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Einiosaurus

Einiosaurus is a genus of herbivorous centrosaurine ceratopsian dinosaur that lived during the Late Cretaceous period, approximately 74 million years ago, in what is now northwestern Montana, United States.^[1] Known from at least 15 individuals recovered from two monospecific bonebeds in the Two Medicine Formation, it represents a medium-sized ceratopsid, estimated to be about 6 meters (20 feet) in length.^[1]^[2] The genus is distinguished by its unique cranial ornamentation, including a prominent nasal horn that curves forward and downward in adults, paired supraorbital horns, and a short parietosquamosal frill adorned with elongate epiparietals forming straight spikes along the posterior margin.^[3]^[4] Named in 1995 by paleontologist Scott D. Sampson based on specimens discovered by Jack Horner and teams from the Museum of the Rockies between 1985 and 1989, Einiosaurus procurvicornis (meaning "buffalo lizard with a forward-curving horn") is the type and only species.^[3]^[2] The holotype skull (MOR 456) and associated postcranial remains indicate a robust build adapted for browsing low vegetation, with a parrot-like beak and dental battery for processing tough plant matter typical of ceratopsians.^[3] The bonebeds suggest gregarious behavior, with evidence of mass mortality events possibly linked to environmental stresses like drought, and histological studies indicate rapid juvenile growth slowing after 3–5 years, possibly around the onset of sexual maturity.^[5]^[2] As a transitional form within centrosaurines, Einiosaurus exhibits morphologies bridging earlier ceratopsids and more derived taxa like Centrosaurus and Styracosaurus, providing insights into the ontogenetic changes and evolutionary diversity of horned dinosaurs in the northern Western Interior of North America.^[3]^[4] All known fossils are housed at the Museum of the Rockies in Bozeman, Montana, underscoring the site's importance for ceratopsian research.^[2]

Discovery and research history

Expeditions and fossil collection

The discovery of Einiosaurus fossils occurred during field expeditions led by paleontologist Jack Horner in the Landslide Butte area of the Two Medicine Formation, northwestern Montana, near the United States-Canada border approximately 40 km northwest of Cut Bank.^[6] These expeditions, conducted by teams from the Museum of the Rockies, began in the early 1980s and intensified in 1985 when Horner and his crew identified two monospecific bonebeds (designated TM-023 and TM-046) containing abundant ceratopsian remains initially thought to represent a new species of Styracosaurus.^[2] Excavation efforts continued annually through 1989, yielding hundreds of bones including three partial adult skulls, with all specimens housed at the Museum of the Rockies under inventory numbers such as MOR 456.^[6] The primary bonebed site, known as the Canyon Bonebed (TM-046), represents a thanatocoenosis—a death assemblage—of at least 15 individuals, predominantly juveniles and subadults based on bone histology and size variation, preserved in a low-diversity floodplain deposit within Lithofacies 5 of the formation, about 45 meters below the Two Medicine-Bearpaw contact.^[6] Taphonomic analysis indicates the assemblage formed due to drought-related mortality, with bones showing minimal weathering, limited trampling, and articulation in some crania, suggesting rapid burial in overbank sediments during seasonal flooding. A second nearby bonebed (TM-023) contributed additional disarticulated material, up to 40 bones per square meter in dense concentrations, further supporting gregarious herd behavior among the animals.^[6] Fieldwork involved systematic quarry mapping to document spatial distribution and orientation of elements, followed by careful jacketing and removal of blocks for transport to the Museum of the Rockies, where initial mechanical preparation occurred using air scribes and consolidants to stabilize fragile frill and horn fragments.^[2] These methods allowed for the recovery of associated postcranial elements like ribs and vertebrae alongside cranial material, preserving contextual taphonomic data. Challenges included the site's remote badlands location along the Milk River, which complicated logistics and access, particularly during wet seasons that could flood the area or erode exposures.^[6] Stratigraphic correlation of the bonebeds to equivalent units like the Judith River Formation in central Montana relied on lithologic matching and biostratigraphic markers, such as the presence of shared fauna, to confirm their late Campanian age around 76-74 million years ago.^[7]

Naming and initial description

The fossils now assigned to Einiosaurus were initially collected during field expeditions led by Jack Horner in the late 1980s from the Two Medicine Formation in Montana.^[5] In 1995, paleontologist Scott D. Sampson formally named the genus and species Einiosaurus procurvicornis, designating the holotype as MOR 456 8-9-6-1, a partial adult skull preserving the nasal horncore, supraorbital horns, and portions of the frill.^[3] Sampson highlighted the diagnostic low, forward-procumbent nasal horn as a key autapomorphy distinguishing it from other ceratopsids, noting its unique curvature that projected anteriorly and slightly ventrally.^[3] The generic name Einiosaurus derives from the Blackfeet word eini (meaning "buffalo") combined with the Greek sauros (lizard), reflecting the animal's bison-like profile due to its robust build and curving horn; the specific epithet procurvicornis refers to the forward-curving horn, from Latin pro (forward), curvus (curved), and cornis (horn).^[3] Prior to formal naming, Horner and colleagues in the late 1980s and early 1990s referred to the material as "Transitional Taxon B" in preliminary reports, interpreting it as an intermediate form between earlier centrosaurines like Centrosaurus and later ones such as Styracosaurus, based on shared frill spikes but differing nasal and postorbital horn shapes.^[5] Sampson's description expanded on these observations, confirming Einiosaurus as a centrosaurine ceratopsid through comparisons of skull morphology, including the elongate, triangular parietal spikes and rounded postorbital bosses, which aligned it closely with Styracosaurus while emphasizing ontogenetic variations in horn development.^[3] Early analyses of the associated bonebeds, detailed in Sampson's 1995 monograph, revealed taphonomic evidence of gregarious herd behavior, with remains from at least 15 individuals concentrated in low-diversity assemblages suggesting mass mortality events, possibly from drought or flooding.^[3] Age profiling indicated a predominance of subadults, comprising approximately 75% of the sample based on skull and long bone metrics, implying segregated or mixed-age herds similar to modern ungulates.^[5]

Referred and potential specimens

In addition to the holotype, numerous specimens from the Canyon Bonebed (TM-046) and Dino Ridge Quarry (TM-023) in the lower Two Medicine Formation of Montana have been referred to Einiosaurus procurvicornis. These include at least 15 individuals represented by multiple partial skulls, postcranial elements such as vertebrae, ribs, limb bones, and pelvic girdle fragments, all recovered from these monospecific bonebeds approximately 45–47 meters below the Two Medicine-Bearpaw contact.^[4] A notable referred specimen is the articulated subadult skull MOR 456 8-8-87-1 from TM-046, which preserves a forward-curving nasal horncore and developing supraorbital bosses, confirming its assignment through shared diagnostic features with the holotype.^[4] Post-1995 analyses, including ontogenetic studies, have solidified these referrals by demonstrating consistent morphological progression in ornamentation across the bonebed assemblages.^[4] Referral criteria emphasize autapomorphic traits unique to Einiosaurus, such as the procurvicorn (forward-curving) nasal horn and a specific epiparietal arrangement featuring a low, triangular P3 epiparietal along the posterior frill margin. These features distinguish referred material from contemporaneous centrosaurines like Centrosaurus or Styracosaurus, with postcranial elements further corroborated by proportional similarities in limb robusticity and frill texture transitions observed in the bonebeds.^[4] Potential specimens include the subadult skull MOR 591 from locality TM-077, higher in the formation (within 50 meters of the Two Medicine-Bearpaw contact), which exhibits ambiguous traits like a low nasal horncore and elongate supraorbital bosses that could align with late-stage Einiosaurus ontogeny or early Achelousaurus horneri.^[4] Initially referred to Achelousaurus in early assessments, its assignment remains debated due to stratigraphic separation from the main bonebeds and overlapping morphological variation, with recent reviews favoring cautious attribution pending further comparison.^[4] No major new discoveries or reassignments have emerged in 2000s–2020s research beyond refinements in these bonebed materials.^[4]

Description

Size and overall morphology

Einiosaurus was a medium-sized centrosaurine ceratopsid, with adult specimens estimated to measure 4.5 to 6 meters in length and weigh between 1 and 2 metric tons, estimates derived from skeletal comparisons with closely related genera like Centrosaurus and Styracosaurus.^[1] The overall body plan followed the typical centrosaurine form, characterized by a robust, low-slung build reminiscent of a modern bison, supported by strong fore- and hindlimbs that enabled quadrupedal locomotion in adults while facilitating weight-bearing for low-level herbivory.^[3]^[5] A distinguishing feature of its morphology was the short, deep tail, which contributed to a compact posterior profile without the elongated vertebral column seen in some other ceratopsids, and the absence of an extremely elongated nasal horn, instead featuring a moderately sized, forward-curving structure.^[3] Juveniles exhibited bipedal posture and were notably smaller, with body sizes scaling proportionally to ontogenetic stage, and less pronounced cranial ornamentation that developed progressively with maturity.^[8] This robust construction, including powerful limbs, supported efficient foraging on tough vegetation in its Late Cretaceous habitat.^[5]

Skull and ornamentation

The skull of Einiosaurus procurvicornis is characteristic of centrosaurine ceratopsids, featuring a robust construction adapted to its herbivorous lifestyle, with adult specimens reaching lengths of up to approximately 1.5 meters from the tip of the nasal horn to the rear of the frill.^[3] The facial region is relatively narrow and pointed anteriorly, with the premaxillae forming a beak-like structure for cropping vegetation. A key autapomorphy is the low, forward-projecting nasal horncore, known as a procurvicorn, which in adults curves strongly downward, distinguishing it from the more upright nasal horns of related taxa like Styracosaurus.^[4]^[3] The frill is rectangular in outline, extending posteriorly from the occiput, and is adorned with three pairs of well-developed epiparietals along the posterior and lateral margins, including notably elongate, straight, and posteriorly oriented P3 processes that project backward.^[3] The squamosals form the lateral portions of the frill, reaching lengths of around 29–31 cm in adults and subadults, with fenestrae positioned typically for ceratopsids: paired parietal fenestrae centrally and smaller squamosal fenestrae laterally. Supraorbital horns are small and subdued, appearing as rounded, spheroid bosses over the postorbitals in adults, measuring approximately 12 cm in length, while the jugals contribute to the cheek region without prominent ornamentation.^[4] Ornamentation undergoes significant ontogenetic progression, with juveniles exhibiting smaller, more upright nasal and supraorbital horns that lack the pronounced curvature seen in adults. In subadults, the nasal horn is erect and recurved, reaching about 13 cm, while supraorbital ornamentation transitions from low, transversely flattened platforms to more prominent, rounded masses.^[4] By adulthood, the nasal horn becomes highly procurved, and the squamosals elongate further relative to the face, with the frill achieving near-adult proportions early in development before facial features fully mature.^[4]^[3] Sensory adaptations include large orbits that suggest good binocular vision, bordered by robust postorbital and lacrimal bones, and expansive nasal passages forming a deep, vaulted cavity that likely supported olfaction through proximity to the olfactory epithelium.^[9] These features, common to ceratopsids, indicate an acute sense of smell for detecting food or environmental cues, with the nasal chamber's ventral channels potentially enhancing airflow efficiency.^[9]

Postcranial skeleton

The postcranial skeleton of Einiosaurus procurvicornis follows the conservative morphology typical of centrosaurine ceratopsids, known primarily from partial skeletons preserved in monospecific bonebeds containing elements from multiple individuals.^[4] These assemblages include vertebrae, ribs, limb bones, and pelvic elements, though the caudal series is often incomplete due to taphonomic biases in the deposits.^[4] The axial skeleton comprises 10 cervical vertebrae, with the first three fused into a robust syncervical that provides structural support for the heavy skull and frill; the cervical centra are short and amphicoelous, featuring tall neural spines that enhance rigidity in the neck region.^[10] Dorsal vertebrae number approximately 12, exhibiting robust construction with prominent transverse processes for rib articulation, contributing to a compact torso suited for quadrupedal locomotion. The sacrum consists of five fused vertebrae, and while over 40 caudal vertebrae are inferred based on ceratopsid patterns, preserved specimens show gaps in the mid- and distal tail regions.^[4] Limb elements indicate a weight-bearing quadruped with powerful fore- and hindlimbs of subequal length. The humerus exceeds the radius in length, with strong, cylindrical shafts adapted for load-bearing; metacarpals are robust and straight, supporting a manus with five digits where the central three bear weight.^[11] In the hindlimb, the femur measures up to 644 mm in adults, slightly longer than the tibia (up to 495 mm), terminating in a pes with three functional digits equipped with hoof-like phalanges for terrestrial stability.^[11] The pelvic girdle features broad, flaring ilia that articulate with the sacrum to provide a wide base for stability during movement, a trait shared among centrosaurines for accommodating the massive cranial mass.^[3]

Classification and evolution

Phylogenetic position

Einiosaurus is recognized as a derived member of the Ceratopsidae family, specifically within the subfamily Centrosaurinae, based on cladistic analyses of cranial and postcranial features.^[3] This placement is supported by shared synapomorphies of Centrosaurinae, including reduced postorbital horns relative to those of Chasmosaurinae and elaborate epiparietal ornamentation on the parietal frill, such as spike-like processes at the P3 locus.^[3] These traits distinguish centrosaurines from other ceratopsids and position Einiosaurus among the more specialized North American taxa from the Late Cretaceous.^[12] In the original phylogenetic analysis by Sampson (1995), Einiosaurus formed part of a "pachyrhinosaur-grade" clade with Achelousaurus and Pachyrhinosaurus, which was positioned as sister to a Centrosaurus-Styracosaurus clade, reflecting an early view of its affinities toward taxa with modified nasal and frill structures.^[3] Subsequent studies in the 2010s refined this positioning within a North American centrosaurine radiation; for instance, Ryan et al. (2012) recovered Einiosaurus in a derived group with Achelousaurus and Pachyrhinosaurus, while emphasizing its distinction from basal forms like Wendiceratops through the absence of certain epiparietal processes (e.g., P1).^[12] Many analyses, including those incorporating additional taxa, resolve Einiosaurus as part of the Pachyrhinosaurini tribe, sister to Achelousaurus and Pachyrhinosaurus, with Styracosaurus as a more basal eucentrosaurine, supported by similarities in frill elaboration and nasal horn curvature.^[13] Recent phylogenetic work through 2020 confirms the stability of this placement, with no significant revisions in the literature from 2023 to 2025, maintaining Einiosaurus as a eucentrosaurine in the broader Centrosaurinae framework.^[13] This consistent positioning underscores its role in the evolutionary diversification of centrosaurines during the Campanian stage.^[14]

Anagenesis hypothesis

In the early 1990s, paleontologist Jack Horner proposed an anagenetic evolutionary sequence within centrosaurine ceratopsids, suggesting that Einiosaurus procurvicornis gradually transformed into Achelousaurus horneri through reduction and fusion of cranial horns into bosses, and subsequently into the smaller, more juvenile-like Brachyceratops montanus, all within a single lineage spanning approximately 1 million years during the late Campanian stage. This hypothesis posited no branching speciation (cladogenesis) but rather continuous, directional morphological change driven by environmental pressures, such as marine transgressions influencing habitat and selection in the Two Medicine Formation of Montana.^[15] Key evidence supporting this model includes the stratigraphic superposition of fossil bonebeds in the Two Medicine Formation, where Einiosaurus specimens occur in lower horizons (Hagans Crossing Member), Achelousaurus in intermediate levels (roughly 25 meters higher), and Brachyceratops in uppermost deposits (Flag Butte Member), indicating temporal succession without overlap.^[7] Additionally, ontogenetic series from Einiosaurus bonebeds reveal progressive changes in horn curvature, size, and fusion that parallel the differences observed between the taxa, such as the shift from forward-curving supraorbital horns in Einiosaurus to flattened bosses in Achelousaurus, mimicking paedomorphic retention in Brachyceratops. While some early analyses, including those by Scott Sampson, acknowledged the stratigraphic and ontogenetic patterns as suggestive of close evolutionary ties, subsequent cladistic phylogenetic studies have recovered Einiosaurus, Achelousaurus, and Brachyceratops as closely related but distinct species within a broader eucentrosauran clade. However, recent stratigraphic and morphological studies, including the 2020 description of Stellasaurus ancellae from the Two Medicine Formation as a transitional form between Styracosaurus and Einiosaurus, have reinforced support for an anagenetic sequence (Styracosaurus → Stellasaurus → Einiosaurus → Achelousaurus), emphasizing continuous evolution over branching. Critics also note that Brachyceratops may represent juvenile forms of other centrosaurines, complicating the linear progression, though insufficient overlapping material prevents full synonymy.^[13] No definitive refutations or confirmations of the hypothesis have emerged from studies between 2023 and 2025, with ongoing assessments emphasizing the need for additional specimens from intermediate stratigraphic levels to test taxon ranges and morphological transitions.^[7] If validated, this model would imply rapid, non-branching evolution in ceratopsian cranial ornamentation, highlighting how short-term geological events could drive significant adaptive shifts in centrosaurine display structures over limited timescales.^[15]

Relationships to other ceratopsids

Einiosaurus is classified within the centrosaurine subfamily of ceratopsids, forming part of the Pachyrhinosaurini tribe alongside Achelousaurus and Pachyrhinosaurus, with Styracosaurus positioned as a basal member of the eucentrosaurine lineage featuring a prominent spiked frill.^[16] In Bayesian phylogenetic analyses, Einiosaurus shares a close clade with these taxa, exhibiting transitional nasal horn curvature and reduced parietal processes compared to the more elongate, erect nasal horn of Styracosaurus.^[16] Diabloceratops, an earlier and more basal centrosaurine, represents an ancestral form with plesiomorphic cranial features, such as shorter nasal horns and less elaborate frill ornamentation, highlighting Einiosaurus's position in a derived North American radiation.^[17] In contrast to chasmosaurines like the later Torosaurus, Einiosaurus displays the typical centrosaurine morphology of a long nasal horn, short supraorbital horns, and a solid, posteriorly directed frill with low osteoderms, whereas chasmosaurines exhibit elongated brow horns, reduced nasal horns, and expansive frills with large fenestrae for weight reduction.^[18] These differences underscore the divergent evolutionary paths within Ceratopsidae, with centrosaurines emphasizing nasal and frill-based displays over the prominent orbital horns of chasmosaurines.^[19] As part of the Campanian-stage spike in centrosaurine diversity across northern Laramidia, Einiosaurus exemplifies the rapid morphological experimentation in frill and horn configurations seen in taxa like Centrosaurus and Styracosaurus, contributing to high endemicity in Montana's fluvial settings.^[16] This contrasts with Asian forms such as Sinoceratops, a basal centrosaurine with a robust skull and long nasal horn but lacking the extreme frill spikes of North American derivatives, indicating limited dispersal and distinct biogeographic evolution despite shared primitive traits. No significant new relational discoveries for Einiosaurus have emerged since 2020.^[16]

Paleobiology

Functions of cranial ornamentation

The cranial ornamentation of Einiosaurus procurvicornis, consisting of a low, forward-curving nasal horn, short triangular supraorbital horns, and a large frill with prominent backward-projecting spikes, has been hypothesized to primarily serve functions related to species recognition and intraspecific display rather than aggressive combat or defense against predators. Sampson (1995) proposed that the distinctive morphology of the nasal horn, which curves anteriorly and downward in adults, was ill-suited for goring opponents or deterring predators, instead facilitating visual signaling for mate attraction and conspecific identification during reproductive behaviors.^[3] This interpretation aligns with the overall centrosaurine pattern, where elaborate frill ornamentation emphasizes display over weaponry, contrasting with the longer, straighter brow horns of chasmosaurines that may have played a greater role in anti-predator defense.^[3] Biomechanical analyses of centrosaurine skulls, including comparative studies on related taxa like Centrosaurus, indicate that the frill's robust construction and internal buttressing could withstand forces from intraspecific head-to-head or side-to-side butting. However, the low profile of the nasal and supraorbital horns in Einiosaurus suggests limited involvement in direct physical confrontations, supporting a primary role in non-contact displays such as head-bobbing or posturing to establish dominance hierarchies.^[20] Ontogenetic evidence further bolsters this, as ornamentation develops late in maturity and shows high variability, consistent with sexual selection pressures for visual cues in mate competition and species recognition.^[20] Alternative hypotheses, such as thermoregulation or protection against injury, have been considered but receive less support for Einiosaurus. Early proposals suggested the frill acted as a heat-exchange surface due to its vascularization and surface area, potentially stabilizing body temperature in variable climates (Farlow, 1974). However, isotopic analyses of ceratopsian bone indicate modest thermoregulatory capacity at best, with display functions better explaining the structure's complexity and energy investment.^[21] Similarly, while the frill might offer passive shielding for the neck, evidence from Einiosaurus bonebeds reveals low rates of healed trauma or pathology on horns and frills, implying infrequent aggressive use and favoring non-violent signaling over combat or injury protection. The absence of puncture wounds or fractures in the monospecific assemblages further underscores that predatory defense was unlikely a key role, given the orientation of ornaments toward conspecific interactions.^[3]

Growth patterns and ontogeny

The ontogenetic series of Einiosaurus procurvicornis is well-documented from the monospecific bonebed in the Two Medicine Formation, which preserves individuals across multiple growth stages from juveniles to near-adults, allowing reconstruction of developmental trajectories.^[6] This assemblage reveals rapid somatic growth during the early life stages, with bone histology indicating fibrolamellar tissue deposition that supports high metabolic rates and fast skeletal expansion, consistent with patterns observed in other ceratopsids like Centrosaurus.^[8] Growth spurts are most pronounced in the first 3–5 years, during which individuals likely reached approximately 4–5 meters in length and approached sexual maturity, after which the transition to parallel-fibered bone signals a deceleration in growth rate.^[8]^[22] Cranial ornamentation undergoes significant transformations during ontogeny, with the parietosquamosal frill expanding dramatically post-hatching to achieve near-adult proportions early in development, often preceding full maturation of other features.^[6] The nasal horncore begins as an erect or slightly recurved structure in juveniles and subadults, laterally compressed and sagittally divided, before fusing and curving strongly forward in adults to form the characteristic procurved morphology.^[6] Supraorbital horncores similarly evolve from pointed, plesiomorphic forms in early stages to inflated, flattened platforms in subadults and finally to rounded, spheroid bosses in adults, highlighting a sequence where frill and supraorbital development outpaces nasal horn modification.^[6] Histological analysis of long bones from the bonebed confirms these patterns, with lines of arrested growth (LAGs) present in tibiae and other elements indicating annual cyclicity and a lifespan potentially extending 20–30 years in ceratopsids, though the preserved Einiosaurus specimens rarely exceed 6 years of age and represent subadult stages.^[8] Evidence for sexual dimorphism in horn robustness remains debated, with some variation in ornamentation possibly attributable to sex rather than age alone, but insufficient sample sizes limit conclusive interpretations.

Behavioral inferences

Fossil evidence from the Two Medicine Formation indicates that Einiosaurus exhibited gregarious behavior, as demonstrated by two monospecific bonebeds preserving remains of at least 15 individuals of varying ages.^[4] These assemblages suggest that Einiosaurus lived in herds typically comprising 10-20 individuals, potentially including juveniles and adults to provide protection against predators through collective vigilance and defense.^[4] Such social structure is consistent with patterns observed in other centrosaurine ceratopsids, where bonebeds reflect group mortality events, possibly from environmental catastrophes or predation.^[23] As a low-level browser, Einiosaurus likely foraged on vegetation at heights of 1-2 meters, utilizing its robust beak and shearing dentition to process tough plant matter. Skull mechanics, including a deep jaw and powerful adductor musculature, enabled efficient slicing and crushing of fibrous materials such as ferns and cycads, which were prevalent in its floodplain environment. This feeding strategy minimized competition with taller herbivores like hadrosaurids, allowing Einiosaurus to exploit understory plants effectively. Einiosaurus faced significant predation pressure from large tyrannosaurids, particularly Daspletosaurus horneri, the apex predator of the Two Medicine Formation. Herd dynamics likely played a key role in defense, with adults forming protective circles around juveniles during attacks, leveraging their forward-curving nasal horns and frill spikes to deter assailants. Evidence of healed injuries on ceratopsid fossils supports the idea of intra- and inter-specific combat, but gregarious living would have enhanced survival against solitary or small-group predators like tyrannosaurids. Cranial ornamentation in Einiosaurus, including the distinctive low-slung frill and forward-projecting nasal horn, supports inferences of mating displays analogous to those in modern ungulates. These structures may have been used in visual or auditory rituals to attract mates or establish dominance within herds, with vibrant coloration or posturing emphasizing species-specific traits. Such behaviors align with sexual selection hypotheses for ceratopsid frills, promoting reproductive success in social groups.

Paleoecology

Geological context

Einiosaurus fossils are primarily recovered from the upper unit of the Two Medicine Formation in northwestern Montana, a geologic unit deposited during the late Campanian stage of the Late Cretaceous, approximately 75.2–75.1 million years ago.^[16] This formation consists of sediments from an alluvial floodplain environment, featuring meandering river channels, overbank fines, and associated paleosols. Radiometric dating of volcanic ash layers (bentonites) using CA-ID-TIMS U-Pb methods has provided precise constraints on the timing of deposition in this upper section.^[7] The characteristic Einiosaurus bonebeds occur within fine-grained overbank siltstones and mudstones, indicative of rapid burial by crevasse splay or flood deposits that preserved large assemblages of individuals, including juveniles and adults. These monospecific accumulations suggest localized mass mortality events followed by quick sedimentation in low-energy fluvial settings. Taphonomic analysis of the bonebeds reveals disarticulated but largely unabraded skeletal elements, consistent with minimal transport prior to burial.^[24] The depositional environment reflects a semi-arid paleoclimate with marked seasonality in precipitation, as evidenced by caliche-bearing paleosols and growth interruptions in preserved conifer wood. This climate supported a lowland vegetation community dominated by ferns, interspersed with conifers, horsetails, and early angiosperms adapted to periodic water availability along riverine corridors. No additional stratigraphic localities yielding Einiosaurus have been documented since 2022.^[25]^[26]

Contemporaneous biota

Einiosaurus inhabited the Two Medicine Formation of northwestern Montana during the late Campanian stage of the Late Cretaceous, approximately 75 million years ago, where it coexisted with a diverse community of herbivores in a floodplain ecosystem. Other ceratopsians, such as the centrosaurine Achelousaurus horneri, shared this environment, representing a succession of horned dinosaurs within the formation. Hadrosaurs were abundant, including the lambeosaurine Hypacrosaurus stebingeri and the hadrosaurine Prosaurolophus maximus, which likely browsed at varying heights. Ankylosaurs like Scolosaurus cutleri and Edmontonia rugosidens also occurred, contributing to a herbivore assemblage dominated by large ornithischians adapted to the lush, riverine habitats.^[27] Predators in this ecosystem included tyrannosaurids, notably Daspletosaurus, which served as apex carnivores capable of preying on or scavenging large herbivores like Einiosaurus. Smaller theropods, such as troodontids (e.g., Troodon), occupied niche roles as agile hunters or scavengers. Evidence of theropod bite marks on Einiosaurus bones from bonebeds suggests interactions involving predation, scavenging, or post-mortem feeding, often linked to drought-stressed mortality events that concentrated carcasses.^[28] The flora of the Two Medicine Formation, reconstructed from megafossils and palynological data, was characteristic of a coastal plain with ferns (e.g., Osmundaceae), horsetails, conifers, ginkgophytes, and early angiosperms comprising up to 30 species, including members of Aceraceae. This diverse vegetation supported a productive ecosystem, with angiosperms becoming increasingly prominent. Palynomorph assemblages confirm the presence of monocolpate and tricolpate pollen, indicating a mix of gymnosperms and emerging flowering plants.^[29]^[30]^[31] As a mid-level browser, Einiosaurus likely targeted tougher, woody vegetation such as ferns and low angiosperms, partitioning resources with taller-feeding hadrosaurs and low-browsing ankylosaurs in the broader Laramidian floodplain community. This niche integration minimized direct competition while enabling coexistence in a dynamic, seasonally variable environment.^[27]

References

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Einiosaurus | Natural History Museum
Einiosaurus, a ceratopsian, was 6.0m long, herbivorous, lived 74 million years ago, and had a large downward-curving nasal horn.Missing: primary sources
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Thus, in ceratopsids, the ventral channels of the nasal cavity which run parallel to the highly vaulted oral cavity have been also elaborated.<|control11|><|separator|>
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A new, transitional centrosaurine ceratopsid from the Upper ...
Apr 29, 2020 · ... Einiosaurus procurvicornis and Achelousaurus horneri. Here, we reassess the previous referral of specimens to Rubeousaurus ovatus and ...<|control11|><|separator|>
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Nov 8, 2012 · At approximately 78 Ma, it is the oldest known Canadian ceratopsid, approximately 0.5 Ma older than Albertaceratops from the lower Oldman ...
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Variation in the shape and mechanical performance of the lower ...
Sep 11, 2015 · 2014), a few contributions have highlighted possible lower jaw shape differences between centrosaurines and chasmosaurines and within each clade ...
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Centrosaurine ceratopsid bearing the following autapomorphies: hypertrophied narial region (approx. 75% preorbital skull length); caudal portion of fused ...
[20]
[PDF] VERTEBRATE PALEONTOLOGY OF MONTANA - MBMG
(1) INTRODUCTION. Montana is renowned for its rich paleontological treasures, particularly those of vertebrate animals such as fishes, dinosaurs, ...
[21]
[PDF] Craniofacial Ontogeny in Centrosaurine Dinosaurs (Ornithischia
Sampson (1995a) recently erected two additional genera, Einiosaurus and. Achelousaurus, from the late Campanian of Montana. Vertebrate paleontologists must ...
[22]
The thermoregulatory functions of the Triceratops frill and horns
The results suggest very high and uniform heat flow through the parietal frill, maintaining mean frill temperatures between 0–4°C below the body core.Missing: Farlow | Show results with:Farlow
[23]
Bone histology and growth curve of the earliest ceratopsian Yinlong ...
Dec 19, 2024 · The humeri of Triceratops display a mix of longitudinal and circumferential vascular canals and bone remodeling happens in a very early stage, ...
[24]
The First Triceratops Bonebed and its Implications for Gregarious ...
Aug 10, 2025 · Two new ceratopsid dinosaurs, Einiosaurus procurvicornis and ... A peculiar bonebed reinforces gregarious behaviour for the Triassic dicynodont ...
[25]
Taphonomy of Three Dinosaur Bone Beds in the Upper Cretaceous ...
Taphonomic studies have explored the fossilization process as it pertains to Two Medicine Formation fossils and revealed aspects of the ancient Two Medicine ...Missing: Einiosaurus | Show results with:Einiosaurus
[26]
Paleoecological utility of insect trace fossils in dinosaur nesting sites ...
Their trace fossils are consistent with previous inferences of semi-arid conditions and seasonality for the Two Medicine Formation. Moreover, apocritan ...
[27]
The early Campanian flora of the Two Medicine Formation ...
The fossil plants occur in the lower part of the Two Medicine Formation ... The flora contains two species of hepatics, one sphenophyte, six ferns, two ...Missing: paleoflora | Show results with:paleoflora
[28]
Competition structured a Late Cretaceous megaherbivorous ...
Oct 28, 2019 · Note that herbivore:carnivore size ratios are closer to 1 in the dinosaur ecosystems; that is, predator and prey are more nearly equal in size.
[29]
Paleontology - Egg Mountain - SERC (Carleton)
Apr 10, 2018 · ... overview of dinosaurs and paleontology along the Rocky Mountain Front. You can't miss the brightly colored Einiosaurus and T. rex in the ...Missing: description | Show results with:description
[30]
"The early Campanian flora of the Two Medicine Formation ...
The early Campanian flora of the Two Medicine Formation northcentral Montana ; Department or School/College. Department of Botany ; Committee Chair. Charles N.Missing: paleoflora | Show results with:paleoflora
[31]
ECOLOGICAL ASPECTS OF THE CRETACEOUS ... - Annual Reviews
The Two Medicine flora has 30 angiosperm, 2 conifer, and. 6 fern species (Crabtree 1987). Although the angiosperms are the most diverse element of the flora ...
[32]
Late Cretaceous (Campanian) leaf and palynoflora from southern ...
Angiosperm pollen comprise up to 15% of the assemblage, represented by monocolpate, tricolpate and periporate pollen and the extinct Normapolles group.