocco to <italic>Spinosaurus aegyptiacus</italic>, including the material previously referred to <italic>Sigilmassasaurus</italic> (<xref rid="ref-48" ref-type="bibr">McFeeters et al., 2013</xref>). This hypothesis was recently challenged by <xref rid="ref-29" ref-type="bibr">Evers et al. (2015)</xref>, who referred part of the material of <italic>Spinosaurus</italic> (<italic>sensu</italic><xref rid="ref-36" ref-type="bibr">Ibrahim et al., 2014</xref>) to <italic>Sigilmassasaurus</italic>, the latter considered a distinct spinosaurid taxon. <xref rid="ref-29" ref-type="bibr">Evers et al. (2015)</xref> and <xref rid="ref-32" ref-type="bibr">Hendrickx, Mateus & Buffetaut (2016)</xref> provided evidence for the presence of more than one spinosaurid taxon in the KKCA. Accordingly, in this study, we distinguish between the material introduced by <xref rid="ref-36" ref-type="bibr">Ibrahim et al. (2014)</xref> and the material of <italic>Sigilmassasaurus</italic> (<italic>sensu</italic><xref rid="ref-29" ref-type="bibr">Evers et al., 2015</xref>), and provisionally restrict the name <italic>Spinosaurus aegyptiacus</italic> to the now lost holotype from Egypt, described by <xref rid="ref-66" ref-type="bibr">Stromer (1915)</xref>. We agree with <xref rid="ref-29" ref-type="bibr">Evers et al. (2015)</xref> that the erection of a neotype for <italic>S. aegyptiacus</italic> based on the material from Morocco described by <xref rid="ref-36" ref-type="bibr">Ibrahim et al. (2014)</xref> is not adequately justified. It should be noted that <xref rid="ref-29" ref-type="bibr">Evers et al. (2015)</xref> have rediagnosed <italic>Si. brevicollis</italic> based on comparison with the known presacral vertebrae of Spinosauridae, and listed a set of characters that does not completely overlap with that used by <xref rid="ref-57" ref-type="bibr">Russell (1996)</xref>. Accordingly, the taxon <italic>Sigilmassasaurus</italic> (sensu <xref rid="ref-29" ref-type="bibr">Evers et al., 2015</xref>) is less inclusive than <italic>Sigilmassasaurus</italic> (sensu <xref rid="ref-57" ref-type="bibr">Russell, 1996</xref>) because some of the diagnostic features of the latter are now known to be shared by other spinosaurid taxa (e.g., <italic>Baryonyx</italic>, <italic>Ichthyovenator</italic>; see <xref rid="ref-29" ref-type="bibr">Evers et al., 2015</xref>). Therefore, we cannot dismiss that some “<italic>Sigilmassasaurus</italic>-like” vertebrae from the KKCA, referred to <italic>Sigilmassasaurus</italic> by <xref rid="ref-57" ref-type="bibr">Russell (1996)</xref>, may eventually prove to not belong to <italic>Sigilmassasaurus</italic> (sensu <xref rid="ref-29" ref-type="bibr">Evers et al., 2015</xref>) but to other coeval spinosaurids, such as <italic>Spinosaurus</italic> (see <xref rid="ref-36" ref-type="bibr">Ibrahim et al., 2014</xref>; <xref rid="ref-32" ref-type="bibr">Hendrickx, Mateus & Buffetaut, 2016</xref>). Furthermore, we note that, following the distinction between <italic>Spinosaurus</italic> and <italic>Sigilmassasaurus</italic> proposed by <xref rid="ref-29" ref-type="bibr">Evers et al. (2015)</xref> and <xref rid="ref-32" ref-type="bibr">Hendrickx, Mateus & Buffetaut (2016)</xref>, the large and well-preserved spinosaurid snout from the KKCA described by <xref rid="ref-24" ref-type="bibr">Dal Sasso et al. (2005)</xref> cannot be referred unambiguously to the former taxon rather than the latter (see also the lack of resolution among spinosaurid taxa in the phylogenetic topology of <xref rid="ref-29" ref-type="bibr">Evers et al. (2015)</xref>). It is worth noting that <xref rid="ref-50" ref-type="bibr">Milner (2001)</xref> described a large spinosaurid dentary from the KKCA, comparable in length to the type dentary of <xref rid="ref-66" ref-type="bibr">Stromer (1915)</xref>, that differs from the latter in the overall stouter proportion of the bone, in the shape of the alveolar margin, and in the number and placement of the alveoli (at least 17, compared to 15 in the Egyptian specimen). This find further supports the hypothesis that the Moroccan material includes at least one spinosaurine taxon distinct from the Egyptian species. Since a discussion of the inclusiveness of the name <italic>Spinosaurus aegyptiacus</italic> (<xref rid="ref-36" ref-type="bibr">Ibrahim et al., 2014</xref>; <xref rid="ref-29" ref-type="bibr">Evers et al., 2015</xref>; <xref rid="ref-32" ref-type="bibr">Hendrickx, Mateus & Buffetaut, 2016</xref>) is beyond the aims of this study, and pending a taxonomic revision of the spinosaurid material from the Cenomanian of Morocco (in particular, the material introduced by <xref rid="ref-36" ref-type="bibr">Ibrahim et al. (2014)</xref>, (Maganuco, 2014, personal communication), (N. Ibrahim, personal communication in <xref rid="ref-32" ref-type="bibr">Hendrickx, Mateus & Buffetaut (2016)</xref>), we suggest to refer the KKCA material that cannot be referred unambiguously to either <italic>Spinosaurus</italic> or <italic>Sigilmassasaurus</italic> to Spinosaurinae indet., the least inclusive taxonomic unit all authors agree that material belongs to <xref rid="ref-32" ref-type="bibr">Hendrickx, Mateus & Buffetaut (2016)</xref>.</p><p><xref rid="ref-57" ref-type="bibr">Russell (1996)</xref> described the partial femur of an indeterminate theropod (“bone taxon M”), characterized by a robust shaft, declined head, distally placed lesser trochanter, and hypertrophied fourth trochanter. <xref rid="ref-13" ref-type="bibr">Carrano & Sampson (2008)</xref> noted the overall similarities to femora of basal theropods, including abelisaurids. As outlined below, based on presence of unique features of the femur referred to <italic>Spinosaurus</italic> by <xref rid="ref-36" ref-type="bibr">Ibrahim et al. (2014)</xref>, we refer “bone taxon M” to Spinosauridae.</p></sec><sec><title>Ceratosauria</title><p><italic>Deltadromeus agilis</italic> is based on a single, partial skeleton from the KKCA (<xref rid="ref-63" ref-type="bibr">Sereno et al., 1996</xref>) including the femora, the latter showing autapomorphic features. Originally interpreted as a coelurosaur (<xref rid="ref-63" ref-type="bibr">Sereno et al., 1996</xref>), more recent phylogenetic analyses agree in placing it among Ceratosauria (e.g., <xref rid="ref-61" ref-type="bibr">Sereno, Wilson & Conrad, 2004</xref>; <xref rid="ref-13" ref-type="bibr">Carrano & Sampson, 2008</xref>; <xref rid="ref-18" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2012</xref>). <xref rid="ref-63" ref-type="bibr">Sereno et al. (1996, note 32)</xref> distinguished <italic>D. agilis</italic> from <italic>Bahariasaurus ingens</italic> (from penecontemporary levels of Egypt, <xref rid="ref-68" ref-type="bibr">Stromer, 1934</xref>) on the basis of three features in the pubis and ischium, and referred part of the Egyptian material, that was first referred to <italic>Bahariasaurus</italic> by <xref rid="ref-68" ref-type="bibr">Stromer (1934)</xref>, to the Moroccan taxon. This interpretation was challenged by <xref rid="ref-13" ref-type="bibr">Carrano & Sampson (2008)</xref>, who suggested (without providing justification) that the bone interpreted by <xref rid="ref-63" ref-type="bibr">Sereno et al. (1996)</xref> as the distal end of the pubis of the holotype of <italic>Deltadromeus agilis</italic> may pertain to the ischium, thus invalidating the differences from the type material of <italic>Bahariasaurus ingens</italic>. The majority of the elements referred alternatively to <italic>Bahariasaurus</italic> or <italic>Deltadromeus</italic> share basal ceratosaurian and abelisauroid synapomorphies (<xref rid="ref-13" ref-type="bibr">Carrano & Sampson, 2008</xref>), including elongate, rectangular anterior caudal neural spines, dorsoventrally expanded acromion and coracoid, gracile and straight humerus with reduced deltopectoral crest, triangular obturator flanges on pubis and/or ischium, expanded ischial foot, prominent muscular insertions on laterodistal margin of femur, large fossa on proximomedial surface of fibula bounded posteriorly by a lip, and gracile fourth metatarsal with reduced distal end (<xref rid="ref-37" ref-type="bibr">Janensch, 1925</xref>; <xref rid="ref-68" ref-type="bibr">Stromer, 1934</xref>; <xref rid="ref-63" ref-type="bibr">Sereno et al., 1996</xref>; <xref rid="ref-14" ref-type="bibr">Carrano, Sampson & Forster, 2002</xref>; <xref rid="ref-13" ref-type="bibr">Carrano & Sampson, 2008</xref>; <xref rid="ref-53" ref-type="bibr">Novas et al., 2008</xref>). Therefore, even if not synonymous, the two taxa may be related to noasaurids or form a clade of mid- to large-bodied and gracile-limbed basal ceratosaurians, including <italic>Limusaurus</italic> and <italic>Elaphrosaurus</italic> (<xref rid="ref-13" ref-type="bibr">Carrano & Sampson, 2008</xref>; <xref rid="ref-19" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2013</xref>), for which the name Bahariasauridae (<xref rid="ref-34" ref-type="bibr">Huene, 1948</xref>) is available. Additional information on the femoral morphology of <italic>Deltadromeus</italic> was recently provided by <xref rid="ref-28" ref-type="bibr">Evans et al. (2015)</xref>. A large theropod femur from the Cenomanian of Egypt was assigned by <xref rid="ref-68" ref-type="bibr">Stromer (1934: 36, pl. 3, fig. 5)</xref> to <italic>Bahariasaurus</italic> (<xref ref-type="fig" rid="fig-3">Figs. 3D</xref> and <xref ref-type="fig" rid="fig-3">3E</xref>). Nevertheless, the type material of <italic>B. ingens</italic> lacks femora (<xref rid="ref-68" ref-type="bibr">Stromer, 1934</xref>); therefore, no direct evidence for referring the former specimen to that species is available. <xref rid="ref-63" ref-type="bibr">Sereno et al. (1996)</xref> referred that femur to <italic>Deltadromeus</italic> (<xref ref-type="fig" rid="fig-3">Figs. 3G</xref> and <xref ref-type="fig" rid="fig-3">3H</xref>), based on their resemblance to the Moroccan material and shared presence of autapomorphies of the latter (<xref rid="ref-63" ref-type="bibr">Sereno et al., 1996, note 5</xref>). Although this referral may further support a close relationship (if not synonymy) between <italic>Deltadromeus</italic> and an Egyptian gracile-limbed theropod (that may be <italic>Bahariasaurus</italic> itself), the Egyptian femur differs from the published holotype femur of <italic>D. agilis</italic> because it appears proportionally stouter, lacks a proximally directed head, and shows a proximodistally shorter lateral accessory crest on the distal end (<xref rid="ref-68" ref-type="bibr">Stromer, 1934</xref>; <xref rid="ref-28" ref-type="bibr">Evans et al., 2015</xref>). Some of these differences, in particular the stouter overall proportions, may be size-related because the Egyptian specimen is about one time and a half larger than the Moroccan specimen. Other differences are more difficult to explain as due to ontogenetic change. In particular, the Egyptian specimen (<xref rid="ref-68" ref-type="bibr">Stromer, 1934, Table III, fig. 5a</xref>; <xref rid="ref-54" ref-type="bibr">Rauhut, 1995, fig. 5F</xref>) shows a neck that is not particularly inclined proximally compared to <italic>Deltadromeus</italic> (see <xref rid="ref-28" ref-type="bibr">Evans et al., 2015, fig. 3B</xref>). Since the proximal inclination of the femoral neck is a weight-bearing adaptation shared by several large-bodied dinosaurs (<xref rid="ref-54" ref-type="bibr">Rauhut, 1995</xref>; <xref rid="ref-10" ref-type="bibr">Carrano, 1998</xref>), the absence of this feature in the more massive Egyptian specimen compared to the more gracile Moroccan specimen is unexpected if we assume that the two femora belong to the same ontogenetic trajectory, and raises question for the referral of the former to the same species of the latter.</p><p>Among the isolated bones from the KKCA described by <xref rid="ref-57" ref-type="bibr">Russell (1996)</xref>, one posterior dorsal vertebra (“bone taxon C”) was referred by the latter author to a large-bodied taxon distinct from <italic>Carcharodontosaurus</italic>, <italic>Sigilmassasaurus</italic> and <italic>Spinosaurus</italic> due to its unique combination of features. Among them, the vertebra is unusual in the relatively large size of the neural canal and the shape of the latter, described as dorsally separated into two halves by a low longitudinal ridge extending along the neural canal roof, and ventrally incised deeply into the centrum (<xref rid="ref-57" ref-type="bibr">Russell, 1996: 378</xref>). Both the large size and “heart-like” outline of the neural canal are shared by the posterior dorsal vertebra of a fragmentary theropod from the Lower Cretaceous of Libya (<xref rid="ref-65" ref-type="bibr">Smith et al., 2010</xref>), suggesting a possible relationship between these taxa. The Libyan taxon is referred to a large-bodied (estimated body length: 7–9 m, <xref rid="ref-65" ref-type="bibr">Smith et al., 2010, table 1</xref>) and gracile-limbed ceratosaurian based on the morphology of the femur and tibia and shows a unique combination of features that supports its referral to a new taxon (<xref rid="ref-65" ref-type="bibr">Smith et al., 2010</xref>).</p><p>Several isolated bone elements from the KKCA have been referred to Abelisauridae (<xref rid="ref-57" ref-type="bibr">Russell, 1996</xref>; <xref rid="ref-45" ref-type="bibr">Mahler, 2005</xref>; <xref rid="ref-13" ref-type="bibr">Carrano & Sampson, 2008</xref>; <xref rid="ref-26" ref-type="bibr">D’Orazi Porchetti et al., 2011</xref>). One abelisaurid, <italic>Rugops primus</italic>, is present in penecontemporary levels from Niger (<xref rid="ref-61" ref-type="bibr">Sereno, Wilson & Conrad, 2004</xref>). It is noteworthy that no abelisaurid material is known from the Baharjie assemblage (<xref rid="ref-67" ref-type="bibr">Stromer, 1931</xref>; <xref rid="ref-68" ref-type="bibr">Stromer, 1934</xref>; <xref rid="ref-13" ref-type="bibr">Carrano & Sampson, 2008</xref>), whereas the same clade is reported in the majority of North African “middle” Cretaceous localities (e.g., <xref rid="ref-61" ref-type="bibr">Sereno, Wilson & Conrad, 2004</xref>; <xref rid="ref-62" ref-type="bibr">Sereno & Brusatte, 2008</xref>; <xref rid="ref-30" ref-type="bibr">Fanti et al., 2014</xref>). <xref rid="ref-13" ref-type="bibr">Carrano & Sampson (2008)</xref> questioned the referral of the isolated maxillary fragment from the KKCA described by <xref rid="ref-45" ref-type="bibr">Mahler (2005)</xref> to Abelisauridae, noting that most of the features discussed by the latter author are shared by carcharodontosaurids. Nevertheless, additional abelisaurid synapomorphies, differentiating it from carcharodontosaurids, are present in this specimen (<xref rid="ref-17" ref-type="bibr">Cau & Maganuco, 2009</xref>).</p></sec><sec><title>Problematic material from the KKCA referred to Theropoda</title><p><xref rid="ref-47" ref-type="bibr">McFeeters (2013)</xref> reviewed the record of small-sized bones of theropods from the KKCA, concluding that most of the elements cannot be unambiguously referred to small-bodied taxa rather than immature individuals of large-bodied species. Among these elements, <xref rid="ref-56" ref-type="bibr">Riff et al. (2004)</xref> referred a small dorsal vertebra to Paraves, noting overall similarities with <italic>Rahonavis</italic>. Nevertheless, the specimen lacks unambiguous paravian or avialan synapomorphies. In particular, the large size of the neural canal, considered by <xref rid="ref-56" ref-type="bibr">Riff et al. (2004)</xref> as an avian synapomorphy, is a size-related feature homoplastically present among all small-bodied theropods (including small abelisauroids; see <xref rid="ref-14" ref-type="bibr">Carrano, Sampson & Forster, 2002</xref>) and also non-theropod taxa (e.g., crocodyliforms; see <xref rid="ref-42" ref-type="bibr">Lio et al., 2012</xref>).</p><p><xref rid="ref-17" ref-type="bibr">Cau & Maganuco (2009)</xref> referred an isolated distal caudal vertebra from the KKCA to a new mid-sized theropod, that they named <italic>Kemkemia auditorei</italic>. Most of the unique features (among theropods) present in this specimen are shared by crocodyliforms, challenging the referral of that vertebra to Theropoda (<xref rid="ref-42" ref-type="bibr">Lio et al., 2012</xref>). Among the unique features of <italic>K. auditorei</italic>, the robust (mediolaterally thick) neural spine with a concave dorsal surface is currently unreported among crocodyliform distal caudal vertebrae (<xref rid="ref-42" ref-type="bibr">Lio et al., 2012</xref>) and may represent an autapomorphic feature of this taxon. Although unreported among crocodyliforms, the unusual mediolateral broadening of the neural spine of <italic>K. auditorei</italic> is shared by a series of isolated caudal vertebrae from the KKCA referred to either <italic>Sigilmassasaurus</italic> by <xref rid="ref-57" ref-type="bibr">Russell (1996, figs. 12F–12G)</xref> or to an indeterminate dinosaur by <xref rid="ref-48" ref-type="bibr">McFeeters et al. (2013, fig. 10)</xref>, and, most recently, to <italic>Spinosaurus</italic> by <xref rid="ref-36" ref-type="bibr">Ibrahim et al. (2014)</xref>. <xref rid="ref-68" ref-type="bibr">Stromer (1934)</xref> described a similar caudal vertebral morphotype among the material of “<italic>Spinosaurus</italic> B” (<xref rid="ref-57" ref-type="bibr">Russell, 1996</xref>; <xref rid="ref-36" ref-type="bibr">Ibrahim et al., 2014</xref>; <xref rid="ref-29" ref-type="bibr">Evers et al., 2015</xref>). It is noteworthy that the Egyptian vertebra illustrated by <xref rid="ref-68" ref-type="bibr">Stromer (1934)</xref> differs from the Moroccan vertebrae of <xref rid="ref-57" ref-type="bibr">Russell (1996</xref>; see also <xref rid="ref-48" ref-type="bibr">McFeeters et al., 2013, fig. 10)</xref> in the unusual transversal broadening of the neural spine, the latter showing lateral margins that diverge apically in anterior view (in the Moroccan material, the lateral margins of the neural spine are subparallel in anterior view, <xref rid="ref-57" ref-type="bibr">Russell, 1996</xref>; <xref rid="ref-48" ref-type="bibr">McFeeters et al., 2013, fig. 10W</xref>). It is unclear whether this difference among the Moroccan and Egyptian vertebrae is merely positional, taxonomically significant, or–as suggested by <xref rid="ref-57" ref-type="bibr">Russell (1996)</xref>–a pathological feature of the Egyptian specimen. The holotype of <italic>K. auditorei</italic> also shares with the KKCA caudal vertebrae described by <xref rid="ref-57" ref-type="bibr">Russell (1996)</xref> the absence of a ventral sulcus in the centrum, the marked reduction of the zygapophyses, and the combination of a well-developed neural spine even in distal vertebrae lacking the ribs; whereas it differs from them in the presence of pre- and postspinal laminae (<xref rid="ref-17" ref-type="bibr">Cau & Maganuco, 2009</xref>; <xref rid="ref-48" ref-type="bibr">McFeeters et al., 2013</xref>). All the known caudal vertebrae referred to <italic>Sigilmassasaurus</italic> and/or <italic>Spinosaurus</italic> pertain to the proximal and middle parts of the tail and thus cannot be compared directly with the more-distally placed holotype of <italic>K. auditorei</italic> (<xref rid="ref-17" ref-type="bibr">Cau & Maganuco, 2009</xref>). Given the series of morphological convergences between spinosaurines and crocodyliforms (<xref rid="ref-36" ref-type="bibr">Ibrahim et al., 2014</xref>), the combination of crocodyliform-like and “<italic>Sigilmassasaurus</italic>-like” features in <italic>Kemkemia</italic> is intriguing: therefore, it is currently unclear whether the holotype of <italic>K. auditorei</italic> is referable to a crocodyliform or a spinosaurid.</p></sec><sec><title>Affinities of OLPH 025</title><p>The combination of large size, presence of both lesser trochanter and large medullary cavity in the shaft unambiguously indicates that OLPH 025 belongs to a theropod dinosaur (<xref rid="ref-59" ref-type="bibr">Sereno, 1999</xref>). <xref rid="ref-57" ref-type="bibr">Russell (1996)</xref> described the proximal portion of a femur from the ‘Kem Kem beds’ of Morocco (NMC 41869; <xref ref-type="fig" rid="fig-3">Figs. 3I</xref>–<xref ref-type="fig" rid="fig-3">3J</xref>) and referred it to an indeterminate theropod. OLPH 025 differs from NMC 41869 in having a larger medullary cavity, a more reclined head that is directed anteromedially, and in the presence of a distinct anterior corner of the shaft in distal view (<xref rid="ref-57" ref-type="bibr">Russell, 1996, figs. 25A–25C</xref>). Based on <xref rid="ref-57" ref-type="bibr">Russell (1996, fig. 25C)</xref>, NMC 41869 shows the head that is directed perpendicular to the anteroposterior axis of the shaft (indicated by the placement of the lesser and fourth trochanters), thus medially directed as in tetanurans and not anteromedially as in abelisauroids and OLPH 025. <xref rid="ref-57" ref-type="bibr">Russell (1996)</xref> described the fourth trochanter of NMC 41869 as “heavily developed.” Furthermore, the cross-section of the shaft depicted by <xref rid="ref-57" ref-type="bibr">Russell (1996, fig. 25C)</xref> shows a smaller medullary cavity than OLPH 025. Since the latter two features are reported exclusively in <italic>Spinosaurus</italic> (<italic>sensu</italic><xref rid="ref-36" ref-type="bibr">Ibrahim et al., 2014</xref>) among large-bodied theropods, we refer NMC 41859 to Spinosauridae. In overall features, OLPH 025 is more robust than a theropod femur from the Cenomanian of Egypt assigned by <xref rid="ref-68" ref-type="bibr">Stromer (1934: 36, pl. 3, fig. 5)</xref> to <italic>Bahariasaurus</italic>. Similarly to NMC 41869, the lesser trochanter of OLPH 025 lies more distally relative to the femoral head, a condition that differs from cf. <italic>Bahariasaurus</italic> and <italic>Carcharodontosaurus</italic> (<xref rid="ref-67" ref-type="bibr">Stromer, 1931, pl. 1, fig. 14</xref>). Furthermore, OLPH 025 differs from the large femur referred to <italic>Bahariasaurus</italic> by <xref rid="ref-68" ref-type="bibr">Stromer (1934)</xref> in the more distally placed lesser trochanter and the absence of a distinct accessory trochanter. OLPH 025 differs from <italic>Deltadromeus</italic> in the more reclined (distally directed) projection of the head, in the more distal placement of the lesser trochanter, and in the overall stouter proportions of the bone (<xref rid="ref-28" ref-type="bibr">Evans et al., 2015</xref>).</p><p>The other large-bodied theropods based on isolated material from the KKCA (i.e., <italic>Sauroniops pachytholus</italic> and <italic>Sigilmassasaurus brevicollis</italic>) cannot be directly compared to the Palermo specimen since no femora are known for either taxon. Both <italic>Sauroniops</italic> and <italic>Sigilmassasaurus</italic> are interpreted as tetanurans (i.e., respectively, a carcharodontosaurid and a spinosaurid, possibly synonymous with <italic>Spinosaurus</italic>; <xref rid="ref-18" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2012</xref>; <xref rid="ref-19" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2013</xref>; <xref rid="ref-48" ref-type="bibr">McFeeters et al., 2013</xref>; <xref rid="ref-36" ref-type="bibr">Ibrahim et al., 2014</xref>; <xref rid="ref-29" ref-type="bibr">Evers et al., 2015</xref>; <xref rid="ref-32" ref-type="bibr">Hendrickx, Mateus & Buffetaut, 2016</xref>). Since no synapomorphies of either Carcharodontosauridae or Spinosauridae (and other tetanuran clades) are present in OLPH 025, it is provisionally considered distinct from these taxa.</p><p>Most of the features present in the Palermo specimen are shared by ceratosaurid ceratosaurians (e.g., <xref rid="ref-44" ref-type="bibr">Madsen & Welles, 2000</xref>), a clade reported in the Aptian-Albian of South America (<xref rid="ref-55" ref-type="bibr">Rauhut, 2004</xref>) and possibly North Africa (<xref rid="ref-30" ref-type="bibr">Fanti et al., 2014</xref>). Nevertheless, the “ceratosaurid-like” features in OLPH 025 (e.g., distally reclined head, low lesser trochanter placed distally) are symplesiomorphies shared by most non-tetanuran neotheropods. Furthermore, the Palermo specimen apparently lacks the distinct trochanteric shelf present in <italic>Ceratosaurus</italic> (<xref rid="ref-44" ref-type="bibr">Madsen & Welles, 2000</xref>). Among non-tetanuran theropods, OLPH 025 is comparable in overall morphology to the femora of Abelisauridae (e.g., <xref rid="ref-12" ref-type="bibr">Carrano, 2007</xref>; <xref rid="ref-13" ref-type="bibr">Carrano & Sampson, 2008</xref>), as both show a distally reclined head, non-prominent trochanteric shelf, distally placed lesser trochanter of stout, alariform shape, a stocky shaft with the fourth trochanter placed proximally, and rugose muscular insertion areas (e.g., <xref rid="ref-12" ref-type="bibr">Carrano, 2007</xref>). Since the latter group is the only known Late Cretaceous clade of large-bodied non-tetanuran theropods (<xref rid="ref-13" ref-type="bibr">Carrano & Sampson, 2008</xref>) and abelisaurid material is already known from the KKCA (<xref rid="ref-57" ref-type="bibr">Russell, 1996</xref>; <xref rid="ref-45" ref-type="bibr">Mahler, 2005</xref>; <xref rid="ref-26" ref-type="bibr">D’Orazi Porchetti et al., 2011</xref>), we consider it most parsimonious to refer OLPH 025 to Abelisauridae.</p></sec><sec><title>Body size estimation of OLPH 025</title><p>Although incompletely preserved, the distal end of OLPH 025 provides information on the minimal mediolateral diameter of the femoral shaft, which we estimate as no less than 115 mm. The same diameter in a 1018 mm long femur of the large abelisaurid <italic>Carnotaurus</italic> measures 95 mm (<xref rid="ref-11" ref-type="bibr">Carrano, 2006</xref>), which may indicate a 1200 mm long femur for the Moroccan individual, comparable to the adult femora of cf. <italic>Bahariasaurus</italic>, <italic>Carcharodontosaurus</italic>, and <italic>Tyrannosaurus</italic> (<xref rid="ref-12" ref-type="bibr">Carrano, 2007</xref>). A length of 1041 mm results using the only known femur of <italic>Xenotarsosaurus</italic> as reference (<xref rid="ref-38" ref-type="bibr">Juarez-Valieri, Porfiri & Calvo, 2011, table 1</xref>). Nevertheless, other abelisaurids show hindlimb proportions stockier than those of <italic>Carnotaurus</italic> and <italic>Xenotarsosaurus</italic> (e.g., <italic>Majungasaurus</italic>, see <xref rid="ref-12" ref-type="bibr">Carrano, 2007</xref>; <italic>Ekrixinatosaurus</italic>, <xref rid="ref-38" ref-type="bibr">Juarez-Valieri, Porfiri & Calvo, 2011</xref>). Therefore, using the gracile-limbed taxa as reference may overestimate the actual length of the Moroccan bone if the latter pertained to the robust morphotype. In particular, the shaft diameter of OLPH 025 is approximately the same as that reported for the type femur of <italic>Ekrixinatosaurus novasi</italic> (shaft diameter, 115 mm; total length, 776 mm), a taxon considered among the most massive abelisauroids by <xref rid="ref-38" ref-type="bibr">Juarez-Valieri, Porfiri & Calvo (2011)</xref>. Based on a large sample of theropod femora known from both total length and mediolateral diameter of shaft, we estimate the minimal total length of OLPH 025 as 924 mm (data from <xref rid="ref-11" ref-type="bibr">Carrano (2006)</xref>, <italic>N</italic> = 55, <italic>r</italic><sup>2</sup> = 0.97). Therefore, we consider a value between 776 and 924 mm as the most conservative estimate for the total length of this Moroccan femur. Using the equation in <xref rid="ref-21" ref-type="bibr">Christiansen & Farina (2004)</xref> to infer total body mass from femur length, a value up to 1850 kg is suggested for this individual, making it among the largest ceratosaurians ever found.</p></sec><sec><title>Palaeoecological implications</title><p>The presence in the KKCA of one of the largest known specimens of Abelisauridae confirms that this clade had reached its largest known body size no later than the early Cenomanian (<xref rid="ref-65" ref-type="bibr">Smith et al., 2010</xref>; <xref rid="ref-38" ref-type="bibr">Juarez-Valieri, Porfiri & Calvo, 2011</xref>), and that large-bodied abelisaurids co-existed with giant carcharodontosaurids and spinosaurids in North Africa (<xref rid="ref-57" ref-type="bibr">Russell, 1996</xref>; <xref rid="ref-61" ref-type="bibr">Sereno, Wilson & Conrad, 2004</xref>; <xref rid="ref-6" ref-type="bibr">Brusatte & Sereno, 2007</xref>; <xref rid="ref-19" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2013</xref>). Unfortunately, the majority of theropod-bearing localities from North Africa lacks detailed information on the geological context of the dinosaurian material (<xref rid="ref-49" ref-type="bibr">McGowan & Dyke, 2009</xref>; <xref rid="ref-20" ref-type="bibr">Cavin et al., 2010</xref>; <xref rid="ref-30" ref-type="bibr">Fanti et al., 2014</xref>). In absence of detailed stratigraphic, taphonomic, and palaeoecological data, it is unclear whether these large-bodied theropod lineages were sympatric and ecologically overlapping or, on the contrary, each group was constrained to a distinct environmental context, with their co-occurrence in the same depositional setting being mainly due to taphonomic factors (see <xref rid="ref-33" ref-type="bibr">Hone, Xu & Wang, 2010</xref>; <xref rid="ref-30" ref-type="bibr">Fanti et al., 2014</xref>; <xref rid="ref-32" ref-type="bibr">Hendrickx, Mateus & Buffetaut, 2016</xref>). The co-occurrence of giant carcharodontosaurids and large abelisaurids in the KKCA recalls the faunal composition of the Candeleros Formation (Neuquén Basin, Argentina), where both <italic>Giganotosaurus</italic> and <italic>Ekrixinatosaurus</italic> are reported (<xref rid="ref-38" ref-type="bibr">Juarez-Valieri, Porfiri & Calvo, 2011</xref>). In this regards, the Moroccan and Niger assemblages are more similar to the Aptian-Cenomanian faunas from South America (see <xref rid="ref-51" ref-type="bibr">Novas et al., 2013</xref>, and reference therein) than the Cenomanian fauna from Egypt, where no abelisaurids are known (<xref rid="ref-67" ref-type="bibr">Stromer, 1931</xref>; <xref rid="ref-68" ref-type="bibr">Stromer, 1934</xref>; <xref rid="ref-13" ref-type="bibr">Carrano & Sampson, 2008</xref>; <xref rid="ref-62" ref-type="bibr">Sereno & Brusatte, 2008</xref>). On the contrary, the KKCA recalls the Baharjie fauna in the presence of large-bodied and gracile-limbed ceratosaurians (bahariasaurids), the latter unknown from Niger and South America. Among non-theropod dinosaurs, both the Candeleros Formation and the KKCA include rebbachisaurid and basal titanosaurian sauropods (<xref rid="ref-57" ref-type="bibr">Russell, 1996</xref>; <xref rid="ref-9" ref-type="bibr">Calvo, Rubilar-Rogers & Moreno, 2004</xref>): on the contrary, rebbachisaurids appear absent from both Niger and Egypt, whereas titanosaurians are reported in Egypt (<xref rid="ref-67" ref-type="bibr">Stromer, 1931</xref>; <xref rid="ref-64" ref-type="bibr">Smith et al., 2001</xref>). Given the small number of collected individuals belonging to the aforementioned clades, the differences among these faunal assemblages may be artifacts due to sampling bias. Nevertheless, it is worth noting that spinosaurids, abundantly recorded in the KKCA and other African assemblages (<xref rid="ref-57" ref-type="bibr">Russell, 1996</xref>; <xref rid="ref-24" ref-type="bibr">Dal Sasso et al., 2005</xref>; <xref rid="ref-33" ref-type="bibr">Hone, Xu & Wang, 2010</xref>; <xref rid="ref-36" ref-type="bibr">Ibrahim et al., 2014</xref>), are currently absent from the Candeleros Formation (<xref rid="ref-38" ref-type="bibr">Juarez-Valieri, Porfiri & Calvo, 2011</xref>). We therefore consider this faunal difference among the large theropods from the KKCA and the Candeleros Formation as not biased by collecting or taphonomic factors. A possible explanation of the anomalous distribution of spinosaurids, when compared to the other mentioned saurischians, is provided by the theropod record in the Ain El Guettar Formation (Albian of Tunisia, <xref rid="ref-30" ref-type="bibr">Fanti et al., 2014</xref>). In the Ain El Guettar Formation, an abelisaurid-carcharodontosaurid association dominates the lower Chenini Member, characterized by wadi-like channels and arid alluvial plain deposits, whereas spinosaurids dominate the upper Oum ed Diab Member, characterized by estuarine and embayment deposits (<xref rid="ref-30" ref-type="bibr">Fanti et al., 2014</xref>). Assuming that this stratigraphic (and, inferred, ecological and environmental) partition between the large-bodied theropods also characterized other “mid-Cretaceous” associations from Africa and South America, we conclude that spinosaurids were ecologically and environmentally segregated to other large-bodied theropods (<xref rid="ref-33" ref-type="bibr">Hone, Xu & Wang, 2010</xref>). This hypothesis is supported by the morphological specializations of spinosaurids (and, in particular, spinosaurines; <xref rid="ref-3" ref-type="bibr">Amiot et al., 2010</xref>; <xref rid="ref-36" ref-type="bibr">Ibrahim et al., 2014</xref>; <xref rid="ref-32" ref-type="bibr">Hendrickx, Mateus & Buffetaut, 2016</xref>) that suggest a mode of life distinct from that of other theropods. As discussed by <xref rid="ref-20" ref-type="bibr">Cavin et al. (2010)</xref>, the KKCA includes at least two distinct formations (the Ifezouane Formation and the overlying Aoufous Formation), with the vast majority of the dinosaurian remains recovered without detailed taphonomic information and often with ambiguous stratigraphic placement. <xref rid="ref-63" ref-type="bibr">Sereno et al. (1996)</xref>, distinguished between a lower and upper units of their “Kem Kem beds,” but it is unclear how these two units fit the Ifezouane and Aoufous formations of <xref rid="ref-20" ref-type="bibr">Cavin et al. (2010)</xref> and <xref rid="ref-32" ref-type="bibr">Hendrickx, Mateus & Buffetaut (2016, supplemental material)</xref>. Therefore, we conclude that the reported co-occurrence of spinosaurids with abelisaurids and carcharodontosaurids in the KKCA may reflect the lack of stratigraphic resolution in a heterogeneous sample recovered from multiple units rather than a genuine evidence of sympatry and ecological overlap between these theropods.</p></sec></sec><sec sec-type="conclusions"><title>Conclusions</title><p>The taxonomy and inclusiveness of the theropod clades from the “middle” Cretaceous of North Africa is complex and problematic. Since <xref rid="ref-67" ref-type="bibr">Stromer (1931)</xref> and <xref rid="ref-68" ref-type="bibr">Stromer (1934)</xref> the minimum number of taxa recovered from these fossil associations has been considered controversial, in particular due to the fragmentary nature of most of the specimens found. Stromer himself (1934) was aware of this as one of the main problems in North African dinosaur palaeontology. Several factors, both biological and geological, may bias the taxonomic composition of the North African theropod faunas. Most North African units are poorly constrained stratigraphically (see <xref rid="ref-20" ref-type="bibr">Cavin et al., 2010</xref>; <xref rid="ref-30" ref-type="bibr">Fanti et al., 2014</xref>), thus preventing detailed correlations between the various localities. For example, the age of the KKCA has been alternatively placed between the Aptian and the Cenomanian (<xref rid="ref-57" ref-type="bibr">Russell, 1996</xref>; <xref rid="ref-20" ref-type="bibr">Cavin et al., 2010</xref>), and both number of and relationships among the units represented by that assemblage remain controversial (<xref rid="ref-63" ref-type="bibr">Sereno et al., 1996</xref>; <xref rid="ref-20" ref-type="bibr">Cavin et al., 2010</xref>). The temporal extent of these assemblages is uncertain, possibly spanning several million years (<xref rid="ref-20" ref-type="bibr">Cavin et al., 2010</xref>). Therefore, the application of biological (neontological) “rules,” based on ecological models and data from modern ecosystems (in order to constrain the number of carnivorous taxa included in a fossil assemblage) is often not adequately justified or not testable. This is particularly problematic for fossil assemblages, like the KKCA, that lack present-day analogues and where an unusually unbalanced ecological web has been suggested (e.g., <xref rid="ref-40" ref-type="bibr">Läng et al., 2013</xref>). Since the co-occurrence in the same North African theropod associations of distinct species belonging to the same clade has been documented (e.g., spinosaurids, <xref rid="ref-30" ref-type="bibr">Fanti et al., 2014</xref>; <xref rid="ref-32" ref-type="bibr">Hendrickx, Mateus & Buffetaut, 2016</xref>; carcharodontosaurids, <xref rid="ref-18" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2012</xref>; <xref rid="ref-19" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2013</xref>), the referral of all isolated elements of one lineage to a single species cannot be justified. Furthermore, the referral of isolated and non-overlapping material to the same species is a phylogenetic hypothesis itself that needs to be explicitly tested by numerical analyses. In absence of positive evidence supporting the referral of such material to a particular species, the inclusion of non-overlapping elements into a single taxon may led to the creation of a potential chimera, with unpredictable effects on the phylogenetic and palaeoecological interpretation of these faunas.</p><p>We have described the fragmentary femur of a large-bodied theropod from the “Kem Kem Compound Assemblage” of Morocco. The specimen lacks tetanuran synapomorphies and is referred to Abelisauridae as it shares the overall morphology of the femora of ceratosaurians and the stocky robust proportions of some Late Cretaceous abelisaurids (e.g., <italic>Ekrixinatosaurus</italic>, <italic>Majungasaurus</italic>, <xref rid="ref-12" ref-type="bibr">Carrano, 2007</xref>; <xref rid="ref-38" ref-type="bibr">Juarez-Valieri, Porfiri & Calvo, 2011</xref>). The large size of the preserved femur suggests an individual comparable in body size with the type specimens of <italic>Carnotaurus sastrei</italic> and <italic>Ekrixinatosaurus novasi</italic>, both estimated to reach 9 meters in length and approaching two tons in body mass (<xref rid="ref-38" ref-type="bibr">Juarez-Valieri, Porfiri & Calvo, 2011</xref>). This discovery further supports that abelisaurids had evolved their largest size no later than the “mid-Cretaceous” (<xref rid="ref-65" ref-type="bibr">Smith et al., 2010</xref>; <xref rid="ref-38" ref-type="bibr">Juarez-Valieri, Porfiri & Calvo, 2011</xref>) and that abelisaurids and carcharodontosaurids co-existed and ecologically overlapped in both North Africa and South America during the Aptian-Turonian. Based on comparison with other “middle Cretaceous” units (<xref rid="ref-38" ref-type="bibr">Juarez-Valieri, Porfiri & Calvo, 2011</xref>; <xref rid="ref-30" ref-type="bibr">Fanti et al., 2014</xref>), we suggest that the co-occurrence of spinosaurids and other large theropods (abelisaurids and carcharodontosaurids) in the KKCA may be mainly an artefact due to poor stratigraphic resolution rather than genuine evidence of ecological and environmental overlap. Given the convergent evolution of several craniodental features among abelisaurids and carcharodontosaurids (<xref rid="ref-39" ref-type="bibr">Lamanna, Martinez & Smith, 2002</xref>; <xref rid="ref-58" ref-type="bibr">Sampson & Witmer, 2007</xref>; <xref rid="ref-13" ref-type="bibr">Carrano & Sampson, 2008</xref>; <xref rid="ref-19" ref-type="bibr">Cau, Dalla Vecchia & Fabbri, 2013</xref>), suggesting similar ecological adaptations in these clades, how these apparently competing groups co-existed for at least 30 million years in both Africa and South America remains to be resolved.</p></sec></body><back><ack><p>We thank Carolina D’Arpa (Museo Geologico e Paleontologico “Gaetano Giorgio Gemmellaro” di Palermo) for access to the specimen, Antonietta Rosso (Università di Catania) for technical help at an early stage of this project, Joanna Wolstenholme (Imperial College London) for linguistic advice, Serjoscha Evers (University of Oxford) and Philip Mannion (Imperial College London) for valuable discussion. Stephen Brusatte (University of Edinburgh) and Oliver Rauhut (Bayerische Staatssammlung für Paläontologie und Geologie) are greatly acknowledged for their helpful reviews. We also thank Hans-Dieter Sues (National Museum of Natural History of the Smithsonian Institution) for his editorial guidance and constructive comments that significantly improved the quality of the manuscript.</p></ack><sec sec-type="additional-information"><title>Additional Information and Declarations</title><fn-group content-type="competing-interests"><title>Competing Interests</title><fn fn-type="conflict" id="conflict-1"><p>The authors declare that they have no competing interests.</p></fn></fn-group><fn-group content-type="author-contributions"><title>Author Contributions</title><fn fn-type="con" id="contribution-1"><p><xref ref-type="contrib" rid="author-1">Alfio Alessandro Chiarenza</xref> conceived and designed the experiments, performed the experiments, analyzed the data, contributed reagents/materials/analysis tools, wrote the paper, prepared figures and/or tables, reviewed drafts of the paper.</p></fn><fn fn-type="con" id="contribution-2"><p><xref ref-type="contrib" rid="author-2">Andrea Cau</xref> conceived and designed the experiments, performed the experiments, analyzed the data, contributed reagents/materials/analysis tools, wrote the paper, prepared figures and/or tables, reviewed drafts of the paper.</p></fn></fn-group><fn-group content-type="other"><title>Data Deposition</title><fn id="addinfo-1"><p>The following information was supplied regarding data availability:</p><p>The research in this article did not generate any raw data.</p></fn></fn-group></sec><ref-list content-type="authoryear"><title>References</title><ref id="ref-1"><label>Accarie et al. 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