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Aucasaurus

Aucasaurus garridoi is a species of medium-sized abelisaurid theropod known from a nearly complete subadult discovered , . It lived during the early stage of the epoch, approximately 83 million years ago, in what is now the Neuquén Basin. The genus name refers to the nearby Auca Mahuevo locality, a significant site for fossils including titanosaur eggs, while the specific honors Argentine geologist and fossil collector Alberto Garrido. The specimen (MCF-PVPH 236), unearthed in 1999 during a joint expedition by the Museo Carmen Funes and the , preserves about 80% of the skeleton, including the , vertebrae, ribs, and limbs, making it one of the best-understood abelisaurids. Estimated at around 5–6 meters in total length, the individual is considered subadult based on some open neurocentral sutures, but bone histology indicates it had achieved somatic maturity with having ceased, as evidenced by the external fundamental . As a bipedal , A. garridoi featured a robust with short, deep , reduced forelimbs bearing two fingers, and a adapted for , traits typical of abelisaurids that suggest it was an agile predator capable of high-speed pursuits. Phylogenetically, Aucasaurus belongs to the tribe Carnotaurini within , closely related to sastrei, another Patagonian abelisaurid, based on shared features like elongated hindlimbs and a shortened . The specimen's preservation in the Anacleto Formation, a fluvial deposit, indicates it inhabited a semi-arid environment with rivers and forests, where it likely preyed on ornithopods and smaller theropods alongside abundant sauropods. Recent analyses of its have further refined its phylogenetic position within . Its discovery has advanced understanding of abelisaurid diversity and development in Gondwanan ecosystems during the .

Discovery

Geological context

The Aucasaurus fossils were recovered from the Anacleto Formation, the uppermost unit of the Neuquén Group within the Río Colorado Subgroup of the Neuquén Basin in Patagonia, . This formation dates to the lower stage of the , approximately 83–80 million years ago. It consists primarily of purple to reddish-brown siltstones, mudstones, and fine-grained sandstones, with siliceous and calcareous concretions, representing deposits from low-energy fluvial systems and extensive alluvial plains. The prevalence of indicates a with seasonal river activity and periodic flooding on floodplains. The stratigraphic position of the Anacleto Formation has been refined through magnetostratigraphic correlation to polarity zone Chron C33R, confirming its early age. Recent U-Pb detrital dating further supports a maximum depositional age around 78–80 Ma, aligning with the broader infill of the Basin during this interval. The formation's thickness varies from 50 to 90 meters, with outcrops primarily in the provinces of , Río Negro, and Mendoza, reflecting a of meandering rivers in a continental setting influenced by tectonic . The Anacleto Formation hosts a diverse vertebrate assemblage, including co-occurring dinosaurs such as titanosaurian sauropods like Rinconsaurus caudamirus, indeterminate forms, and the recently described Chadititan calvoi, other abelisaurid theropods, and ornithopods such as Gasparinisaura. The nearby Auca Mahuevo locality within the formation is renowned for its titanosaur nesting sites, yielding hundreds of megaloolithid eggs and embryos that highlight the reproductive behavior and ecological dominance of sauropods in this ecosystem. This faunal diversity underscores a complex terrestrial community in a floodplain habitat supporting large herbivores, predators, and smaller vertebrates like crocodylomorphs and .

Fossil material

The specimen of Aucasaurus garridoi, designated MCF-PVPH-236, was discovered in 1999 by Argentine Alberto Garrido at the Auca Mahuevo locality in , . The find occurred during fieldwork in the badlands of the Anacleto Formation, where the was partially exposed on the surface. It was subsequently excavated and prepared by a joint Argentine-American team led by paleontologists Rodolfo A. Coria and Luis M. Chiappe, in collaboration with Lowell Dingus.022[0460:ANCROC]2.0.CO;2.short) The specimen, representing a nearly complete of a subadult individual, is housed at the Museo Carmen Funes in Plaza Huincul, . Osteohistological analysis of elements including the and indicates the individual attained somatic maturity at a minimum age of 11 years at death, though continued slow growth suggests it was not fully senescent. The is approximately 85% complete, preserving a significant portion of the axial column (including , , sacral, and caudal vertebrae, along with , , and haemal arches) in articulation, as well as partial fore- and hind limbs and cranial material.022[0460:ANCROC]2.0.CO;2.short) Preparation efforts revealed minimal in key regions, pointing to rapid entombment that limited post-mortem disturbance. No additional complete skeletons of Aucasaurus have been identified as of 2025. Referred material is limited to isolated abelisaurid elements from the Anacleto Formation, such as the partial , vertebrae fragments, humeri, pubis, and pedal ungual comprising specimen MPCN-PV 69, which may pertain to a distinct within . Taphonomic features, including the lack of bite marks or extensive weathering on the bones, support deposition in a low-energy setting conducive to quick burial and preservation of articulated structures.

Description

General morphology

Aucasaurus garridoi was a medium-sized, bipedal theropod within the family, exhibiting a robust yet relatively gracile build compared to bulkier relatives such as sastrei. The specimen (MCF-PVPH-236), a nearly complete from the Anacleto Formation in , , indicates an overall body length of approximately 5–6 meters and an estimated body mass of around 180 kg for this subadult individual, calculated using femoral circumference measurements. This size places Aucasaurus as shorter than Carnotaurus (estimated at 7.5–9 meters) but similarly stocky in its hindlimb-dominated , with a adapted for terrestrial predation in a environment. Osteohistological evidence indicates continued rapid growth, suggesting adults may have reached larger sizes. Distinguishing external features include a shallower and more elongated profile relative to the deeper, shorter rostrum of , contributing to a less robust cranial appearance overall. The supraorbital ridges are low and rounded, lacking the prominent, horn-like projections seen in and some other abelisaurids, which may have reduced the emphasis on display structures. Forelimbs are notably reduced in size, typical of abelisaurids, but proportionally longer than those of , with the reaching about 15–20% of length, suggesting slightly greater potential for manipulative functions despite their diminutive scale. The postcranial further highlights a balanced, stocky supported by powerful hindlimbs, while the elongated caudal series—comprising over 30 vertebrae—likely enhanced balance and agility during pursuits. No confirmed evidence of exists in the known fossil material of Aucasaurus, as the single well-preserved provides limited sample size for assessing intraspecific variation. However, the 's proportions, including its relatively elongated forelimbs and less exaggerated cranial ornamentation, have been interpreted as potentially retaining juvenile-like traits into maturity, hinting at ontogenetic stability in rather than marked allometric changes. These metrics underscore Aucasaurus's position as a versatile predator, with indicators like the extended tail supporting maneuvers in diverse terrains. Body mass estimates for the specifically, derived from femoral circumference measurements, suggest a range of 178–180 , indicating the specimen represents a subadult individual.

Cranial and braincase features

The skull of Aucasaurus garridoi is partially preserved, featuring a characteristically short and deep rostrum typical of abelisaurids, which contrasts with the more elongate snouts of basal theropods. This rostrum houses a reduced , a derived among abelisaurids that minimizes the size of the opening while maintaining structural integrity for biting forces. Unlike relatives such as sastrei, which possess prominent supraorbital horns, Aucasaurus exhibits low supraorbital crests, providing a less ornamented but potentially lighter cranial profile. The braincase of Aucasaurus, examined through CT scanning of the holotype specimen (MCF-PVPH-236), reveals endocranial features closely resembling those of Majungasaurus crenatissimus, including a compact forebrain, midbrain, and hindbrain configuration. A notably large floccular process, larger than in Majungasaurus and enclosed within an 8-shaped recess, suggests adaptations for enhanced balance and oculomotor control during rapid head movements. The flexible atlanto-occipital joint further indicates a wider range of head mobility compared to more rigid-jointed theropods, potentially aiding in prey detection and capture. Sensory structures within the braincase highlight specialized adaptations; the olfactory bulbs are large, oval, and divergent, with an olfactory tract measuring approximately 30 mm in length and 15 mm in width, proportions that imply acute olfactory capabilities for tracking prey or carrion in its environment. The semicircular canals of the inner ear, while similar overall to those of Majungasaurus, feature a relatively shorter lateral canal, yet their configuration supports enhanced vestibular function, facilitating quick turns and agile maneuvers. There is no anatomical evidence for venom delivery mechanisms, such as grooved teeth or specialized glands, distinguishing Aucasaurus from certain other theropods hypothesized to possess such traits. In comparisons to other abelisaurids, the cranium of Aucasaurus is less robust than that of , with further reductions in fenestral openings representing a more derived condition within the , possibly linked to increased skull strength for terrestrial predation. The endocranial of approximately 47.6 cm³ is notably smaller than the ~70% larger estimated for , consistent with Aucasaurus representing a subadult individual.

Postcranial skeleton

The of Aucasaurus garridoi is characterized by robust adapted for supporting the relatively heavy typical of abelisaurids. The middle exhibit well-developed epipophyses with prominent anterior and posterior processes, providing attachment sites for strong neck muscles such as the m. complexus and m. longus colli dorsalis, which facilitate head stabilization and movement. The consists of six fused vertebrae, forming a rigid structure with continuous neural spines and robust transverse processes that enhance pelvic stability and weight transfer during locomotion. In the caudal series, the 12th and 13th vertebrae are notably elongated, featuring prominent chevron facets that indicate robust attachment for haemal arches and strong tail musculature, contributing to and . The forelimbs of Aucasaurus are extremely reduced, consistent with the trend in derived abelisaurids toward vestigial upper limbs. The is significantly shorter than the , measuring approximately 15-20% of the length, with a deltopectoral crest that is poorly developed..022[0068:ANCROC]2.0.CO;2) The manus retains four metacarpals, but only digits II and III are functional, lacking phalanges or claws on the others, which limits manipulative capabilities..022[0068:ANCROC]2.0.CO;2) Osteohistological analysis of the reveals a dominated by with a woven-fibered matrix, indicative of rapid, uninterrupted growth during despite the limb's small size. The hindlimbs exhibit adaptations suggestive of locomotion, with a gracile and that are proportionally long relative to the body, enabling efficient strides. The features a straight shaft and a reduced fourth , while the has a prominent, hooked cnemial for muscle leverage. The pes is tridactyl, with three functional toes (-), and displays an arctometatarsal condition where metatarsal III is pinched proximally between II and IV, optimizing and foot flexibility during rapid movement. The is correspondingly reduced, with a short, slender that correlates with the diminutive forelimbs and minimal mobility..022[0068:ANCROC]2.0.CO;2) In contrast, the supports more robust function, featuring a broad, laterally flared ilium with an elongated preacetabular process that accommodates powerful extensors for forceful strides. The and pubis are robust, with the pubis showing a slight twist distally, contributing to overall pelvic stability..022[0068:ANCROC]2.0.CO;2)

Pathological features

The holotype specimen of Aucasaurus garridoi (MCF-PVPH-236) displays a prominent pathological condition in its axial skeleton, characterized by the complete fusion of the fifth and sixth caudal vertebrae, along with the associated fifth haemal arch. This anomaly results in block vertebrae, a congenital malformation arising from disrupted somitogenesis during early embryonic development, where individual vertebral precursors fail to segment properly. Computed tomography scans of the fused elements reveal smooth sutural surfaces without evidence of bone spurs, proliferative new bone formation, or inflammatory remodeling, distinguishing it from traumatic or infectious origins and confirming its antemortem, developmental nature. This marks the earliest documented case of congenital vertebral in non-avian theropods, with prior records limited to more derived tetanurans and ornithischians. In the subadult , estimated at least 11 years old based on osteohistological , the occurs in the proximal region, potentially reflecting localized genetic or environmental influences on formation without broader skeletal impacts. Although the block structure slightly alters local flexibility, the overall and lack of associated deformities suggest negligible effects on or predatory . Osteohistological examination of the holotype's long bones, including the , reveals secondary remodeling with Haversian systems in the outer , indicative of localized mechanical stress or minor trauma response during growth, though no overt fractures or bite traces are preserved. This contrasts with taphonomic alterations elsewhere in the specimen, such as postmortem cracking, emphasizing the antemortem origin of the vertebral fusion. In comparison to other theropods, similar congenital fusions appear sporadically in tyrannosaurids like Tyrannosaurus rex but remain undocumented in abelisaurids beyond this instance, highlighting A. garridoi as a unique case among South American ceratosaurs and underscoring variability in developmental health within the group.

Classification

Relationships within Abelisauridae

Aucasaurus garridoi was formally named and described in 2002 by Rodolfo A. Coria, Luis M. Chiappe, and Lowell Dingus, based on a nearly complete subadult skeleton recovered from the Auca Mahuevo locality in Patagonia, Argentina. The genus name combines "Auca," from the Mapudungun language in reference to the Auca Mahuevo locality, with the Greek "saurus" for lizard. The specific epithet honors Alberto Garrido, the and fossil collector who discovered the specimen. The initial description, published in the Journal of Vertebrate , established Aucasaurus as a new member of , highlighting its close affinity to Carnotaurus sastrei within the tribe Carnotaurini, supported by shared derived traits such as reduced forelimbs and specialized caudal vertebrae. Early phylogenetic analyses positioned it as the sister taxon to , emphasizing synapomorphies in the and limb reduction. A brief taxonomic debate arose in 2010 when proposed synonymizing Aucasaurus with Abelisaurus comahuensis due to similarities in fragmentary cranial material, but this has not gained acceptance; distinctions in humeral morphology, including differences in the development of the deltopectoral crest, support their separation. Subsequent cladistic studies have refined its placement within . A 2023 phylogenetic analysis incorporating an expanded character matrix recovered Aucasaurus within the derived Brachyrostra, in a with Carnotaurus and other South American abelisaurids such as Viavenator and Llukalkan, based on shared brachyrostran apomorphies like robust neural arches and increased axial rigidity. This positioning underscores its role in Carnotaurini, characterized by extreme reduction and ornate cranial features. No major taxonomic revisions have occurred as of 2025, maintaining its validity as a distinct . Indeterminate abelisaurid fossils from the Anacleto Formation, including the partial skeleton MPCN-PV 69 (comprising premaxilla, vertebrae, humeri, and pubis), share stratigraphic provenance with Aucasaurus and exhibit comparable humeral proportions, leading to suggestions of possible congenericity; however, subtle differences in forelimb robusticity prevent formal assignment.

Comparisons to other abelisaurids

Aucasaurus garridoi is notably smaller than its close relative Carnotaurus sastrei, with body size estimates indicating it was approximately 30% smaller overall, reaching lengths of about 5.5–6 meters compared to Carnotaurus's 8–9 meters. Unlike Carnotaurus, which preserves extensive skin impressions revealing a mosaic of small scales and larger osteoderms, no such soft tissue evidence is known for Aucasaurus. In cranial morphology, Aucasaurus exhibits a shallower, longer rostrum and a sigmoidal dentigerous margin, contrasting with the taller, more convex rostrum of Carnotaurus; additionally, Aucasaurus lacks the prominent supraorbital horns present in Carnotaurus, instead showing subtle frontal swells. Postcranially, Aucasaurus has relatively longer forelimbs despite their reduction, with a slender, compressed humerus featuring well-defined condyles and an ulna about one-third the humerus length, compared to the shorter, stouter forelimb elements in Carnotaurus where the ulna is only one-quarter the humerus length. These proportions suggest Aucasaurus may have been more agile than the bulkier Carnotaurus, particularly in cursorial capabilities inferred from its axial skeleton and limb ratios. Comparisons with crenatissimus reveal similarities in braincase structure, including comparable proportions of the , , and , as well as overall length around 160 mm, but Aucasaurus displays a slightly more closed cerebral flexure and a less developed dorsal sagittal sinus. The of Aucasaurus appears less robust overall than that of , which features a wider and more heavily sculptured cranium with pronounced maxillary and nasal ornamentation. In the , both taxa share a similar shape, though Aucasaurus has a shorter and less laterally curved lateral semicircular , potentially indicating subtle differences in head movement . Osteohistological analysis of Aucasaurus indicates a cyclical strategy with fibrolamellar bone tissue and lines of arrested , reaching maturity by at least 11 years; this pattern aligns with other Gondwanan abelisaurids but suggests relatively rapid early rates within the lineage, distinguishing it from slower-maturing forms like . Relative to Abelisaurus comahuensis, Aucasaurus possesses a smaller , with the former estimated to have had a cranium up to 85 cm long versus Aucasaurus's approximately 60 cm, though Aucasaurus is far better preserved, allowing detailed study of its . Differences in the orbital region further distinguish the two: while both exhibit low parietal eminences and wide sagittal , Aucasaurus shows more pronounced orbital bulges without hypertrophied horns, and its floccular recess is 8-shaped and similar in size to Abelisaurus but integrated differently with surrounding cranial elements, confirming their separation as distinct taxa. In broader evolutionary terms, Aucasaurus represents a transitional form in the trend of forelimb reduction among ceratosaurs, with its moderately elongate yet reduced arms bridging earlier, more robust-limbed abelisauroids and the extremely diminutive of later taxa like , as evidenced by recent analyses of evolutionary rates in . This positions Aucasaurus as emblematic of Gondwanan endemism within , highlighting rapid diversification and specialization of these predators across southern continents during the .

Paleobiology

Growth and ontogeny

The holotype specimen of Aucasaurus garridoi (MCF-PVPH 236) was estimated to be approximately 11 years old at death through osteohistological analysis involving the counting of annuli in the . This age assessment indicates that the individual had reached maturity, as evidenced by the presence of an external fundamental system (EFS) in the compacta, consisting of parallel-fibered with densely packed circumferential vascular canals. However, the specimen exhibits subadult characteristics, including unfused epiphyses on the long s, suggesting incomplete skeletal maturity. Histological examination reveals a primarily fibrolamellar structure throughout the , characteristic of rapid growth in dinosaurs, with a woven-fibered matrix and high dominated by circumferential canals in early ontogenetic stages. Growth patterns show accelerated juvenile development, marked by dense vascular networks, which slowed in later life as indicated by decreasing vascular density and the accumulation of lines of arrested growth (LAGs). These LAGs, numbering up to 11 in the sampled elements, reflect cyclical interruptions possibly linked to seasonal environmental stresses. Secondary osteonal remodeling is prominent in the caudal vertebrae, contributing to structural reinforcement during maturation. Ontogenetic changes in Aucasaurus garridoi include the retention of juvenile-like proportions, such as relatively longer fore- and hind limbs compared to other abelisaurids, potentially representing paedomorphic traits preserved into subadulthood. This , combined with the histological evidence of ongoing but decelerating growth, underscores a developmental trajectory typical of ceratosaurs, where rapid early expansion supports predatory adaptations before stabilizing in later stages.

Locomotion and behavior

Aucasaurus garridoi exhibited adaptations in its postcranial , with relatively longer and more gracile limbs compared to other abelisaurids like crenatissimus, indicating potential for agile, bipedal . The anterior caudal vertebrae feature strongly dorsally oriented ribs with interlocking tips, conferring tail rigidity that likely enhanced straight-line propulsion via a large caudofemoralis muscle for short bursts of speed, though this may have limited maneuverability during turns. A prominently enlarged floccular process in the , larger relative to that in , points to advanced vestibular input for balance and coordination, supporting rapid head movements during pursuit or navigation. The forelimbs were markedly reduced and robust, with an articulated permitting an extreme but lacking features for effective grasping or manipulation. Muscle attachment sites on the , , , and manual elements reveal retained but specialized musculature, consistent with non-locomotory roles such as intraspecific , , or sensory functions during rather than prey handling. Predatory behavior in Aucasaurus is inferred from cranial and axial features, with bite estimates for closely related abelisaurids ranging from 3 to 8 along the row, sufficient for puncturing and dismembering medium-sized prey. The suggest a flexible capable of lateral excursions, enabling slashing bites and tearing motions to process carcasses. Direct evidence for is absent, but the discovery of the Aucasaurus in proximity to extensive titanosaur nesting colonies at Auca Mahuevo implies opportunistic hunting strategies targeting vulnerable individuals or eggs in these aggregated sites.

and diet

Aucasaurus garridoi inhabited the deposits of the Anacleto Formation (early ) at Auca Mahuevo in northern , , where the consisted of low-gradient fluvial channels, overbank mudstones, and paleovertisols formed under semi-arid to sub-humid conditions with seasonal wet-dry cycles. These paleosols, characterized by slickensides and cracking, indicate periodic aridity that likely concentrated water sources and prey animals around riverine habitats during dry seasons. The site was a major titanosaur nesting ground, with clutches of 15–40 eggs preserved in surface depressions later buried by floods, suggesting colonial breeding in open, vegetated floodplains. As a carnivorous abelisaurid theropod, Aucasaurus possessed moderately recurved, labiolingually compressed teeth with well-developed serrations (denticles spaced ~10 per 5 mm) along the carinae, adaptations for slicing and tearing from prey. Its diet likely included juvenile titanosaurs, whose eggs and embryos are abundant at Auca Mahuevo, as well as small to medium-sized ornithopods such as Gasparinisaura cincosaltensis; co-occurrence of Aucasaurus remains with these nesting sites supports opportunistic predation on hatchlings. Scavenging may have supplemented hunting, given the predator's mid-sized body (estimated 5–6 m long) and the availability of carcasses in a setting prone to flash floods. Within the mid-sized theropod guild of the Anacleto Formation, Aucasaurus occupied an apex predatory niche, potentially competing with other abelisaurids for similar prey resources in a dominated by titanosaurs like Bonatitan reigi and Chadititan calvoi. It likely avoided confrontations with adult sauropods, focusing instead on smaller herbivores and young, while seasonal aridity influenced by limiting patches and intensifying predator-prey interactions around reliable water sources. The specimen exhibits a congenital involving fusion of the 5th and 6th caudal vertebrae (block vertebra), which reduced tail flexibility and may have compromised during pursuits, potentially lowering individual in this agile predator.

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