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Paralleling the caudal margin of the coracoid tuber, but cranially displaced from it, the groove has a more expanded, rounded craniodorsal portion, from which it tapers caudoventrally. The whole structure possibly represents a modified version of the subglenoid ridge seen in Maj. Indeed, as already mentioned, that part of the bone is particularly expanded laterally, with the ventrally tapering surface medial to the groove representing the subglenoid fossa Such a groove is, to our knowledge, unknown in any related dinosaur and potentially represents an autapomorphy of Ves.
A left humerus MPCO. V j; Fig. V f.
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Based on the proportions seen in Li. A partial right hand MPCO. Taken together, the forelimb elements of Ves. The humerus is a nearly straight bone, slightly bent caudally and medially within its proximal third. This differs from the laterally arching element of Mas. The proximal and distal ends are not expanded craniocaudally relative to the midline breadth of the shaft, but are lateromedially expanded to about 2.
These expansions occur along the same plane, so that the long axes of both articulations are nearly aligned. This results in the non-twisted humeral shaft, typical of noasaurids 4. However, the proximal expansion is somewhat rotated ca. This differs from both the more globular humeral head of abelisaurids 4 , 75 , 77 and the less than two times wider than long humeral head of Mas. The proximal expansion of the head is slightly medially displaced, leaving a lateral space proximal to the greater tubercle 2 , 3 , 4 , It is distally placed and positioned at about the same level as the internal tuberosity, a condition considered synapomorphic for abelisauroids As in most noasaurids 2 , 4 , 57 , the internal tuberosity is highly reduced compared to that of other abelisauroids 4 , 75 , It corresponds to the pinched medial margin of the bone, as seen in proximal view.
The deltopectoral crest starts as a faint ridge immediately distal to the greater tubercle. In general form, the deltopectoral crest is a lateromedially compressed lamina that, in cranial view, arches laterally at its mid-length. This arched portion is also the most cranially projected i. At this point, the crest has a slightly flattened cranial margin, merging smoothly into the shaft both proximally and distally.
Similarly unexpanded deltopectoral crests are typical of abelisauroids 4. The cranial surface of the humerus is smoothly depressed medial to the deltopectoral crest for the insertion of M. Likewise, the lateral surface of the crest is also depressed proximal to its point of maximal inflection, probably for the insertion of M.
Caudal to that, a straight, rugose ridge extends from the lateral margin of the greater tubercle distally along the caudolateral corner of the bone, possibly representing the insertion area of M. Its proximal end is in the same position as the caudolateral tubercle of E. Vespersaurus paranaensis also lacks the medially broad caudal tuberosity reported in a Late Cretaceous Argentine abelisauroid A small foramen is visible on the craniomedial corner of the shaft, immediately opposite to the distal end of the deltopectoral crest, as also seen in Mas.
The distal part of the humerus expands smoothly both laterally and medially. Accordingly, as also occurs in Eo. The distal condyles are separated caudally, distally, and cranially by a continuous shallow depression. This is most prominent on the cranial surface, where it forms a low, but well delimited semi-circular brachial fossa, which is absent in El. The ectepicondyle, on the other hand, is laterally smooth, lacking any marked structure. The ulnar condyle is more craniocaudally expanded than the radial condyle, bearing a more rounded distal outline.
The associated left radius MPCO. V j of Ves. This differs from the stouter element of abelisaurids 75 , 78 and even other ceratosaurs 13 , 79 , approaching instead the condition of Li. The radius represents ca. The radius is slightly arched both medially and caudally, although this morphology is enhanced by its expanded proximal and distal articulations and by the longitudinally concave cranial and lateral margins. On the contrary, the caudal and medial margins are nearly straight.
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The proximal outline is eye-shaped, with the long axis almost twice broader than the short axis, and the caudal portion more proximally expanded. The distal articulation has an ovoid outline, with the medial margin representing the distal articulation of the ulna. The articulation facets for the ulna at the distal and proximal portions of the radius are not in the same plane. In any case, the part of the distal articulation opposite to the ulnar facet is more distally expanded, as also seen in various other theropods 80 , Three digits are preserved in the recovered right hand MPCO.
The ungual phalanx and its proximally articulating element were preserved for both medialmost digits, whereas three bones were preserved in the lateralmost digit, including what is interpreted to be the terminal phalanx see below. As such, regardless of its poor preservation, the presence of a biconcave proximal ginglymoid articulation indicates that the penultimate element preserved in digit I is a phalanx, not a metacarpal as in deeply-nested abelisaurids Its proximal articulation is much more expanded, both lateromedially and on the extensor-flexor axis, than the distal articulation, and bears a subtle extensor tuber.
This expansion is more marked towards the flexor side, as seen in isolated noasaurid manual phalanges 3 , The bone is also only about 1. Further comparisons to noasaurids and abelisaurids are hampered by the highly modified manus of the latter group 75 and the lack of well-preserved hands in the former.
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The medial condyle of the distal articulation of the first phalanx is broken away, although it can be inferred that the whole articulation was flattened on the extensor-flexor axis. The ungual is not lateromedially flattened at its proximal portion, which tapper in all dimensions. It bears a distinctive, but minor ventral curvature, as reported for Mas. The ungual and its previous phalanx are preserved in manual digit two. These are respectively exposed mainly in lateral and dorsal views, precluding assessment of many details. The non-ungual phalanx is only slightly longer than lateromedially broad at its proximal end, which is also only slightly broader than the distal end and bears a subtle extensor tuber.
The distal articulation forms a symmetric ginglymus, with rounded condyles separated by a shallow extensor fossa, and a collateral pit is seen on the lateral side the medial surface is covered by sediment. Although uncommon among averostrans, a non-elongated penultimate phalanx was also reported for Li. As in that taxon, the ungual of the second digit is longer than the preceding phalanx. Its biconcave proximal articulation is broader lateromedially than dorsoventrally deep. The bone is less recurved than that of the first digit, not nearly approaching the curvature of the possible second or third manual ungual of N.
This configuration suggests that Ves. The more proximal element preserved in digit III has a planar proximal surface, with a broader than deep subtriangular outline deeper medially and laterally ponied. As such, it is interpreted as a metacarpal. The bone is twice longer than broad at mid-length, with the medial distal condyle slightly more expanded distally than the lateral.
The first phalanx is not block-shaped as in most abelisauroids 13 , 57 , being longer than twice its midline breadth. The terminal phalanx has a subtriangular proximal outline and is distally pinched, but not recurved, differing markedly from the larger recurved digit III ungual of Li. As such, although it retains a relatively elongated first phalanx, digit III of Ves. Obviously, the incompleteness of the Ves. The holotypic left ilium MPCO. V d11 Fig. The latter ala is better preserved, but lacks most of its caudal margin, as well as the bone cover of the dorsal surface.
It is strongly expanded lateromedially towards its caudal end, with a maximal breadth nearly twice that of the bone at the level of the acetabulum, resembling the condition in Mas. The ventral surface of the ala is entirely occupied by a deep funnel-shaped brevis fossa. Although possibly exaggerated by compressive deformation, such a morphology seems to represent a further development, seen in both noasaurines 3 and G. Such a connection between the brevis shelf and the supraacetabular crest via a strong ridge is typical of some early neotheropods and ceratosaurs 3 , The caudolateral portion of the dorsal surface of the brevis shelf is covered by longitudinal scars, possibly representing the origin of the flexor tibialis musculature.
Pelvic girdle elements of Vespersaurus paranaensis gen. V d11, holotype in lateral a and ventral b views. V in dorsal c , lateral d , medial e , and proximal f views. V d2; holotype in lateral view. V in ventral h , medial i , proximal j , lateral k , and dorsal l views. Anatomical abbreviations: acr, acetabular roof; am, acetabular margin; ai, acetabular insisure; ao, M. Dashed lines represent reconstructed margins. The iliac body is mostly preserved, including the supraacetabular crest, acetabular wall, as well as the pubic and ischiadic peduncles, although the distalmost end of the pubic peduncle is damaged.
The acetabulum is fully open, with vestiges of its medial wall only at the craniodorsal corner. The supraacetabular crest is well-developed and ventrally overhanging, so that the dorsal margin of the acetabular opening is hidden in lateral view, resembling the condition in Mas. In contrast, the same profile is subrectangular in El.
From the point of maximal lateral expansion, the caudal continuation of the crest becomes abruptly less laterally projected, forming the caudodorsal corner of the acetabulum and bifurcates into two lower ridges. The most conspicuous of these ridges extends caudally to join the brevis shelf, whereas a much subtler ventral extension forms the medial margin of the caudal surface of the acetabulum. Cranial to the point of maximal lateral projection, the supracetabular crest continues as a laterally expansive and ventrally overhanging flange.
It bows dorsally, following the craniodorsal contour of the acetabulum, and extends along the pubic peduncle, where it merges smoothly to its lateral surface. The acetabular roof is also exposed in ventral view in another specimen of Ves. V a , along with a partial sacral series. The exposed portion of this ilium matches the previously described bone in all details, demonstrating that the lateromedial breadth of the acetabulum is greater than half the space between the iliac pair when they are in articulation.
Unlike Mas. Although its cranial surface is broken, it is possible to infer that the pubic peduncle had a subtriangular ventral outline, with lateral, craniomedial, and caudal margins. The former of those corners is continuous with a ridge that extends caudodorsally, forming the sharp medial margin of the cranial surface of the acetabulum. The caudolateral of those corners is caudoventral to the cranioventral tip of the supraacetabular crest, which does not reach the articulation facet itself. The better preserved ischiadic peduncle is also subtriangular in distal outline, with a pointed lateral corner and a convex medial margin.
The preserved medial surface of the ilium is mostly flat, with shallow depressed areas above the peduncles and a more medially expanded area dorsal to the acetabular aperture. An ovoid depression dorsal to the ischiadic peduncle and below an oblique shelf that extends along the postacetabular ala possibly represents the fourth sacral rib articulation, as is seen in Mas. Likewise, another oblique shelf that follows the dorsal contour of the pubic peduncle probably represents the dorsal margin of the more craniocaudally extensive articulation of the second sacral rib.
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The isolated right pubis MPCO. The body is dorsoventrally expanded and lateromedially compressed. These elements are enhanced in medial view, in which two markedly convex margins meet at a nearly right angle, directly medial to a deep excavation on the articular surface. The dorsal of these convex margins represents the dorsal two-thirds of the iliac articulation, whereas the lower encompasses the lower part of that articulation, as well as the acetabular margin and the ischiadic articulation.
In proximal view, the iliac articulation arches laterally around the deep medial excavation, dorsal to which the articular facet is dorsoventrally convex. The entire proximal articulation area of the pubis is, however, more lateromedial compressed in Ves. The dorsal margin of the pubic body of Ves.
This, along with a depressed area positioned proximoventrally from it, on the lateral surface of the bone, represents the origin of M. This contrasts with the origin of this muscle in Mas. The obturator plate is incomplete ventrodistally, but it is possible to infer a rounded profile and the presence of single a suboval obturator foramen, as occurs in Mas. In medial view, the proximal and dorsal margins of the foramen in Ves.
In contrast, that surface bears a proximodistally elongated groove in Mas. Dorsal to that ridge, the proximal part of the shaft bears a subtle longitudinal groove, which extends as a shallow fossa on the medial surface of the pubic body, dorsal to the obturator plate. More distally along the shaft, that medial ridge is more expanded, forming the laminar medial contact of the pubic pair, i.
The partial left ischia MPCO. V d2 and MPCO. Both lack the distal half of the shaft as well as the ventral margin of the obturator flange and the proximal margin of the pubic peduncle. Their proximal bodies are composed of well-developed iliac and pubic peduncles, separated by a deeply incised acetabular margin that bows slightly laterally in proximal view.
The pubic peduncle is longer, but not as dorsoventrally deep as the iliac peduncle. The latter is also much more lateromedially expanded than both the pubic peduncle and the more laminar portion of the bone that extends ventrally from it. The proximal ends of both peduncles have rugose and striated outer margins, indicating the presence of ligamentous attachments. As seen in MPCO. The margins around the socket are mostly eroded in MPCO.
V , but MPCO. V d2 shows that they expand proximally, especially at the dorsal and ventral parts of the articulation, which acquires a concave lateral profile. The most conspicuous feature of the iliac peduncle is the rugose origin point of M. This is better developed than that of Mas. The tip of that projection forms a low angle between the straight dorsal margin of the iliac peduncle and the concave margin of the bone distal to that.
As such, the proximal portion of the iliac peduncle has a slight ventral lean relative to the long axis of the bone in lateral view , as is also manifested by the dorsal margin of the ischial acetabulum. Indeed, as in Mas. The ventral margin of the pubic peduncle extends distally to form a low obturator flange. As is typical of noasaurids, that flange lacks a distal notch separating it from the ischial shaft. However, a subtle distal incision is present Fig.
Likewise, a secondary notch, as seen in Mas. The latter also arches slightly ventrally in lateral view. Immediately distal to the obturator flange, the ischial shaft has a dorsoventrally elongated cross section. Yet, as it extends distally, the ventral portion of the shaft twists medially, so that the cross section becomes lateromedially elongated, although a laminar medial flange is lacking.
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Instead, the symphyseal medial articulation to the antimere is deep and striated, marking the distal half of the persevered part of the shaft. The symphyseal articulation occupies the entire dorsoventral depth of the shaft more distally, but tapers to its ventral margin as it extends proximally. The cross section of the most distally preserved part of the shaft is ovoid, with a lateromedial long axis.
A single right tibial shaft MPCO. The mid-shaft cross-section is ovoid, with a craniomedially to caudolaterally oriented long axis and a flattened craniolateral margin. More proximally, however, due to the cranial expansion of the cnemial crest, the tibial cross section is subtriangular.
Its most conspicuous feature is the sharp fibular crest, which extends along the lateral surface of the proximal part of the shaft, turning slightly caudally at its distal end.