Anatosuchus minor Sereno 2003
Creators: name: Sereno, Paul
name: Larsson, Hans
Publisher: name: Zenodo
Year: 2009
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Reptilia
Crocodylia
Uruguaysuchidae
Anatosuchus

description: <i>Anatosuchus minor</i> Sereno et al., 2003 Figs. 4 –10, 12, 13 Tables 2–6 Sereno et al. (2001, figs. 1, 2) <b>Holotype.</b> MNN GAD603; nearly complete skull with lower jaws of a subadult individual; margins of the skull are eroded away. The holotype was previously catalogued as “ GDF603 ” (Sereno et al. 2003). <b>Type locality.</b> Gadoufaoua, Agadez District, Niger Republic (N 16° 46’, E 9° 22’) (Fig. 1A, C). <b>Horizon.</b> Elrhaz Formation, Tegama Series; Lower Cretaceous (Aptian-Albian), ca. 110 Mya (Taquet 1976). In association with a diverse dinosaurian fauna (Taquet 1976; Sereno et al. 1998, 1999, 2007; Taquet and Russell 1999; Sereno and Brusatte 2008) and the crocodyliforms <i>Sarcosuchus imperator</i> (Broin and Taquet 1966; Sereno et al. 2001), <i>Araripesuchus wegeneri</i> (Buffetaut and Taquet 1979), and <i>Stolokrosuchus lapparenti</i> (Larsson and Gado 2000). At a single field locality (G109), specimens were recovered that are referable to <i>Anatosuchus minor</i> (MNN GAD 18) and <i>Araripesuchus wegeneri</i> (MNN GAD 19). <b>Referred material.</b> MNN GAD 17 (Figs. 4–8, 12, 13), nearly complete skull with lower jaws lacking only the anterolateral corner of the snout in articulation with a postcranial skeleton lacking the right pectoral girdle and forelimb, most of both hind limbs, sacrum, and tail; MNN GAD 18 (Fig. 9), mid-section of the left dentary preserving alveoli 7–14 and the anterior tip of the left splenial. <b>Revised diagnosis.</b> Small-bodied metasuchian (&lt;1.0 m) with low transversely expanded snout that forms the broadest portion of the cranium, broad-based anteriorly projecting pointed internarial bar, lenticular-shaped external nares, elevated narial bridge which expands transversely behind the external nares, prominent median edentulous dentary margin, laterally projecting vascularized dentary shelf on parasagittal portion of dentary ramus, enlarged neurovascular foramina located along the anterior snout margin, anterior snout margin smooth, vertical and sharply defined on the premaxilla and maxilla, oval splenial fenestra on the anterior transverse portion of the lower jaw, six premaxillary teeth, premaxillary and anterior maxillary tooth row that angles ventrolaterally toward the corner of the snout at approximately 25°, largest upper and lower teeth positioned along the bend in the L-shaped tooth row (m4, d12), three pairs of cervical osteoderms that decrease in size posteriorly, large manus (30% skull length), elongate poorly recurved manual unguals on digits I-III, and manual digit IV with six phalanges. The initial description was based on an immature skull embedded in a hematitic concretion (MNN GAD 603). The concretion was discovered on the surface with prominent edges of the skull, such as the anterior end of the snout, trimmed by erosion (Sereno et al. 2003). Th e likeness drawn between <i>Anatosuchus</i> and the South American genus <i>Comahuesuchus</i> was based on a few seemingly unique features, such as a diastema between the premaxillary tooth rows, which we can now say arose in the immature skull of <i>Anatosuchus</i> as an artifact of erosion. Th e revised diagnosis is based mainly on a referred adult skull and partial articulated postcranium (MNN GAD 17) that preserves an intact portion of the paravertebral shield (Fig. 4). Th is well preserved skull was found embedded in sandstone, the right corner of the snout, right limbs, sacrum and tail lost to erosion. Th e additional information available for both <i>Anatosuchus</i> and <i>Comahuesuchus</i> confirms Martinelli’s (2003) view that these genera are not closest relatives among known notosuchians. <b>Dorsal skull roof.</b> In <i>A. minor</i> the snout becomes relatively broader and longer during growth. In the juvenile holotype specimen MNN GAD603, the width of the skull across the rounded anterior corner of the snout is subequal to that across the suborbital ramus of the jugal (Sereno et al. 2003). Preorbital length, in addition, is subequal to that of the remainder of the skull. In mature individuals, in contrast, the anterior snout corner is the broadest region of the skull, and preorbital length is approximately 20% greater than the posterior portion of the skull (MNN GAD17; Figs. 5, 6; Tables 2, 3). Th e following description is based primarily on this specimen. The <i>premaxilla</i> is a broad bone housing six recurved teeth. Th e base of the internarial process is broad, unlike that in <i>Araripesuchus,</i> but similar in this regard to <i>Simosuchus</i> (Buckley et al. 2000). It extends anteriorly at approximately 30° above the horizontal, and tapers to a point, where it joins at a sharp angle the nearly horizontal internarial process of the nasal (Figs. 5, 6). Th e external nares, as a result, are dorsoventrally compressed and appear as a narrow slit in lateral view. In dorsal view, the external nares are elliptical, the floor of the narial passage broadly exposed to each side of the tapering internarial process of the nasal. The floor of the narial passage, which is formed by the premaxilla, is raised and slightly extended anterolaterally by a short tongue-shaped flange (Figs. 5B, 6B, 7A). The anterior half of the external nares projects beyond the first premaxillary tooth, a narial structure that projects anteriorly more prominently than in any other crocodyliform. The narial fossa is clearly demarcated as a smooth subtriangular surface located lateral to the external nares and restricted to the premaxilla. In glancing light, a subtle division of the surface is visible. A teardrop-shaped fossa within the narial fossa is the largest surface, its tip emerging from under the lip of the rim of the external naris. In ventral view, the anterior projection is smooth and incorporates into the narial fossa the alveolar margin dorsal to premaxillary teeth 1–3. The lateral margin of the narial fossa is delimited by a shallow trough from the smooth, highly vascularized, vertical alveolar margin, which extends laterally toward the premaxilla-maxilla suture. No other crocodyliform known thus far closely approaches the form and orientation of the external nares in <i>A. minor</i>. The remainder of the external surface of the premaxilla can be divided into the alveolar margin and the ramus that tapers between the nasal and maxilla. The alveolar are measured on left side except as indicated. 1 Measurement from right side. margin faces primarily anteriorly, has a vertical orientation, and is gently transversely convex (Fig. 7A, B). As in <i>Araripesuchus wegeneri</i>, two large neurovascular foramina are situated between the narial fossa and the premaxilla-maxilla foramen. The ventral margin is scalloped to match the position of the lateral three premaxillary teeth (Fig. 7B) as occurs in <i>Simosuchus,</i> but unlike the straight margin in <i>Araripesuchus</i>. The dorsal margin meets the dorsal surface of the snout at nearly a right angle along a rugose edge. Small foramina and grooves for impressed vessels are visible on the dorsal surface of the snout near the narial fossa and alveolar margin. Th at texture becomes deeply pitted as the premaxilla tapers to a point on the lateral aspect of the nasal bridge. In ventral view, the premaxilla is divided between the transversely convex surface of the internarial bar, the raised edges of the alveoli that scallop the alveolar margin, and the flat palatal surface, which is only partially exposed (Figs. 5C, 6C). The <i>maxilla</i> is the most expansive bone in the skull and forms most of the snout. Its external surface is composed of a narrow alveolar margin and broader posterodorsal and posteroventral rami that extend above and below the antorbital opening, respectively. Like the premaxilla, the alveolar surface is vertical (Fig. 7B). It faces anterolaterally, borders the premaxilla-maxilla foramen, and gives passage to one additional large neurovascular foramen. Th e dorsal edge protrudes over this foramen before curving posteroventrally to join the scalloped ventral margin near the overhanging corner of the snout adjacent to the fourth maxillary tooth. Several large foramina are present just above this edge on the corner of the snout (Fig. 7B). The dorsal surface of the maxilla remains lightly textured along a band near the sharp anterior margin of the snout from the narial fossa to the anterolateral corner. This same low texture is present across the posteroventral ramus lateral to the antorbital depression, a muted textural pattern that resembles that seen in <i>Simosuchus</i>. In both taxa most of the maxilla below the antorbital opening is only lightly textured. In <i>Araripesuchus</i>, by contrast, the comparable region of the maxilla above m3 and m4 is more deeply sculpted with pits (Figs. 14A, 15A). As in most crocodyliforms, in <i>A. minor</i> a row of neurovascular foramina runs above the alveolar margin along the posteroventral ramus, although these are smaller than those at the anterior end of the snout. The maxilla forms the smooth and elongate anterior wall of the antorbital fossa, which is pierced by a foramen (Fig. 7C). Th e posterodorsal ramus of the maxilla is deeply pitted and slightly elevated as it passes over the antorbital depression to join the lacrimal and prefrontal. The <i>nasal</i> extends from the tip of the internarial bar anteriorly to a subquadrate process posteriorly. Th e texture is reduced on the nasals immediately posterior to the external nares. Nonetheless, shallow sculpting is present, and the nasals do not contribute to the smooth narial fossa, which is isolated on the premaxilla as in <i>Araripesuchus</i> (Fig. 16A), <i>Simosuchus</i> (Buckley et al. 2000) and other crocodyliforms. The elevated nasal bridge is narrowest in width at mid-length along the snout, after which it broadens slightly to equal interorbital width (Figs. 5B, 6B). A narrow median trough is present from mid-snout to the subrectangular interdigitating ends of the nasals. The L-shaped <i>lacrimal</i> has anterior and ventral rami, which join near a laterally prominent process for articulation with a missing anterior palpebral (Fig. 7C). The lacrimal foramen is tucked under this process within the orbit. Th e anterior ramus is deeply pitted and joins the maxilla along a subrectangular suture. Th e ventral ramus is smooth and divided into an orbital margin and medially inset posterior margin of the antorbital fossa. The <i>palpebrals</i> are disarticulated in both known skulls. In the adult skull, however, they have fallen into orbital and temporal spaces, where they are partially exposed. A pair of articular fossae, the anterior on the lacrimal and <i>prefrontal</i> and the posterior on the postorbital, supported anterior and posterior palpebrals, respectively, as in many crocodyliforms (Fig. 7C, D). Th e prefrontal-frontal suture courses anteriorly, extending parallel to the inset of the fossa for the anterior palpebral. Th e prefrontal narrows in mid-section, where it contacts the lacrimal, and then extends anteriorly to contact the maxilla, effectively separating the nasal and lacrimal. Th e prefrontal pillar angles ventromedially and slightly posteriorly, tapering strongly from the skull roof to the palate. The <i>frontal</i> and <i>parietal</i> are fused to their opposites and joined to each other by an interdigitating frontoparietal suture in both the adult and subadult skulls. The deeply pitted frontals have a median crest. The flat skull table formed by the parietals is also deeply pitted and separates the supratemporal fossae to a greater degree than in <i>Simosuchus</i> (Buckley et al. 2000). During growth in <i>A. minor</i>, interorbital width expands relative to the width of the skull table, such that the two measurements are subequal in a subadult (Sereno et al. 2003) whereas the former is nearly twice the latter in an adult (Figs. 5B, 6B). In the adult skull the frontal forms the anteromedial rim and distinctive corner of the supratemporal fossa, which is not the case in the subadult skull. Th at corner, in addition, is invaded by diverticulae from the supratemporal fossa. Although there is a similar corner in the rim of the fossa in <i>Araripesuchus wegeneri</i>, the rim is not undercut by pneumatic invagination. <i>Simosuchus</i>, on the other hand, has diverticulae resembling the condition in <i>A. minor</i> that undercut the anterior rim of the supratemporal fossa, a condition that has arisen a few times among crocodyliforms. The frontal contributes to the rim of the supratemporal fossa and reaches the fossa in dorsal view. Frontal participation in these supratemporal structures seems to occur with maturity, given the exclusion of the frontal in a subadult skull (Sereno et al. 2003). Th e posterior margin of the skull table is scalloped to each side of a short posteromedian projection formed by the supraoccipital, which joins the parietals along a shallow V-shaped suture. <i>Simosuchus</i>, in contrast, is shown with a nearly straight posteromedian margin. In this case, notching of the posterior margin of the parietals by the supraoccipital may have been obliterated by coossification. The right side of the skull has rotated slightly posterolaterally, an asymmetry best seen in dorsal view (Figs. 5B, 6B). Because there is no pattern of postmortem distortion of the skull, this asymmetry appears to be pathological rather than preservational in origin. Th e articular notch for the posterior palpebral on the right side is shifted posterolaterally, altering the shape of the supratemporal fossa. Th e right fossa has a convex lateral margin and its maximum parasagittal length is about 10% longer than the left side. The <i>postorbital</i> is notched by an articular facet for a small posterior palpebral. The surface of the postorbital between the facet and the supratemporal fossa varies, remaining textured with pits in some species, such as <i>A. gomesii</i> (Price 1959) and <i>A. tsangatsangana</i> (Turner 2006), and smooth in others such as <i>A. patagonicus</i> (Ortega et al. 2000). In <i>A. wegeneri</i> that surface between the palpebral facet and supratemporal fossa is smooth and convex (Figs. 14B, 15B). The <i>squamosal</i> is distinctly triradiate in dorsal view, the anterior process that contacts the postorbital the most slender. The dorsal surface of the anterior process is deeply pitted and depressed to form a shallow arcuate fossa (Figs. 5B, 6B). The posterior process is offset below the skull table and has a more subdued texture. The <i>jugal</i> approaches, but does not contact, the posteroventral corner of the antorbital fossa (Fig. 7C). Th e anterior ramus is moderately expanded dorsoventrally toward its anterior end and is deeply pitted, with an oval fossa located beneath the orbit (Fig. 7D). The relatively slender postorbital process is inset at its base, the location for a very small siphonal opening. Th e posterior ramus is also relatively slender under the laterotemporal fenestra, where it terminates in a shallow inset articulation on the quadratojugal. The L-shaped <i>quadratojugal</i> is partially fused to the quadrate near the quadrate condyle, where it approaches, but does not contribute to, the jaw articulation. The suture with the quadrate shaft is relatively straight, and surface texture is low and limited to the anterior portion of the bone. <b>Palate.</b> The configuration of palatal sutures, shape and position of the suborbital fenestra, form of the mandibular rami of the pterygoid and ectopterygoid, position of the choanae, and form of the choanal septum (Figs. 5C, 6C) correspond well with those of <i>Araripesuchus</i> (Price 1959) (Figs. 14C, 15C) and differ markedly from the palatal configuration described in <i>Simosuchus</i> (Buckley et al. 2000). In these regards, <i>A. minor</i> is less derived than <i>Simosuchus</i>. The premaxillary portion of the palate is restricted to a broad-based triangle near the anterior margin. Th e premaxilla-maxilla suture, however, is exposed only near the alveolar margin. Th e premaxilla-maxilla foramen may communicate with the palate as in <i>A. wegeneri</i>; a foramen is present at the anterior margin of the maxilla just posterior to the premaxilla-maxilla suture, as is the case on one side of a cranium of <i>A. wegeneri</i> (Figs. 14C, 15C). Furthermore, as in another skull of that species (Fig. 20B), this palatal foramen appears to be associated with the tip of the fourth dentary crown (MNN GAD 17, GAD 603) The maxilla and <i>palatine</i> form the majority of the palate in <i>A. minor</i> (Figs. 5C, 6C). Th e median one-third appears to preserve its natural arching toward the midline, whereas the lateral one-third on each side lies closer to the horizontal. Neither the vomer nor pterygoid are exposed in the midline as in <i>Simosuchus</i> (Buckley et al. 2000). A slit-shaped foramen opens on the maxilla. Canted along an anterolateral-posteromedial axis, opening anterolaterally, and associated with a small palatal fossa, the foramen is far from the alveolar margin and may not correspond to maxillary foramina associated with the alveolar margin in other notosuchians. The <i>pterygoid</i> and <i>ectopterygoid</i> form the posterior portion of the palate, including the posteroventrally projecting mandibular rami. Th e distal end of this process is modestly expanded as in <i>Araripesuchus</i> and lies in its natural position adjacent to the adductor fossa of the lower jaw. The ectopterygoid overlaps the ventral aspect of the pterygoid on the lateral edge of the palate. The suborbital fenestra, which is best exposed in the subadult skull (Sereno et al. 2003), is subequal in size to the paired choanae and located farther anteriorly. The palatine-pterygoid suture, preserved on the right side, courses across a broad palatal border lateral to the choanae. In the midline of the adult skull, the posterior one-half of the very thin choanal septum is exposed, the remainder covered from view by extraneous bone pieces. Th e choanae are located as far posterior on the pterygoids as possible, butting against a posterior palatal ridge formed by the pterygoids. During growth the sigmoid curve of the posterior palatal ridge in the subadult becomes a broad arch in the adult (Figs. 5C, 6C). Unlike in some other species of <i>Araripesuchus</i> (<i>A. gomesii, A. wegeneri</i>), there is no development of a pair of parasagittal flanges extending from the posterior palatal ridge. The <i>quadrate</i> angles posteroventrally from the recessed otic region toward the quadrate condyles. In the otic region, a large opening constitutes the fenestra ovalis and confluent cranioquadrate passage. Anterior to this opening is the preotic siphonium, ventral to which is a circular fossa (Fig. 7D) as in <i>Araripesuchus wegeneri</i>. A sharp vertical crest on the quadrate contributes to the posterior skull margin, joining the paroccipital process with the rim of the medial condyle. In posterior view, a foramen aërum opens on the posterior aspect of the quadrate shaft just above the medial condyle (Fig. 8). In lateral view, the posterior margin of the quadrate angles anteroventrally as in <i>Simosuchus</i> (Buckley et al. 2000) rather than posteroventrally as in nearly all other crocodyliforms. Th e quadrate condyles are relatively flat and separated by a marked V-shaped cleft (Fig. 7D). <b>Braincase.</b> The braincase is well preserved and exposed in the holotype and referred skull
description: Published as part of <i>Sereno, Paul &amp; Larsson, Hans, 2009, Cretaceous Crocodyliforms from the Sahara, pp. 1-143 in ZooKeys 28 (28)</i> on pages 8-33, DOI: 10.3897/zookeys.28.325, http://zenodo.org/record/576570
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