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Hypacrosaurus

Hypacrosaurus is an extinct genus of lambeosaurine hadrosaurid dinosaur known from the Late Cretaceous period of western North America, characterized by its large size, duck-billed snout, and distinctive tall, hollow crest on the head.^[1] The genus includes two recognized species: H. altispinus, described by Barnum Brown in 1913 based on postcranial remains from the Horseshoe Canyon Formation in Alberta, Canada, and H. stebingeri, named by John R. Horner and Philip J. Currie in 1994 from embryonic, neonatal, and adult specimens from the Two Medicine Formation in Montana, USA.^[1]^[2] Both species date to the Campanian and Maastrichtian stages, approximately 75 to 67 million years ago, and inhabited coastal floodplains and forested environments.^[3] Adults of Hypacrosaurus reached lengths of about 9 meters (30 feet) and weights of around 4 metric tons. It was a facultatively bipedal/quadrupedal animal, with longer hind limbs relative to forelimbs and tall neural spines along the back.^[4]^[1] As herbivorous animals, they likely fed on low-lying vegetation using their complex dental batteries, and the crest may have served for vocalization or display.^[5] Fossil evidence, including nests and bonebeds, suggests social behavior with parental care and possible age segregation in H. stebingeri.^[2]^[6]

Discovery and species

Initial discovery and naming

The initial fossils of Hypacrosaurus were discovered in 1910 by paleontologist Barnum Brown during an American Museum of Natural History expedition in the Horseshoe Canyon Formation of Alberta, Canada.^[7] In 1913, Brown formally described and named the type species Hypacrosaurus altispinus, with the genus name deriving from Greek roots meaning "near the highest lizard" (hypo- for "near," akros for "highest," and sauros for "lizard"), in reference to its large size, nearly rivaling the largest known carnivorous dinosaur, Tyrannosaurus, at the time.^[1]^[7] The species epithet altispinus further emphasizes these prominent spines, from Latin alti- ("high") and spinus ("spine").^[1] The description was based on the holotype specimen AMNH 5204, a partial adult postcranial skeleton that includes the last eight dorsal vertebrae, the first two caudal vertebrae, both ilia, the right ischium and pubis, several ribs, and elements of the limbs.^[7] No skull was preserved in this initial find, limiting early assessments to skeletal proportions and vertebral morphology.^[1] This naming took place amid a surge in hadrosaur discoveries during the early 20th century, often termed the "dinosaur rush," when researchers like Brown were rapidly expanding knowledge of duck-billed dinosaurs through fieldwork in western North America.^[7] Hypacrosaurus was initially compared to contemporaries such as Corythosaurus, sharing features like a presumed hollow cranial crest and lambeosaurine affinities, though its taller neural spines set it apart in Brown's analysis.^[7]

Subsequent specimens and species distinction

Following the initial description of Hypacrosaurus altispinus from the Horseshoe Canyon Formation in Alberta, subsequent research in the 1970s led to the referral of fragmentary hadrosaurid material from the Two Medicine Formation in Montana to the genus. Peter Dodson's analysis of ontogenetic changes in lambeosaurine skull morphology demonstrated that previously identified taxa like Procheneosaurus represented growth stages of Hypacrosaurus, allowing for the confident assignment of these Montana specimens based on shared crest development and cranial proportions. In the 1980s, paleontologist Jack Horner identified a major bonebed and nesting site in the Two Medicine Formation near Choteau, Montana, yielding numerous juvenile, hatchling, and embryonic remains of a lambeosaurine hadrosaurid, including eggshells and articulated skeletons. These discoveries formed the basis for naming a second species, H. stebingeri, in 1994 by Horner and Currie. The holotype is MOR 549, a nearly complete adult skull exhibiting a rounded, hollow crest; paratypes include MOR 548, a well-preserved juvenile skull, and additional embryonic specimens preserving early ontogenetic features such as unfused nasals and reduced crests. Additional key specimens have expanded the known hypacrosaurid record across formations. In Alberta, partial adult skeletons from the Horseshoe Canyon Formation, such as TMP 1986.46.1 (a postcranial assemblage including vertebrae and limb elements), provide insights into mature morphology for H. altispinus. In Montana, hatchling individuals and eggs from the Two Medicine Formation nesting sites, including articulated embryos with preserved skin impressions, further document early life stages for H. stebingeri. In 2024, the Natural History Museum in Oslo, Norway, acquired two nearly complete skeletons of H. stebingeri (nicknamed Zelda and Zara), offering detailed insights into adult morphology.^[8]^[9] The distinction between H. altispinus and H. stebingeri relies on geographic separation and subtle morphological differences in the skull and crest, with H. stebingeri featuring a broader, more rounded crest shape, while both share tall neural spines along the back. Material from Baja California was initially referred to Hypacrosaurus in 1981 but was later described as the distinct lambeosaurine genus Magnapaulia laticaudus in 2012.^[10]

Description

Overall size and build

Hypacrosaurus was a large hadrosaurid dinosaur, with adult individuals estimated to reach lengths of 7.6 to 9.1 meters from snout to tail tip.^[11] Hip height for adults is estimated at 2.5 to 3 meters, contributing to their substantial stature among Late Cretaceous ornithischians.^[11] Body mass for mature specimens approached 4 tonnes, reflecting the robust scaling typical of lambeosaurine hadrosaurids.^[12] The overall build of Hypacrosaurus was robust, characterized by strong hindlimbs that supported both bipedal and quadrupedal locomotion, allowing facultative shifts in posture for foraging or evasion.^[13] A powerful tail provided balance during movement, while the forelimbs, though less robust than the hindlimbs, enabled quadrupedal support when needed.^[13] This versatile skeletal architecture aligned with the general hadrosaurid body plan, emphasizing stability and efficiency in a terrestrial herbivorous lifestyle. As a herbivore, Hypacrosaurus inferred its diet from a complex dental battery capable of grinding tough vegetation, a trait shared with related lambeosaurines like Corythosaurus.^[14] Ontogenetic changes were pronounced, with hatchlings measuring approximately 1.7 meters in length and rapidly scaling to adult dimensions through accelerated growth phases.^[2]

Skull and crest morphology

The skull of Hypacrosaurus is elongated and robust, featuring a broad, ventrally deflected rostrum that facilitated low-level browsing on vegetation.^[9] This structure includes a complex dental battery comprising hundreds of teeth arranged in functional rows, with secondary ridges present in juveniles but resorbed in adults, enabling efficient processing of tough plant material.^[9] A defining feature of Hypacrosaurus is its hollow cranial crest, formed primarily by the premaxillae and nasals, with the nasals contributing the majority of the ventral tubes and the premaxilla providing an anterior cover via a lateral process.^[9] The nasal passages route through the crest, featuring a hypertrophied, non-olfactory vestibule external to the main cavities.^[9] In H. stebingeri, the crest exhibits ontogenetic variation: juveniles and subadults display a taller, more rectangular profile with an incipient structure dominated by the nasal and elongate narial openings, transitioning to a more rounded, circular shape in adults.^[15]^[9] In contrast, H. altispinus possesses a shorter, broader crest with an anteriorly projecting nasal process overlying the dorsal premaxilla, resulting in a semicircular profile that varies from as tall as long to longer than tall across adult specimens.^[9] Crest development begins around 50% of adult skull length, with juveniles lacking a fully formed structure; growth proceeds isometrically at first, followed by positive allometry, achieving full development by the subadult stage, as evidenced by a decreasing crest-snout angle from approximately 163° in juveniles to 141°–159° in adults.^[9]

Postcranial skeleton

The postcranial skeleton of Hypacrosaurus features prominently elongated neural spines on the dorsal vertebrae, measuring 5 to 7 times the height of the vertebral centra and contributing to a distinctive high-backed profile.^[7] These spines are particularly tall in H. altispinus, the type species named for this trait (altispinus meaning "high-spined"), exceeding those in H. stebingeri and elevating the animal's overall body height substantially in adults.^[1] The holotype of H. altispinus (AMNH 5204) preserves the last eight dorsal vertebrae with incomplete neural spines, while referred specimens confirm their elongation in the shoulder region for both species.^[16] The appendicular skeleton reflects adaptations for both bipedal and facultative quadrupedal locomotion. Forelimbs are robust, with the humerus often shorter than or equal to the scapula in juveniles but becoming substantially longer in adults to support weight-bearing postures; the manual digits exhibit hoof-like phalanges suited for quadrupedality.^[17] Hindlimbs are powerful, featuring a femur that is shorter than or equal to the tibia in young individuals but longer in adults, enabling efficient bipedal speed and propulsion.^[17] The pelvic girdle is broad and robust, with a well-developed ilium for weight support and a distinctive autapomorphy in H. altispinus: an enlarged ischial foot where the depth exceeds 26% of the shaft length.^[16] Caudal vertebrae and associated elements indicate a stiff tail for balance during locomotion. The proximal caudals in the H. altispinus holotype show elongation, paired with extended haemal arches (chevrons) that enhance tail rigidity.^[18] Ontogenetic changes are evident throughout the postcrania, with juveniles displaying proportionally longer limbs relative to the body (e.g., more gracile femur-tibia ratios and shorter neural spines, less than four times centra height) compared to the more robust, proportionately shorter-limbed adults.^[17] These differences underscore rapid growth and shifts toward greater structural support in maturity.^[17]

Classification

Phylogenetic relationships

Hypacrosaurus is classified within the subfamily Lambeosaurinae of the family Hadrosauridae, a group of ornithopod dinosaurs characterized by synapomorphies such as a hollow nasal crest formed by the premaxillae, nasals, and prefrontals, and elongated neural spines on the vertebrae that contribute to a high-backed profile.^[9] These features distinguish lambeosaurines from the hollow-crested Parasaurolophini and the non-crested Saurolophinae.^[19] Phylogenetic analyses consistently place Hypacrosaurus within Lambeosaurinae, often as a member of the tribe Lambeosaurini. A 2013 study on lambeosaurine systematics placed Hypacrosaurus within Lambeosaurini, more closely related to Corythosaurus than to Lambeosaurus.^[9] This positioning highlights its North American affinities during the Late Cretaceous.^[19] More recent analyses have refined this placement. A 2022 phylogenetic analysis incorporating detailed cranial and postcranial data positioned Hypacrosaurus as a close relative to Olorotitan within Lambeosaurinae, emphasizing differences in crest morphology and neural spine proportions.^[20] Ongoing debates surround its relationship to Nipponosaurus sachalinensis, with earlier studies suggesting synonymy or close affinity to H. altispinus based on juvenile similarities, while a 2017 reanalysis using expanded character matrices placed Nipponosaurus as a more basal lambeosaurine, rejecting synonymy due to distinct postcranial proportions.^[21]

Species validity and synonymy

Hypacrosaurus altispinus, the type species, was established based on distinctive vertebral proportions in its holotype specimen (AMNH 5204), particularly the exceptionally tall neural spines on the dorsal vertebrae, which exhibit a height-to-centrum ratio exceeding 3:1, setting it apart from other lambeosaurines. This postcranial feature, described from partial skeletal remains including vertebrae and a pelvis recovered from the Horseshoe Canyon Formation, supports its validity as a unique species within the genus. In contrast, Hypacrosaurus stebingeri was defined primarily through embryonic and neonatal specimens, including well-preserved skulls that reveal a distinctive low, rounded cranial crest formed mainly by the premaxillae and nasals, along with associated eggs and hatchling postcrania from the Two Medicine Formation and Judith River Formation. These morphological traits, including elongate narial openings and embryonic bone histology, distinguish it from H. altispinus and confirm its status as a valid species. Ongoing taxonomic debates highlight potential overlap between H. altispinus and H. stebingeri due to their temporal proximity across the Campanian-Maastrichtian boundary in western North American formations with similar depositional environments, raising questions about whether observed differences reflect true species distinction or ontogenetic variation.^[9] However, no formal synonymy has been proposed, as the two retain distinct autapomorphies in vertebral and cranial morphology, respectively.^[9] Additionally, a 2022 phylogenetic analysis suggested that the Baja California lambeosaurine Magnapaulia laticaudus may nest within Hypacrosaurus, potentially representing an indeterminate species (Hypacrosaurus sp.) based on shared traits such as broad caudal vertebrae and lambeosaurine crest architecture, though this remains an ongoing discussion without full synonymization.^[20]

Paleobiology

Growth and ontogeny

Hypacrosaurus exhibited rapid juvenile growth, with hatchlings measuring approximately 1 to 1.7 meters in length and reaching subadult sizes of 5 to 6 meters within 2 to 3 years after hatching.^[22] This accelerated phase is evidenced by a growth inflection point in long bones, such as the femur at around 2.5 years and the tibia at 2.8 years, marking the transition to slower growth rates.^[22] Full somatic maturity was attained by 10 to 12 years, when individuals approached 95% of their asymptotic body length of about 7 to 9 meters.^[22] Bone histology of Hypacrosaurus reveals fibrolamellar bone tissue in the cortex of long bones, indicative of fast growth rates typical of large dinosaurs. Lines of arrested growth (LAGs) are present but vary in number across skeletal elements, ranging from 0 in some phalanges to up to 8 in the tibia and femur, complicating precise age estimation due to Haversian remodeling that obscures earlier LAGs. The presence of an external fundamental system (EFS) in adult specimens signals the cessation of rapid periosteal deposition and the onset of slower, zonal bone formation. Ontogenetic changes in Hypacrosaurus included post-hatching elongation of the neural spines and the development of the hollow nasal crest, which began as a small prominence in juveniles and expanded significantly with age, potentially altering respiratory or display functions.^[23] Locomotor shifts occurred during growth, with juveniles primarily bipedal and larger subadults and adults adopting a facultatively quadrupedal stance, as inferred from proportional changes in limb bone scaling and robust forelimb development. A 2024 taphonomic analysis of Hypacrosaurus stebingeri bonebeds demonstrates age segregation, with early juveniles under 1 year old and late juveniles around 3 years old forming separate cohorts, suggesting gregarious behavior in monospecific herds that persisted until individuals reached their fourth year before potentially integrating into multigenerational groups.^[24] This pattern implies distinct social structures during ontogeny, with bonebed assemblages reflecting mass mortality events in age-specific herds.^[24]

Reproduction and nesting

Hypacrosaurus stebingeri produced elongated eggs measuring approximately 20 by 18.5 cm, with an estimated volume of 3,900 cm³, arranged in clutches typically containing 15 to 20 eggs.^[25] These eggs were laid in colonial nesting sites, where multiple clutches were positioned in close proximity to facilitate group protection.^[26] Nest sites for Hypacrosaurus have been identified in the Upper Cretaceous Two Medicine Formation of Montana, particularly at localities such as Blacktail Creek (TM-066), where mound-style nests were constructed by scraping shallow depressions in the soil and possibly covering them with vegetation for insulation and moisture retention.^[26] Exceptional preservation at these sites includes embryo specimens like MOR 548, which exhibit the characteristic curled posture of late-stage dinosaur embryos within the egg, indicating advanced development prior to hatching.^[26] Histological analysis of embryonic teeth from similar specimens reveals incremental growth lines (von Ebner lines), supporting an incubation period of approximately 171 days, consistent with reptilian-grade metabolic rates rather than the accelerated avian pattern.^[25] Evidence for parental care in Hypacrosaurus is inferred from the dense clustering of nests at these sites, suggesting colonial breeding that allowed adults to guard multiple clutches collectively against predators.^[26] Hatchlings, emerging at around 1 to 1.7 meters in length, were highly vulnerable to predation due to their small size and limited mobility, implying that proximity to nesting adults provided essential protection during the early post-hatching phase.^[25]^[22] No definitive evidence of sexual dimorphism exists in Hypacrosaurus, though the prominent hollow crest may have exhibited size variations potentially linked to sexual display functions, as hypothesized for other lambeosaurines based on growth patterns in cranial ornamentation.^[23]

Crest functions and sensory adaptations

The hollow nasal crest of Hypacrosaurus, a lambeosaurine hadrosaurid, primarily served acoustic functions by resonating low-frequency sounds produced in the throat, facilitating long-distance communication within herds.^[27] The convoluted nasal passages within the crest amplified frequencies below 1,000 Hz, with juveniles capable of higher pitches around 1.1 kHz due to smaller cavity sizes, enabling species-specific calls for coordination during migration or predator avoidance.^[28] This resonance model is supported by the elongate cochlea in Hypacrosaurus skulls, indicating sensitivity to low frequencies around 80 Hz.^[28] In addition to acoustics, the crest functioned as a visual signal for social and reproductive purposes, with its helmet-like shape and ontogenetic changes from small juvenile structures to prominent adult forms aiding in mate attraction and species recognition.^[29] High variation in crest morphology among lambeosaurines, including Hypacrosaurus, suggests it acted as a premating isolating mechanism, similar to display structures in modern birds like cassowaries, where exaggerated head features enhance intra-specific signaling without impeding basic sensory functions.^[30] The large cerebral hemispheres in Hypacrosaurus endocasts (comprising about 43% of brain volume) further imply complex social behaviors supported by such visual cues.^[28] Debates on alternative functions, such as humidity control or enhanced olfaction, have been largely resolved through CT scans revealing that the olfactory epithelium remained positioned rostromedial to the orbits, outside the crest cavities, with no evidence of respiratory turbinates for moisture retention.^[30] These scans confirm the crest's hypertrophy was tied to behavioral adaptations like vocalization and display rather than physiological needs, aligning with paleoneurological data showing small olfactory bulbs (less than 5% of endocast volume).^[28] Acoustic and visual roles thus appear complementary, promoting herd cohesion in the Late Cretaceous environments of western North America.^[23]

Thermoregulation and metabolism

Analyses of oxygen isotope ratios in the tooth enamel and bone phosphate of juvenile Hypacrosaurus specimens reveal low intrabone and interbone variability, indicating stable body temperatures with fluctuations of less than 2°C during growth, a pattern consistent with endothermic thermoregulation rather than the high variability expected in ectotherms.^[31] This low heterogeneity suggests that Hypacrosaurus maintained internal heat production to regulate its physiology, distinguishing it from modern reptiles that exhibit greater isotopic variation tied to environmental fluctuations.^[31] Further oxygen isotope data from Hypacrosaurus stebingeri enamel yield δ¹⁸O values around 10.5‰, corresponding to estimated body temperatures of 36–38°C, substantially higher than the inferred ambient environmental temperatures of approximately 15–20°C in its high-latitude habitat during the Late Cretaceous.^[32] These values align with broader hadrosaurid patterns, where clumped isotope analyses of related taxa confirm average body temperatures near 37°C, supporting widespread endothermy among ornithischians and refuting earlier inferences of ectothermy.^[33]^[34] In contrast, a 1996 study based on nasal passage morphology in Hypacrosaurus proposed metabolic rates closer to those of extant reptiles, but subsequent isotopic evidence has favored partial to full endothermy.^[35] Histological examination of Hypacrosaurus long bones demonstrates rapid growth rates, with annual increments indicating body mass increases of up to 100% per year in subadults, far exceeding those of modern ectothermic reptiles and implying elevated metabolic demands for sustained high-energy tissue deposition.^[36] Elongated neural spines along the vertebral column may have functioned as heat exchangers to aid thermoregulation, though this hypothesis remains debated in light of the isotopic data supporting overall endothermic physiology.^[36] Embryonic growth lines in Hypacrosaurus eggshells further suggest a developmental shift toward higher metabolic rates mid-incubation, bridging reptilian-like early stages with later endothermic traits observed in juveniles and adults.^[37]

Soft tissue preservation

Exceptional soft tissue preservation in Hypacrosaurus stebingeri has been documented in embryonic specimens, particularly the MOR 548 embryo from the Two Medicine Formation.^[38] In a 2020 study, researchers isolated chondrocytes from the calcified cartilage of this specimen, revealing round cells with microstructures consistent with preserved nuclei containing condensed, chromosome-like material, potentially representing chromatin remnants from cellular processes such as chondroptosis.^[38] Immunohistochemical analyses further identified preserved collagen type II epitopes in the extracellular matrix, appearing as globular structures with fluorescence patterns analogous to those in extant avian cartilage, though with reduced intensity.^[38] Chemical markers suggestive of DNA preservation were detected through histochemical staining; isolated H. stebingeri chondrocytes exhibited positive reactions to propidium iodide (PI) and 4',6-diamidino-2-phenylindole (DAPI), dyes that intercalate with double-stranded DNA, localizing specifically to nuclear regions in a manner comparable to modern cells.^[38] However, these findings have sparked debate regarding whether the signals indicate endogenous biomolecules or artifacts from microbial contamination or non-specific binding, with critics noting the extraordinary age of the material—approximately 75 million years—and the absence of confirmatory sequencing data.^[38] The study authors countered contamination hypotheses by emphasizing the specificity of staining patterns, lack of bacterial biofilms, and controls using extant tissues, but no independent replication of these molecular results has been reported since 2020.^[38] These observations imply that biomolecules in calcified cartilage can persist far longer than previously modeled, potentially up to tens of millions of years under rare taphonomic conditions, challenging conventional degradation timelines for proteins and nucleic acids.^[38] Despite this, no complete genome or extensive sequence recovery has been achieved, limiting applications to paleogenomics, and the controversy underscores the need for advanced analytical techniques to verify such ancient molecular traces.^[38]

Paleopathology and injuries

Fossil evidence of predation on Hypacrosaurus includes tooth score marks on limb bones, such as a deep groove on the fibula of specimen MOR 549, attributed to the serrated teeth of a tyrannosaurid predator. In the Late Cretaceous ecosystems of western North America, Daspletosaurus served as a primary predator of Hypacrosaurus, with bite marks consistent with the dimensions of its dentition supporting scavenging or failed predatory attempts on subadult individuals. A 2025 study by Bertozzo et al. documented recurrent tail pathologies in Hypacrosaurus stebingeri, particularly in the proximo-middle caudal vertebrae, with specimen MOR 549 exhibiting multiple healed fractures and deformations in the neural spines.^[39] These injuries, affecting approximately 15.4% of examined caudal vertebrae across hadrosaurid specimens including Hypacrosaurus, show signs of direct trauma from diagonal compressive forces (30°–60° angle), most plausibly linked to mating behaviors involving side-mounting by conspecifics, as finite element analysis ruled out predation or trampling as primary causes.^[39] The concentration of lesions in adult individuals, with advanced healing stages at death, suggests repeated occurrences potentially indicating sexual dimorphism, where females bore the brunt of such trauma near the cloacal region.^[39] Additional traumatic injuries in Hypacrosaurus fossils include healed rib fractures, as seen in the hatchling specimen MOR 548 from Montana, where a small callus on the rib shaft indicates survival following thoracic trauma likely from a fall or conspecific interaction. Vertebral pathologies in Hypacrosaurus also encompass possible infections, such as osteomyelitis in caudal elements, evidenced by irregular bone overgrowth and periosteal reactions in specimens like MOR 549, though distinguishing infectious from purely traumatic origins remains challenging without histological data.^[39]

Paleoecology

Geological formations and taphonomy

Fossils of Hypacrosaurus altispinus are primarily recovered from the Horseshoe Canyon Formation in Alberta, Canada, a unit of the Edmonton Group deposited during the early Maastrichtian stage of the Late Cretaceous, approximately 71 to 68 million years ago. This formation consists of interbedded sandstones, siltstones, and mudstones indicative of fluvial and coastal plain environments. In contrast, H. stebingeri is known from the Two Medicine Formation in northwestern Montana, United States, and the equivalent Oldman Formation in Alberta, Canada, which spans the Campanian, from about 82 to 74 million years ago, encompassing a range of terrestrial depositional settings including fluvial channels, floodplains, and volcaniclastic layers.^[16]^[40] The biostratigraphic context of Hypacrosaurus fossils aligns with the Judithian land-vertebrate "age" of the late Campanian, particularly for specimens from the upper Two Medicine Formation, where index taxa and magnetostratigraphy support correlation to this interval. Remains from the Horseshoe Canyon Formation fall into the succeeding Edmontonian "age," reflecting a temporal progression across the Campanian-Maastrichtian boundary. These assignments are based on dinosaur faunal assemblages and radiometric dating of volcanic tuffs within the formations.^[41]^[42] Taphonomic processes for Hypacrosaurus vary by ontogenetic stage and depositional setting. Nesting sites, particularly those preserving embryonic and hatchling remains of H. stebingeri, were rapidly entombed by volcanic ashfalls in the Two Medicine Formation, leading to exceptional preservation of fragile structures such as eggshells and skeletal elements with minimal post-mortem alteration. Adult and subadult specimens, however, are typically preserved in fluvial deposits of both formations, where seasonal flooding or drought-induced mass mortalities contributed to bonebed formation, often involving groups of similarly aged individuals. These bonebeds exhibit uniform taphonomic signatures, including low levels of weathering, abrasion, and trampling, suggesting rapid burial in low-energy channel or overbank environments.^[6] Fully articulated skeletons of Hypacrosaurus are rare, with most assemblages showing partial to complete disarticulation due to hydraulic transport in riverine systems, scattering bones across floodplain deposits. Notable exceptions include a partially articulated subadult skull and associated postcrania from the Horseshoe Canyon Formation, preserved in finer-grained sediments that limited dispersal. Bonebeds from mass death events dominate the record, providing insights into gregarious behavior but often with fragmented elements due to subsequent sediment reworking.^[16]^[6]

Paleoenvironment and habitat

Hypacrosaurus inhabited the diverse landscapes of the Late Cretaceous Western Interior of North America, characterized by coastal floodplains, meandering rivers, dense forests, and extensive wetlands. These environments formed part of the alluvial plains bordering the Western Interior Seaway, with sediment deposition dominated by fluvial channel-belt and overbank deposits in the Horseshoe Canyon Formation of Alberta, where H. altispinus is found.^[43] In contrast, the Two Medicine Formation of Montana, home to H. stebingeri, preserved fluvial and floodplain facies interbedded with lacustrine deposits, reflecting a more inland setting influenced by sediment input from nearby highlands.^[40] The regional climate was seasonal, featuring wet-dry cycles driven by monsoonal patterns and rain shadows from the rising Cordilleran mountains, with evidence of periodic droughts interrupting growth in local vegetation.^[44] The vegetation in these habitats supported a rich herbivorous niche for Hypacrosaurus, consisting primarily of conifers such as taxodiaceous trees, ferns, and emerging angiosperms including magnoliids and early eudicots, which formed multilayered forests and understories along riverbanks and floodplains.^[45] As facultative bipeds, Hypacrosaurus individuals could browse at heights of 1-4 meters, accessing mid-level foliage on soft, fibrous plants and fruits while avoiding competition with lower feeders like ceratopsians; quadrupedal stance allowed grazing on ground-level ferns and horsetails during wetter periods.^[46] Wetlands and riparian zones provided additional resources, with seasonal flooding promoting lush growth that sustained large hadrosaur populations. Habitat partitioning between the two species reflected topographic and climatic gradients across the region. H. stebingeri occupied montane foothills in a volcanically active setting, where bentonite ash layers from Cordilleran eruptions enriched floodplain soils and influenced local hydrology in the Two Medicine Formation.^[47] Conversely, H. altispinus thrived in lowland coastal plains closer to the seaway, with river systems and wetlands in the Horseshoe Canyon Formation experiencing tidal influences and higher sedimentation rates.^[48] Paleoclimate reconstructions indicate a warm-temperate regime, with mean annual temperatures around 14°C, warm months averaging 22°C, and cold months near 8°C, supporting year-round activity amid the seasonal precipitation patterns.^[49]

Contemporaneous fauna and interactions

Hypacrosaurus coexisted with diverse vertebrate assemblages across formations such as the Oldman, Two Medicine, and Horseshoe Canyon formations during the Campanian and Maastrichtian stages.^[50] The dominant herbivores included other hadrosaurids such as Maiasaura peeblesorum and Prosaurolophus maximus, alongside ceratopsians like Centrosaurus apertus and Chasmosaurus russelli, which together formed the bulk of the megaherbivorous community.^[50] Ankylosaurians, including species referable to Euoplocephalus, were also present, contributing to the low-level browsing guild.^[50] Apex predators targeting Hypacrosaurus included tyrannosaurids such as Daspletosaurus torosus and Gorgosaurus libratus, which preyed on both juveniles and adults based on comparative growth trajectories and fossil evidence of predation.^[51] Smaller theropods like Troodon formosus and dromaeosaurids (e.g., Saurornitholestes langstoni) likely acted as opportunistic predators or scavengers, particularly on young or vulnerable individuals.^[51]^[50] Ecological interactions among these taxa involved niche overlap in mid- to high-level browsing, where Hypacrosaurus shared foraging resources with other hadrosaurids and ceratopsians on tough vegetation up to 4 meters in height, potentially leading to resource competition.^[52] Herd structures in hadrosaurids, including Hypacrosaurus, may have facilitated collective defense against tyrannosaurid attacks, reducing individual predation risk through group vigilance and evasion behaviors inferred from assemblage patterns.^[51] In the regional food web, Hypacrosaurus occupied a mid-trophic level as a primary herbivore, serving as key prey for large theropods while influencing vegetation dynamics through its grazing habits.^[52]^[51] This position highlights its role in sustaining apex predators, with evidence of bite marks on hadrosaur bones indicating direct trophic links to Daspletosaurus and similar taxa.^[51]

Behavioral inferences from bonebeds

Fossil bonebeds of Hypacrosaurus stebingeri provide key evidence for gregarious behavior, with multiple individuals preserved in monodominant assemblages indicating social grouping for protection and foraging. For instance, the Blacktail Creek North bonebed in the Two Medicine Formation of Montana preserves the remains of at least 23 early juvenile individuals, suggesting they lived and possibly died together in cohorts.^[6] Similarly, the Devil's Coulee Juvenile Hadrosaur Bonebed in the Oldman Formation of Alberta contains at least four late juvenile specimens, while the Lambeosite bonebed in the Two Medicine Formation includes four late juveniles and one adult, further supporting herd-like structures among age-specific groups.^[6] A 2024 taphonomic analysis by Joubarne, Therrien, and Zelenitsky compared these bonebeds and revealed age segregation in H. stebingeri social behavior, where juveniles formed separate cohorts from adults until approximately their fourth year of life, likely rejoining multigenerational herds upon reaching sexual maturity.^[24] This segregation, observed across sites in Montana and Alberta, implies complex social dynamics, with early juveniles (<1 year old) in larger nursery groups for protection, transitioning to smaller late-juvenile cohorts (~3 years old) possibly optimized for independent foraging, before integrating into mixed-age herds.^[24] Such patterns mirror behaviors in modern ungulates and enhance understanding of hadrosaurid social complexity, as the decreasing group sizes in later juvenile bonebeds may reflect higher mortality or shifting ecological needs.^[24] The distribution of Hypacrosaurus bonebeds across contemporaneous formations in Alberta and Montana hints at possible seasonal movements to exploit varying resources, though direct taphonomic evidence remains limited.^[31] Oxygen isotope analyses of juvenile specimens indicate stable body temperatures consistent with endothermy, supporting the capacity for long-distance travel if environmental pressures necessitated it.^[31]

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Testing size–frequency distributions as a method of ontogenetic ...
Aug 1, 2020 · ... Hypacrosaurus MOR 549. To test this large discrepancy, we ... 2015; Wosik and Evans 2015; Wosik et al. 2017a). By 4 years of age ...
[11]
https://pubs.geoscienceworld.org/paleosoc/paleobiol/article/46/3/379/590865/Testing-size-frequency-distributions-as-a-method
[12]
Limb-Bone Scaling Indicates Diverse Stance and Gait in ... - NIH
May 22, 2012 · Our findings suggest that hadrosaurs had a higher locomotor performance than other quadrupedal ornithischians. This could relate to an ability ...
[13]
Ontogeny reveals function and evolution of the hadrosaurid ...
Jul 28, 2016 · Hadrosaurid dinosaurs, dominant Late Cretaceous herbivores, possessed complex dental batteries with up to 300 teeth in each jaw ramus.
[14]
A sub-adult skull of Hypacrosaurus stebingeri (Ornithischia
Juveniles and sub-adults of H. stebingeri are characterised by an incipient cranial crest formed predominantly by the nasal, elongate narial openings with a ...
[15]
Hypacrosaurus altispinus Brown 1913 - Plazi TreatmentBank
Nov 6, 2023 · Hypacrosaurus altispinus has one autapomorphy in the postcranial skeleton, a relatively enlarged ischial foot in which the depth of the foot is over 26% the ...<|control11|><|separator|>
[16]
Placing juvenile specimens in phylogenies: An ontogenetically ...
Oct 31, 2022 · ... Hypacrosaurus, the juvenile scapulae are relatively long, with a ratio of 7.7 in MOR 548 and MOR 355, and 7.2 in the adult MOR 549.<|control11|><|separator|>
[17]
Hypacrosaurus altispinus? Brown from the Two Medicine Formation ...
7948 and type of Hypacrosaurus altispinus (AMNH 5204) and paratype (AMNH 5272) (From Gilmore, 1917). H. altispinus. AMNH. 5204 USNM. Type 7948. Ilium:.Missing: holotype | Show results with:holotype<|separator|>
[18]
Diversity, Relationships, and Biogeography of the Lambeosaurine ...
Jul 26, 2013 · We provide a thorough re-evaluation of the taxonomic diversity, phylogenetic relationships, and historical biogeography of the lambeosaurine ...
[19]
Reanalysis of the phylogenetic status of Nipponosaurus ...
Our phylogenetic analysis reveals that Nipponosaurus is a basal lambeosaurine hadrosaurid, much further down in the tree than previously suggested, and shows a ...
[20]
https://www.tandfonline.com/doi/full/10.1080/14772019.2022.2043924
[21]
Cranial anatomy and systematics of Hypacrosaurus altispinus, and a ...
Two new horned dinosaurs from the Upper Cretaceous Two Medicine Formation of Montana; with a phylogenetic analysis of the Centrosaurinae (Ornithischia ...<|control11|><|separator|>
[22]
https://doi.org/10.1098/rspb.2008.0912
[23]
Dinosaur incubation periods directly determined from growth-line ...
Jan 3, 2017 · We discovered daily forming growth lines in teeth of embryonic nonavian dinosaurs revealing incubation times. These lines show slow reptilian-grade development ...Dinosaur Incubation Periods... · Results · Materials And Methods
[24]
First Evidence of Dinosaurian Secondary Cartilage in the Post ...
First Evidence of Dinosaurian Secondary Cartilage in the Post-Hatching Skull of Hypacrosaurus stebingeri (Dinosauria, Ornithischia) ... Horner JR,Currie PJ (1994) ...
[25]
Oxygen isotope variability in juvenile dinosaurs (Hypacrosaurus)
Feb 8, 2016 · Analyses of juvenile Hypacrosaurus individuals indicate a pattern of low isotopic heterogeneity suggestive of endothermy. Information. Type ...
[26]
Oxygen isotopes from biogenic apatites suggest widespread ...
Oxygen isotopes from biogenic apatites suggest widespread endothermy in Cretaceous dinosaurs ... Amiot et al. [20] for details). The locality of Cherves-de-Cognac ...Missing: hadrosaur | Show results with:hadrosaur
[27]
Dinosaur Body Temperatures Determined from Isotopic ( 13 C
Our data indicate body temperatures of 36° to 38°C, which are similar to those of most modern mammals. This temperature range is 4° to 7°C lower than predicted ...
[28]
Eggshell geochemistry reveals ancestral metabolic ... - Science
Feb 14, 2020 · Variations in assumed model parameters can produce large deviations in temperature estimates (2° to 8°C), even in extant taxa (7). This ...
[29]
The Metabolic Status of Some Late Cretaceous Dinosaurs - Science
... (Hypacrosaurus) showed that their metabolic rates were significantly lower than metabolic rates in modern birds and mammals. In extant endotherms and ...
[30]
Relative growth rates of predator and prey dinosaurs reflect effects ...
Aug 5, 2008 · In this study, we longitudinally sampled a sequence of lines of arrested growth (LAGs) from an essentially full-grown hadrosaur Hypacrosaurus ...
[31]
Embryonic metabolism of the ornithischian dinosaurs Protoceratops ...
Apr 24, 2017 · The embryonic metabolisms of the ornithischian dinosaurs Protoceratops andrewsi and Hypacrosaurus stebingeri have been determined and are in ...Missing: crest morphology validity
[32]
https://www.sciencedirect.com/science/article/abs/pii/S0012821X06003050
[33]
https://www.science.org/doi/10.1126/science.1206196
[34]
Updating the Upper Cretaceous (Campanian) Two Medicine ...
Jul 17, 2024 · New age data confirm that the Two Medicine Formation accumulated during much of the Campanian, with deposition spanning ca. 82.4 Ma to 74.4 Ma.
[35]
Dinosaur biostratigraphy of the Edmonton Group (Upper Cretaceous ...
Mar 4, 2013 · Although a small bonebed containing Hypacrosaurus altispinus is known from near the base of the Tolman Member, specimens occur mostly as single ...
[36]
Untitled Document - Paleofile.com
Genus: Hypacrosaurus BROWN, 1913. Etymology: Greek, bypakros, “nearly the highest”, and Greek, sauros, “lizard”: “near-highest lizard”. = Cheneosaurus LAMBE, ...Missing: naming Lawrence
[37]
Insights from the Dinosaur Park Formation, Alberta, Canada
May 8, 2025 · A floodplain-dominated stratigraphic section of the Dinosaur Park Formation comprises six recurring lithofacies and two facies associations ...Missing: forests | Show results with:forests
[38]
Two Medicine Formation | Life on Our Planet Wiki - Fandom
The site is famous for its fossil eggs of the dinosaur, which are arranged in dug out nests of earth. This showed that dinosaurs did in fact care for their ...
[39]
https://doi.org/10.1016/j.isci.2025.113739
[40]
Feeding height stratification among the herbivorous dinosaurs from ...
Apr 4, 2013 · Hadrosaurids were uniquely capable of feeding up to 2 m quadrupedally, or up to 5 m bipedally. There is no evidence for either feeding height ...
[41]
[PDF] Facies analysis of the volcaniclastic Two Medicine Formation Wolf ...
Montana suggest an volcaniclastic apron depositional environment ... fluvial and flood- plain deposits to lacustrine ... The volcanic member of the Two Medicine ...
[42]
Paleoenvironmental reconstruction of Late Cretaceous rivers ...
Aug 7, 2025 · A floodplain-dominated succession in the northwestern sector of Dinosaur Provincial Park was examined in an interval where channel-belt ...Missing: forests | Show results with:forests
[43]
New Absolute Paleoclimate Estimates from Dinosaur Provincial Park ...
May 23, 2018 · New absolute paleoclimate estimates from dinosaur Provincial Park, Alberta (Campanian, Late Cretaceous): Preliminary results based on a new fossil leaf ...
[44]
Palaeoenvironmental drivers of vertebrate community composition ...
Nov 15, 2016 · The similarity of the dinosaur faunal assemblages between the time-equivalent portions of the Dinosaur Park Formation and Oldman Formation ...
[45]
https://www.britannica.com/science/Cretaceous-Period/Terrestrial-life
[46]
https://bmcecol.biomedcentral.com/articles/10.1186/1472-6785-13-14