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In some cases, the quadrate shaft also encompasses a ventrodorsally elongated depression, or fossa, on the posterior side of the quadrate and known as the posterior fossa. A second depression, the medial fossa, is located on the ventromedial surface of the pterygoid flange and is bounded by the quadrate shaft posteriorly. The quadrate head can be monostylic or bistylic and divided by an intercapitular sulcus into the squamosal and otic capitula. The mandibular articulation includes, in the large majority of theropods, two condyles. The lateral condyle of the mandibular articulation, called ectocondyle, is separated from the medial condyle, or entocondyle, by the intercondylar sulcus. An anterior or posterior intercondylar notch can sometimes be seen either on the anterior or posterior surface of the intercondylar sulcus, respectively, between the two mandibular condyles, 9k yeti como jogar. When pneumatic, the quadrate includes one or several pneumatic openings, i. The theropod phylogeny adopted here follows the classification summarized by Hendrickx et al. Likewise, we follow the phylogenetic definitions compiled by Hendrickx et al. Values are given in millimeters. A phylogenetic analysis was performed to assess the phylogenetic relationships of the quadrate bones from the Kem Kem beds, and the bones were coded in an updated version of the supermatrix of Hendrickx et al. The supermatrix encompasses 98 quadrate-related characters (S1 File) originally associated with six recent datasets (i. All quadrate-based characters were removed from the six datasets. The main changes are the inclusion, in the supermatrix, of the data matrix of Novas et al. The final supermatrix includes 2377 characters and 59 taxa for one outgroup ( Eoraptor ; Table A in S1 File). KK374 (Fig 3A and 3C) and absent in all other isolated quadrates. The deep intercondylar sulcus of Spinosaurus aegyptiacus also extends far dorsally along the posterior surface of the quadrate, forming a posterior intercondylar notch absent in both quadrate morphotypes (but present in Suchomimus ; char. Morphotypes 1 and 2 were recovered in two separate spinosaurine clades in the phylogenetic analysis, the former being closely related to Spinosaurus whereas the latter forms a sister-taxon pair with Irritator (Fig 6). Nevertheless, the two ambiguous synapomorphies uniting Morphotype 2 and Irritator (i. Consequently, based on the results of the phylogenetic analysis, and given the fact that fossils of Spinosauridae have so far been assigned to the spinosaurid taxa Spinosaurus and Sigilmassasaurus , Morphotype 1 is referred with confidence to Spinosaurus , and Morphotype 2 likely belongs to Sigilmassasaurus , a referral that will, however, be thoroughly discussed in another section below. Likewise, given the almost identical morphology of quadrates of Morphotype 1 and FSAC-KK 11888, Morphotype 1 is confidently assigned to the species Spinosaurus aegyptiacus , an opinion followed by Ibrahim et al, 9k yeti como jogar. Ontogenetic variation occurring in the spinosaurid quadrate was briefly investigated by Hendrickx and Mateus [88] based on the material examined here. Given the relative size of the specimens, the level of coossification between the quadrate and quadratojugal and the development of dorsal and mandibular condyles, it is assumed that Morphotype 1 includes quadrates belonging to juvenile, immature, sub-adult and adult individuals, so that a sequence list of ontogenetic character transformations (maturity-dependent characters) can be provided for the quadrate of Spinosaurus aegyptiacus. Due to the scarcity of spinosaurid cranial material, no destructive methods were employed to conduct histological analysis and to investigate growth series in the Spinosaurus quadrates. At a juvenile stage represented by MHNM. KK374, the quadrate lacks several deep grooves on the lateral side of the ventral quadratojugal contact, and two grooves on the dorsal quadratojugal contact (Fig 10A’10C). The ectocondyle is also poorly developed and the dorsal quadratojugal contact lacks a ventral projection. A ‘ C , Left quadrate MHNM. KK374 representing ontogenetic stage 1 (juvenile) with A , close up on the smooth lateral surface of the dorsal quadratojugal contact in lateral view; B , quadrate foramen and absence of a ventral projection of the dorsal quadratojugal contact in posterior view; C , and non-delimited mandibular condyles in ventral view. D ‘ E , Left quadrates of specimens D, F , MHNM.
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