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Alamosaurus

Alamosaurus sanjuanensis is a species of titanosaurian sauropod dinosaur known from the late Maastrichtian stage of the Late Cretaceous period, approximately 66 million years ago, in southwestern North America.^[1]^[2] Named in 1922 by Charles W. Gilmore based on a holotype consisting of a left scapula and a paratype right ischium collected from the Ojo Alamo Formation in the San Juan Basin of New Mexico, the genus derives its name from this formation, reflecting its initial discovery site.^[3] Subsequent discoveries have expanded the known fossil record to include more complete material, such as an articulated series of cervical vertebrae from Big Bend National Park in Texas and an incomplete skeleton from southern Utah, revealing a long-necked herbivore with robust limb bones and a whip-like tail typical of advanced titanosaurs.^[1] Phylogenetic analyses place Alamosaurus within the Lithostrotia clade of Titanosauria, potentially allied with the South American Lognkosauria group rather than the more derived Saltasauridae, suggesting possible migratory connections between North and South America via a late Cretaceous land bridge.^[1] Fossils have been recovered from multiple formations across New Mexico, Texas, Utah, and possibly Wyoming and Mexico, indicating a wide distribution in the final dinosaur ecosystems of the continent.^[1] Size estimates for adults vary, but reconstructions based on limb elements suggest a body length of around 26 meters and a height of 5 meters at the shoulder, with some specimens implying masses up to 75 tons, making it one of the largest terrestrial animals ever to inhabit North America.^[1]^[4] As the only conclusively identified titanosaurian sauropod in North America during the Late Cretaceous, Alamosaurus marks the end of the "sauropod hiatus"—a gap in large sauropod records from the mid-Cretaceous—and underscores the resilience of dinosaur faunas right up to the Cretaceous-Paleogene boundary.^[1] Recent high-precision U-Pb dating of associated sediments confirms that Alamosaurus and other non-avian dinosaurs thrived in diverse communities without signs of decline until the asteroid impact approximately 66 million years ago.^[2] Its osteology, including somphospondylous vertebrae, provides insights into the adaptations of giant herbivores in the final phases of the Mesozoic era.^[1]

Anatomy

Osteology

The osteology of Alamosaurus sanjuanensis is known from numerous disarticulated and partially articulated skeletal elements recovered primarily from Maastrichtian formations in the southwestern United States, providing a detailed but incomplete picture of its titanosaurian body plan. The axial skeleton features robust vertebrae with pronounced pneumaticity, characteristic of advanced titanosaurs. Cervical vertebrae are elongate and opisthocoelous, with centra that are wider than tall and exhibit broad lateral fossae containing pneumatic pleurocoels; an articulated series of nine posterior cervicals (estimated as C6–C14) from a mature individual shows increasing centrum length toward the mid-neck before a posterior decrease, with the longest centra measuring approximately 800 mm. Neural spines are tall and triangular in lateral view, expanding laterally in posterior cervicals to become wider than the centra, differing from the lower spines in more basal sauropods but aligning with those of other non-lithostrotian titanosaurs like Futalognkosaurus dukei. The tentative cervical count is 14, fewer than the 17 in Rapetosaurus krausei but comparable to other lognkosaur-grade titanosaurs. In 2025, a large vertebra was recovered from Big Bend National Park, potentially associated with previously known material and contributing to the understanding of its axial skeleton.^[5] Dorsal vertebrae display camellate internal pneumaticity and spatulate neural spines with strong prespinal laminae, lacking hyposphene-hypantrum articulations typical of some diplodocoids. Caudal vertebrae are procoelous throughout, with wide, robust anterior centra indicating a deep tail base for muscular support and elongated centra in the mid-to-posterior regions; these features closely resemble those of Neuquensaurus australis, supporting a shared lognkosaurian affinity, though Alamosaurus lacks the deep lateral fossae seen in some saltasaurids. Appendicular elements underscore a quadrupedal stance with robust limb proportions suited to bearing immense body mass. The humerus is slender yet strongly constructed, with lengths up to ~1.5 m in known adults (estimates ~2 m for largest individuals), featuring a ventrally extended ulnar facet and deltopectoral crest similar to Neuquensaurus; the femur, proportionally longer than the humerus (ratio ~1.1:1), has known lengths of ~1.6 m (estimates up to ~2.2 m), with a straight shaft and fourth trochanter positioned distally as in other titanosaurs.^[6]^[7] No cranial material has been recovered for Alamosaurus, leaving its skull morphology inferred from closely related titanosaurs such as Rapetosaurus and Saltasaurus, which possessed boxy skulls with elongated snouts adapted for cropping low vegetation. Isolated teeth attributed to Alamosaurus are rod-shaped, peg-like structures with finely wrinkled enamel and prominent apicobasal wear facets on the labial and lingual surfaces, indicative of a diet of tough, fibrous plants processed by dental batteries similar to those in other titanosaurs. These teeth, measuring up to 10 cm in height, lack the spoon-shaped crowns of more basal sauropods, aligning with derived titanosaur dentition. Evidence for dermal armor includes osteoderm fragments associated with a partial Alamosaurus skeleton from Utah, consisting of bulb-and-root morphotypes with irregular, keeled surfaces up to 20 cm in length; these are comparable to those in armored titanosaurs like Saltasaurus but smaller relative to body size, suggesting sporadic distribution along the back and flanks rather than full-body plating. No skin impressions have been reported, though the presence of osteoderms implies a textured integument consistent with titanosaur relatives. Limb proportions, with forelimbs approximately 80% the length of hindlimbs, further confirm a stable quadrupedal gait akin to that of Dreadnoughtus and other large titanosaurs.

Size and growth

Alamosaurus sanjuanensis was one of the largest dinosaurs known from North America, with total body length estimates ranging from 21 to 30 meters based on partial skeletons and scaling from associated skeletal elements.^[8]^[9] A reconstructed composite skeleton incorporating an articulated cervical series from Big Bend National Park, Texas, yields a length of approximately 26 meters.^[1] Recent discoveries of large limb bones and vertebrae from the same locality suggest some individuals reached up to 30 meters, representing the upper end of the size range for this taxon.^[9] Body mass estimates for adult Alamosaurus vary between 30 and 80 metric tons, derived from volumetric models of 3D reconstructions and scaling equations applied to limb bone dimensions.^[10]^[11] For instance, specimens such as USNM 10486 and associated material have been modeled using profile-based skeletal restorations scaled to known titanosaur proportions, yielding masses around 30-35 tons with a specific gravity adjustment for soft tissues.^[11] Higher estimates, up to 80 tons, incorporate fragmentary large elements like partial femora and tibiae from Big Bend, where limb bone circumferences (e.g., femur midshaft diameter exceeding 50 cm) correlate with body masses over 50 tons via established sauropod scaling relationships that link bone robustness to supporting weight.^[10]^[9] Shoulder height is inferred at 5-6 meters from humerus and scapula proportions in multiple specimens, with the scapula of the holotype (USNM 10486) measuring 1.55 meters long, scaled against related titanosaurs.^[1]^[8] Growth in Alamosaurus was rapid during juvenility, with bone histology revealing lines of arrested growth (LAGs) in limb bones indicative of seasonal pauses but overall fast deposition rates.^[12] Analysis of humeri and femora from Big Bend specimens shows maximum growth rates approaching 1,000-1,090 kg per year during peak ontogeny, allowing individuals to reach adult sizes exceeding 30 tons within approximately 45 years.^[10]^[13] LAG counts in cortical bone sections confirm this timeline, with outer laminar fibrolamellar tissue in mature femora suggesting sustained high growth into adulthood before slowing.^[12]^[14] Ontogenetic variation is evident in comparisons between juvenile and adult specimens, highlighting allometric changes such as disproportionate neck elongation in maturity.^[8] A juvenile individual from Big Bend National Park, represented by partial ischia, caudal vertebrae, and limb elements, exhibits a higher humerus-to-femur length ratio (0.77) compared to adults (0.65), indicating shifts in limb proportions during development.^[8] Smaller vertebrae from a partial skeleton (USNM 15560) collected in Utah further demonstrate these differences, with more compact cervical dimensions relative to body size than in large Texas adults, consistent with accelerated elongation of the neck in later growth stages.^[15]

Discovery

History of research

The history of research on Alamosaurus began in June 1921 when John B. Reeside Jr., a geologist with the U.S. Geological Survey, discovered the holotype specimen—a large left scapula (USNM 10486)—in the Ojo Alamo Formation of San Juan County, New Mexico, during fieldwork related to coal exploration.^[16] This find represented the first evidence of a Late Cretaceous sauropod in southwestern North America. In 1922, Smithsonian Institution paleontologist Charles W. Gilmore formally described and named the new genus and species Alamosaurus sanjuanensis, based on the scapula and an associated right ischium (USNM 10487, paratype) from the same locality; Gilmore noted its titanosaur affinities but had limited material for detailed comparisons. During the 1930s and 1940s, additional fossils expanded the known distribution of Alamosaurus into Texas. In 1940, Barnum Brown of the American Museum of Natural History, accompanied by Roland T. Bird, led an expedition to the Big Bend region where they collected several large dorsal and caudal vertebrae, as well as limb elements, from the Javelina Formation; these specimens confirmed the presence of a massive sauropod in Maastrichtian deposits and were among the most complete early finds.^[17] Brown's discoveries highlighted Alamosaurus as a key taxon bridging the apparent "sauropod hiatus"—a perceived 30-million-year gap in North American sauropod records from the Early to Late Cretaceous—prompting initial discussions on their persistence in southern latitudes.^[18] Mid-20th-century studies from the 1960s to 1980s focused on referrals of fragmentary material from other regions, particularly Utah, and deepened debates over the sauropod hiatus. Fossils from the North Horn Formation, initially collected during a 1937 Smithsonian expedition led by Charles W. Gilmore, were referred to Alamosaurus in the 1980s, including caudal vertebrae and limb bones that supported its identification and suggested a wider Laramidian distribution.^[6] These referrals, notably by Spencer G. Lucas and colleagues in 1987, played a pivotal role in resolving the hiatus controversy by demonstrating that titanosaurs like Alamosaurus had reoccupied North American ecosystems after a mid-Cretaceous decline, likely migrating from South America. In the late 20th and early 21st centuries, excavations in Big Bend National Park, Texas, produced more substantial skeletal material, transforming Alamosaurus from a poorly known taxon into a well-represented Late Cretaceous giant. Starting in the 1990s, teams from the Texas Memorial Museum and National Park Service unearthed partial skeletons, including a juvenile specimen (TMM 43621-1) from the Javelina Formation collected in 1979 but fully described later.^[8] In the 2010s, description of an articulated series of nine cervical vertebrae (BIBE 45854) from the lower Black Peaks Formation, discovered in 1997 and published in 2016 by Steven J. Jasinski and colleagues, offered the first detailed view of the neck anatomy, revealing robust neural arches and pneumatic features consistent with derived titanosaurs.^[1] Recent research in 2025 has further illuminated Alamosaurus populations near the Cretaceous-Paleogene boundary through targeted excavations in New Mexico. A multidisciplinary team, including paleontologists from the University of Nebraska-Lincoln and collaborators supported by National Geographic, analyzed new fossils from the Naashoibito Member of the Ojo Alamo Formation, including partial Alamosaurus skeletons with vertebrae and limb elements alongside other dinosaurs; precise U-Pb dating placed these assemblages at approximately 66.0 to 66.4 million years old, confirming thriving sauropod communities persisted until moments before the end-Cretaceous extinction.^[2] This study, published in Science on October 23, 2025, underscores Alamosaurus as a resilient apex herbivore in the final Maastrichtian ecosystems of the American Southwest.^[19]

Geological context

Alamosaurus fossils are primarily preserved in the Naashoibito Member of the Ojo Alamo Formation in New Mexico, the Javelina and Black Peaks Formations in Texas, and the North Horn Formation in Utah. These units consist predominantly of sandstones and mudstones deposited in fluvial environments, including channel fills, overbank floodplains, and shallow ponds that reflect low-energy river systems with periodic flooding.^[20]^[21]^[22] The depositional age of these formations falls within the Maastrichtian stage of the Late Cretaceous, spanning approximately 70 to 66 million years ago. Recent U-Pb dating constrains the Naashoibito Member of the Ojo Alamo Formation to approximately 66.4 to 66.0 Ma, positioning Alamosaurus fossils very close to the Chicxulub impact dated to 66.04 Ma.^[2]^[23]^[24] Alamosaurus represents the return of sauropod dinosaurs to North America following a prolonged absence known as the sauropod hiatus, during which no sauropod fossils are recorded from the Cenomanian to Campanian stages (approximately 95 to 72 Ma). This hiatus ended in the latest Cretaceous with the appearance of Alamosaurus, likely an immigrant from South America.^[25]^[26] Taphonomic evidence indicates that Alamosaurus remains are typically disarticulated and occur in floodplain deposits, suggesting transport after death by fluvial processes rather than in situ preservation. No mass death assemblages have been identified, implying isolated mortality events rather than catastrophic group deaths.^[8]^[27] Stratigraphically, the formations bearing Alamosaurus correlate to the global Maastrichtian stage and share temporal overlap with the Hell Creek Formation in the northern Western Interior, but reflect a distinct southern Laramidian faunal province characterized by different vertebrate assemblages.^[23]^[2]

Taxonomy

Classification

Alamosaurus sanjuanensis is the sole recognized species within the genus Alamosaurus, a titanosaurian sauropod dinosaur from the Late Cretaceous of southwestern North America. The genus name derives from the Ojo Alamo Formation, the geologic unit yielding the type material, where "Alamo" alludes to the Spanish term for "cottonwood eye" referencing a local landmark, combined with the Greek "saurus" for lizard; the specific epithet honors San Juan County, New Mexico, the type locality.^[28] The type specimen, designated USNM 10486, consists of a nearly complete left scapula measuring approximately 155 cm in preserved length, while the paratype USNM 10487 is a right ischium about 81 cm long; both were collected in 1921 from sandstones of the Ojo Alamo Formation near Barrel Spring Arroyo in San Juan County.^[28] No junior synonyms have been proposed for the taxon. Referred specimens include partial skeletons and isolated elements from coeval Maastrichtian strata in Texas, such as a series of articulated cervical vertebrae from Big Bend National Park, broadening the known hypodigm beyond the sparse type material. In higher-level taxonomy, Alamosaurus belongs to the sauropod clade Titanosauria, characterized by features such as procoelous caudal vertebrae and robust limb elements adapted for weight support in large-bodied herbivores. It is often placed within Lithostrotia, a derived titanosaurian subgroup including taxa with extensive skeletal pneumatization, though its precise familial affinities remain unresolved and debated, with placements varying from basal Lithostrotia to outside Saltasauridae in different analyses. The validity of Alamosaurus as a monospecific genus has been debated, with ongoing discussions about whether it functions as a wastebasket taxon, potentially encompassing multiple distinct Maastrichtian titanosaurians from the southwestern United States due to limited diagnostic material and historical lumping of fragmentary remains. A 2025 study proposed Utetitan zellaguymondeweyae as a new genus for some previously referred material from Utah and Texas, further highlighting geographic and stratigraphic variations.^[29]

Phylogeny

Alamosaurus is recognized as a derived member of Titanosauria, exhibiting key synapomorphies such as procoelous caudal vertebrae and robust limb elements that align it with advanced titanosaurs.^[1] Phylogenetic studies have variably positioned it within Opisthocoelicaudiinae, a subclade of Saltasauridae, or as basal to Lognkosauria, a group of gigantic South American titanosaurs including Futalognkosaurus and Patagotitan.^[1] These placements are supported by shared features like extensive pneumaticity in the vertebral column and a wide-gauge posture indicated by limb proportions.^[30] A 2016 cladistic analysis incorporating new cervical material recovered Alamosaurus as the sister taxon to Lognkosauria, emphasizing resemblances in neck morphology and pneumatic features to South American forms like Patagotitan.^[1] Earlier studies, such as those by Zaher et al. (2011), had placed it as sister to Saltasauridae, highlighting affinities with armored titanosaurs like Saltasaurus based on overall lithostrotian traits.^[1] More recent 2020s analyses, including Otero et al. (2021), refine this by positioning Alamosaurus in a clade with the Argentine titanosaur Pellegrinisaurus, basal to Saltasauridae, and underscore vertebral pneumaticity patterns that link it closely to Patagotitan and other lognkosaurs.^[31] The evolutionary history of Alamosaurus supports a migration hypothesis from Gondwanan origins, with ancestors likely dispersing northward from South America into Laramidia during the late Campanian to early Maastrichtian via temporary land bridges, thereby ending the long-standing sauropod hiatus in North American fossil records.^[1] This dispersal is evidenced by biogeographic patterns in titanosaur distributions and the abrupt appearance of advanced lithostrotians in southwestern North America after a ~30-million-year absence of sauropods.^[32] Phylogenetic placement of Alamosaurus remains inconsistent across datasets, with some analyses recovering it as relatively basal within Titanosauriformes due to plesiomorphic cranial and axial features, while others nest it deeply within Colossosauria, a diverse South American titanosaur radiation.^[1] These discrepancies arise from varying character scorings and taxon sampling, particularly in handling fragmentary North American material.^[31] As the dominant and final sauropod in Late Cretaceous North America, Alamosaurus exemplifies the last major radiation of Titanosauria on the continent, contributing to high sauropod diversity in Laramidia just prior to the end-Cretaceous mass extinction.^[33] Its presence highlights a late pulse of titanosaurian evolution that bridged Gondwanan and Laurasian faunas, influencing regional paleoecological dynamics.^[1]

Biogeography and Paleoecology

Distribution

Fossils of Alamosaurus sanjuanensis are primarily known from localities in the southwestern United States, reflecting its endemism to southern Laramidia during the Maastrichtian stage of the Late Cretaceous. The holotype specimen, consisting of a partial scapula and ischium, was discovered in the Ojo Alamo Formation of the San Juan Basin in northwestern New Mexico, marking the type locality for the taxon.^[34] Additional significant finds come from the Big Bend region of West Texas, where remains have been recovered from the Javelina, Black Peaks, and El Picacho Formations, including partial skeletons and isolated elements such as vertebrae, limb bones, and pelvic girdle material.^[9]^[23] In Utah, fossils are documented from the North Horn Formation in central regions, though these are less common and typically consist of fragmentary material like a partial skeleton including a pelvis.^[8] Secondary or possible occurrences include debated associations with undescribed titanosaur fossils closely related to Alamosaurus from the Evanston Formation in Wyoming, but these assignments remain unconfirmed and are often reattributed to other titanosaurs or indeterminate taxa. No verified Alamosaurus fossils have been reported from Mexico or regions further south, limiting its known distribution to the North American continent north of the paleoequator.^[35] The geographic range of Alamosaurus was confined to southern Laramidia, spanning approximately 35° to 40° N paleolatitude, from central Utah southward to the Big Bend area of Texas—a north-south extent of roughly 1,000 km. This distribution contrasts sharply with the northern Laramidian fauna dominated by Triceratops, highlighting latitudinal provinciality in Late Cretaceous dinosaur communities. Abundance patterns show Alamosaurus as relatively common in Texas and New Mexico, where dozens of specimens (including multiple partial skeletons) indicate population centers in subtropical lowlands, while Utah yields only a handful of rarer finds.^[36]^[35]^[1] Biogeographically, Alamosaurus defines the "Alamosaurus fauna," a distinctive southern Maastrichtian vertebrate assemblage characterized by endemic titanosaurs and associated taxa, distinct from the northern biota featuring Edmontosaurus and Tyrannosaurus rex. This faunal partitioning underscores endemism driven by paleogeographic barriers and climatic gradients across Laramidia.^[23]^[35]

Paleoenvironment

Alamosaurus inhabited riverine floodplains and coastal deltas in the intermontane basins of the southwestern United States during the late Maastrichtian stage of the Late Cretaceous, approximately 69 to 66 million years ago. These environments featured markedly seasonal, semi-arid conditions with warm temperatures, supporting conifer-angiosperm forests interspersed with ferns and cycads. Paleoclimate reconstructions indicate mean annual temperatures exceeding 20°C.^[2]^[37] As a high-level browser, Alamosaurus primarily foraged on the upper canopy of conifers, tree ferns, and cycads, utilizing its long neck to access foliage 10–15 meters above the ground. This feeding strategy allowed it to exploit resources unavailable to shorter herbivores, with its peg-like teeth suited for cropping tough plant material rather than grinding. The abundance of specimens suggests possible gregarious behavior, with herds potentially forming for protection and foraging efficiency during migrations across floodplains.^[38] Alamosaurus coexisted with a diverse array of predators and competitors in these ecosystems, including tyrannosaurids such as Tyrannosaurus rex in northern regions and the southern T. imperator, which likely preyed on juveniles and weakened adults. It competed for vegetation with hadrosaurids like Kritosaurus and ceratopsians such as Torosaurus, partitioning resources by height and habitat preferences. The broader community included azhdarchid pterosaurs, crocodylomorphs, and multituberculate mammals, reflecting a thriving, regionally endemic fauna.^[37] Recent geochronological studies date Alamosaurus-bearing strata to as young as 66.34 million years ago, indicating it persisted until the final moments before the Cretaceous-Paleogene (K-Pg) boundary impact. A 2025 study further refines the dating of the Naashoibito Member to approximately 66.04 million years ago, indicating Alamosaurus persisted in a diverse, stable ecosystem until the K-Pg boundary.^[2]^[39] Bone histology reveals continuous, rapid growth with lines of arrested growth but no pronounced stress markers, such as dense vascular disruption or resorption lines indicative of environmental instability, supporting a stable paleoenvironment without pre-impact decline. This contradicts earlier hypotheses of faunal impoverishment, highlighting robust diversity right up to the extinction event.^[2]

References

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